SYSTEMATICS OF ZAPOTECA — Héctor M. Herndndex* (LEGUMINOSAE)! “ABSTRACT Zaporcs (Leguionse: Mivonstea) is a tebivey wellckcamscried gems beloging to the wit Ingear tected Gm te wuthvesons United Stam sod norarn Mesizo to aorkeen Arora and the West Ine ‘The hagestconcrtaton of tana occurs ithe dy forest of southern Menico and, vos ser extent a Cntr merce andthe nother Ande. The protominance af alopt dations andthe apparent ow frequency ol tye sugeeat htspecion nis gems ax pana arisn foe get dierenlininn ef ger apicay eed pps Ti ect cena pads othe Peacocece apps tu have had promioeal e'om fhe cre ‘Etrbutonal pacers o€Zaporne ye. Zapotce specs ive ver repraducve loge A ply have tural ethos and ae gliatd by anpecalaod cating Wethe,Sa compat cpyearew be wilpred i is genin, Ztetagina Boag the only known seitinsemwtble species Ardromonocey is nother wedrprend Sapomon o the beting system of Zapoter spies. THs probally maxis the bse ol repraacive energy ‘oor redoing plan suetinn The bri chrtewcme rather f Zapengen r= 13) and wef the promrlaed ‘aractor af te ply (uss aclymime's, deca TS grained ply indice a reanship hewn Zapotocz Sad th seuinog genera oft lageae. wth the shcepten of Calizndra see sacs, wach ir eyaosealy and Sorplogrally soled mth tribes Tsecognie Lv speasio-Lopoeca tas of hie alinae, 2. ending. ad Peta) are ney decid. Eleven singed sce recogni, of wach Z_portancenss np. pelicarpa ‘oly decd. A oiprere anion i propoed, wets zane goupe! mio fr mepholgealy inet ‘SSbgsnre’ Nesta, Lopursen,lewentm and amesonica ‘The sigeners sre based primary ferences in Yogattvecheracars numberof oof plana, mba fost pepo pn, lett te, leat rota, vation aterm, an spl ype RESUMEN Zapotec (Laguicosen: Mime) ox un yénero rlativarnente bien desitade que pectenee al ib Ingese ‘Se dtribuye desde e sarcese do los Eatadne Unde yl rorte de Mako sia el wer de Argentine y las Ane. {i mae elt concentrasin de taxa a0 da en le bogus tropealssoscr del sur de MésiceY, en ence tedida, en CCeotco Aiea y ela pockn worte de lee Andes, La predominancin de dtrbaciones alopitriae, en conbinacion "ds study is pact of my docworal desis at Saint Louie Universiy, Sain: Louis Miswour. lank De- Peer HL oven, Director ofthe Mixeari Botanical Garden, for duectng my riearch and for support and encouragement in all wages ofthe projert. The Missour:Hotanical Garden provide oce space and herkarie, Kerry. and greeshowse Tnclites The asutance of te ial and technics ofthe Minours Uotaniel Gorden was esters wal grateflly secnvedged Tao dank Drs. Rupert Barnsby, Gerrit Davidse, Joba Dwyer, Alnye Geney, Mario Sousa, and ax snenymoue reviewer for reading the manticrpt and providing hpful comments and olor alec: Ph. Gate! R- Keating, S.Keptar, PLowry. ard B. Stom for nelly roading port ofthe searmaonps J. Deer Cor providing the Latin Alserptions of the new taxa C. Belelapacher (orienting the flower veto: J. Myers fr his excallentbotaneal Hurtin: . Vileggs for Figure 48; ML Veit hl ML Reyes for sssitance in We ube uf the SEM at Wanington University and Tete de Geslgie, UNAM, respectively: and D. Thiel for typing par of the ewiceata. Tam ‘epecalygraofl tothe foloning porone wh provided sted sales and other botanieal material: M, Corre, A Delgado, (. Dodson, f. Foro, (. Hghes John, J. Levis, Es Lat, Miler, D, Nel, J Oakes, P. Rebiese D. Smit, W. Stevens, Re'Tortm, J. Vash T. Wen, and Zancai, Wao thank the Surates othe following Iria for providing spocineras Ay BM, BR, CAS, COR, ENG, FG, GH, CORT, K, LE, MA, MEXU, MICK), IY, P, 5, TEX, UG. US, WIS, and XAL Told etusce wou not have been pocle without tho Snanciel acstance of the Micouri Botanical Candon and ‘World Wilts Fond 1, (Tropical Uotany Fellowship) 1 thnk the flog rire hn provided ata the fell M. Arredanio, 8. Artedondo”H. Chutth. M. Qucrea, M: Grayun, P- Gay, L. Me:vdoden, D- Janes, P. Moroso, A: Rater, A. Shin Ten, W. D. Stevca, P. Tenorio, ea R Tore, ‘Tho ve your cay i Sain: Loui, Miscou wos mate possible for my family and mo by a cebelartip provided by the Intute de Halo, Univerided Nacional Auténomea do México, theough ti Deparamente ds Bees. U express sincore gttode lo De ox Sahn K., then diector of the Inst do Blogs, and Dr- Mav Sos Soret ‘trator ofthe Herter Nacional de México|MEXU) for thee confidence and continua support. Thanks ae extended to the Missou Botanical Carden for paral anal support during our last yeat Ie Sunt Louis. Fly, ay dep thanks toy wife Alina and to Pablo fer thes love and emodanal support dating this log experience Miscou! Botanical Carden, P.O, Box 29U, Saint Lou, Miscour 03160, USA. Present edirese Hodbusio Nacional de México, Instituto de Biologia, Universidad Nesoral AUtéroma de Mexico, Apartado Postal 70-367, Ginlad Universitaria, C4510, Maize, DP, Mesico, Ann. Missourt Bor. Ganp. 76: 781-862, 1989. NextPREV 782 ‘Annals of the Missour Botanical Garden on I aparente baja frecuencia de bibrios, sapere que ls mpeciacin en exe pero ha sdo ol rsuiade de un Jprocoro de dierenciaciin gentiea do poblaciono> alas googyitcarente. Los opzodios climdtieo: reciens de Prestoceno pasblementetwveron un pépeldatorminare stro ke gotrnes de dababusge actualoe do las enpesine de Zaputern. Las esjroes ik Zapoteen son bastante wformes en lon regos genereles de = lmogia reproduc, ‘Toles denen ponbletente fares ton anton nectar y 20m paitzadae por palrllan no especialy, Low sles de asxccompatblidad grnition parecrn sar [én eepeewentaion en opener, 2 letragoun send la Goce especie fn donde le tuto ncomperibded ba se dewentrada La aniremonoeea colo compere cords dels shea ‘productos de ls eopsciee da Zopotace; este fonémeno co interpreta com i ieo pra msainitar € wo de ‘erga reprdutiv, sin rodackt le atracctn aloe poliasador=s, El nero comosimica bisa de Zapotecs ( 1} en mucaciin on alguns cactees generalzaon de fas poliades (e.poinies neaimnda, dears, con 16 rates), reff reacones ene Zopasec yo reno los genes de at gan, com la encepin de Calne Sn sro, ginore que fuattemente agico desde al punto de vista ciolgico y motfolegico- En ete tajo se reeonoren 17 eepocks en Zapotece, do ls cuss Z.alinae, 2, cadina 2. tehwana se deserben a coro nace ae fara ln cenci, Tambien oe rovonocen LI eubsepecc, dele eolee Z. portoricense tbip. publonrpa se deecribo emo nueva, Ex base alos patrons de variack {ke pares de fl por pina, tao de lon ‘ipa dep Uktintes: Newose, Zaporeca, desleate y dmazonica earacteres vegetal ron extra elon fin, patrones de ranacin de los fie ae rope sia elasicacén sobgentrica, en done las espeties se agrupan en canto sulgererox urero de petes de pionas, nero Znpoteca (Leguminosae: Mimascieac) is a ge- tus of [7 species im the relatively specialized tbe Ingone, Ite species are mostly erect oF seandent shrubs, of small to modorate size, found in vege tation types ranging from arid, scrubby thorn for. ests to wet, evergreen tropical forests, although mutt species oceut in areas orginally covered by Uuopieal deciduous focests, Zapoteea ranges from southwestern Urited Mexico, ta the West Indies end southward throngh most of Mexico, Central America, and port of South Amer ia, to northern Argentina. Ht attains it greatest diversity in southern Mexico, although in. come trea: of Central Ameriea and the northem Andes diversity is also considerable. tates and north Zaporera was recently separate fom the large emus Calliandra Benth. (Hernander, 1986a). Ex- texsve analyses of the polyad characterises, seed ling morphology. chromosome mumbers. and sev tral roproductive features in Calliandra sensu Bentham revealed differontial variation potterns among its neotrapical species. These variation pat terns were associated with two distinct taxonomic groups: the first, inching the members of Ben- thauw's series Laetevirenies, and the secon, the remaining, neotropical species, these belonging 10 series Macrophyllae, Nividae, Pdicellatac, and Racemosar, The steonely divergent nature ofthee> two groups, along with the absence of intermediate forms horwoon them, suggested that Calliandra, 1s originally conceived by its author, is polyphyletic (ernindez, 19863). In an accompanying paper Romioo (1986) provided supporting evidence based fon differing patterns of nomprotein imino and se hur amino acids in seeds and leaves of Zapotcea and Calliandra. Consequonily. the new genus Za ppoteea was proposed to include the species of series Lactevirentes. Zapoteen and Calliandra are com pred in Table | TTeonsider all the species included by Bentham (1875) in Calliandra see. Lactevirentes to be Za ipotcea. When I eriuinally proposed Zapoteca (Hernindes, 19862), the possiblity of including Calliandra amazonica and C. aciulenta, treated by Bentham in ser. Macrophyllae, was discussed, Since these two species share a number of repro: oetive characters with Zaporeca (i.e, compact. sphorical, horamorphie eapitulas 16-grained dic ‘oid polyade with lens-shaped thiekeningss last cally dehiscent fruits with relatively thin valves), they are transferred in the present work to Za: ppotees, The stipular spines of Z aeuleeca, the compound pseudopanicles of Z. amazonica, and the small number and Inrger sie of the leaflets in hath species are interpreted as advanced charac teristics. These species should not be considered by any moans ab intermediate betweon Zaporoea and Calliendva: they are specialized species within Zapotoea, with divergent morphological charac Zapoteca is dele by its compact, spherical homemorphie esptulas aeslyramote, 16-grained. sliscoid pelyads with lens-shaped thickenings (ex eluding Z nereosal elastically dehiscent fruits with ‘mombranour or eoriaceous valves; and basic chro rmosome number of x = 13. This eombinstion of characters makes Zepoteca a well-cireurmscribed ceatitys however, with he exception of the distinc re circular thickenings in the polyads, each of these characters necurs mdepondently in ather gen xa of tne Ingeae. The general characteristics of the Zapoteca polyads its basic chromosome nur ber, und come ofits morphelogieal characters re fect close relationships of this genus with some NEXTPREV Vokime 76, Number 3 Hemandez 783 1988 Zepoteca Tarts J. Comparison of Zaporoca and Calandra fms Horninde, (860) NEXT Tapes Calliandra Trades ‘hin wembranous arly eoiacemua charac wo coronas Polyads L6prived,deoid Signs copsape, wlth nisrow ares of aha receptivity Pade membrana #0 oriaceo Sewdlings ‘orjedone orate, see, felines ephewel 3 Cheomosome sumbare spherical heads; monic ‘beonifonn, himertie ve eterna ‘grained, binmmmetn, vith Vek a engin “has” gain expanded: dco, or capitate scomeee, rarely spherical ie more rigid einen, liga sary ‘means cvtledonn sagt, petite, See, persbent fet genera of the Ingeae, perhaps with members ofthe Puheceltobium complex; however, to establish del. inite generic affine of Zapoteca ia dificult task due to our poor current understanding about go nerie delimitation within this complex. What seems lear is that Zapoicea has closer affinities with some of these genera than with Calliandra sensu stricto. Several characters in. Calliandra (e.g. polyad features, chromosome numbers, feral sor phology) are atypieal in the tribe Ingeas aed i dlieate that this genus is evclutionariy divergent from the other genera of the tribe This stuly provides an account of the mor phology, biogengraphy, reproductive biology, ey- Aogeneties, andl taxonomy Hf Zaporece. The taxoe nomic revision is based on over 4,000 herbs specimens from the major eallections of the world and on extensive field observations. Field studies were conducted in most of Mexico, Nicaragua, Costa Rice, and Panama during 1983-1985. In audition, individual representing eight species and five subspecies of Zepoteca were grown from sed ina greenhouse at the Missouri Botonical Garden uring 1982-1980; these plants provided infor- mation for the introductory chapters and served to complement the field observations and herbar: im studies when taxonomic decisions had to be made. Based on the critical exainination of cor related! morphological characters, in combination, wth ece-gengraphical factors, recognize 17 species fand 11 subspecies in Zapotec (Table 2). The species are classified into four sugencra, whieh represent diferent ovolutionary lines ae Paws Habit ‘The species of Zapoteca ace relatively uniform in hahit. Most are erect shrubs imately 3-4.ca tall with woody stems, In mest eases {he stems are rather thin (ea. 1~2 em in diaceter), although oceosionlly oividuals of Z. tetragona and Z. portoricensis develop stems up 4 20 em in diameter. ‘ fow species are senndent oF sub. tame Ouiline of the elasdieation of Zuposeen T Silgenus Neveore LZ. eros H. Subgomue Zapotecs Z. media Z inae Z.lamberviane 3 6 Portericensi fulpecies portariceass subspecies pubicerra Ibapecies flava ‘aol abpecies earazasane thlepecios eberbuuer’ retragone ‘ania ‘hepeciesformoce tubepeiessafadoronsis ce wubepecis gracilis 4. aecins ros i Ebepecies mola subspecies sehoral seuteliyire 13.2, MZ. MI, Satgenn Aeudnat 15. 2 oculoata 1, Subgeras mosanien 16. 2. microcepila 17. Z.anasoatcoPREV 784 seanlent. Zapotec portoricensis subsp. flavida and Z: amazonica grow in moist thickets supported by noighboring shrabs, Simi branches near the hase of the primary stems and has spreading branches that become prosteste or subprostrate. The scandont an prostrate habits aro found oxchsively in warm, moist habstts. Z. caracasans Leaves, ‘The leaves ofall species of Zapotec are stipulate and twieo pinnate with opposite leaf lets. The wide variation i leaf characteristics pro vides en exeellent basis for classifying members of the genus. Mature leaflets vary in length from about 0.5 em in Z. alinae and Z, media to a maximum cof 22 em in Z. ameazonica, In addition, there i 0 clear tendeney tonarde 2 reduction in sumber of leaflets concomitant with an increase in leaflet size. In all species of subg. Zapezcea the pinnae have from several to many pairs ol leaflets; by eautratt, in Z. amazoniea the leaves have ory one pair of pinnse with a single pair of large leaflets. Fur thermore, all members of subg. Zapotera have relatively’ membranous lallets, whereas in the re- raining species they are cleady chariaeraus to Extrafloral noetarioe aro unifortaly present in Z. Jilipes, Z.seutellifera, and Z. nervosa. They sre located between the incortion of the pine and. less frequently, between the distal pare of leet In Z. seuselljera and Z. ftipes an additional nee tary is located near the base of the petioles, The pectaries are more or less cup-shaped in Z. seu teller and Z. nervosa and exfiical io Z. fe lipes. According 10 Zimmrman's classifation (Blas, 1983), all of these fll into the “Hoch tariea"” catezory (elevated nectariee). Extrafloral nectates have lng been considered tu be the facal point of a mutually beneficial relationship between the plants that possess them and certain groups of ants [sce summary in Bentley, 1927). The ants exploit the nectar preted by these glands, which rich in sugars, amino acide, proteins, an other compounds (Flas, 1983) and provide the plant with varying degrote of antiherbivere protection. Howeyer, such a relationship has not yet bern aemonsteated in Zapotcea. Venation pauerns. Despite the aetthat leaves hhave numerous taxonomically useful characters, manoacaphere generally limit themselves to super ficial deseriptions of the most eonpicuous features of leat architecture, In this coetion, I describe the venation pattern of the leaflets of several specios (of Zapotrea, in order to obtain @ broad picture of the variation of this important component of the leaf architecture, ‘Annals of the Missouri Botanical Garden Mature leaets of ten species of Zapoteca, rp: ‘resenting all the subgenera (Table 3), were cleared NEXT following Dileher (1974). The terminology fellows Hickey (1973, 197), Variation in leaflet venation patterns is highly consistent wit the subgeneric lsssifeation in Ze pnteci and provides important inforaation for phy logenetic interpretation. All members of the serie essentially have brochidedromous venation. ‘The secondary veins ususlly depact from the primary vein at an acute to right angle, curving sbrptly to join superadjacent secondaries to forta alo In addition, loops are always enclosed hy arches formed by higher-order voine. Zapotera nervosa differs from the remaining examined species in having cucamptodromous venation in the hasel half cf each leaflet and brochidodromous venation in the upper half (Fig, 30). Furthermore, the tertiary domains in the leaflets ofthis species are elongate and parallel 10 the primacy vein, and the free: ending veinlets are move abundant in the distal ares and usually are once- of (wice-branelied The marginal venstion is oped in all Zepoteea species with the exception af subg. Amazoniea, where it is clearly frbriated, Similarly, all species Of the genus have incompletely clsed areulen ex- cept Z amazonica and Z. micracephalo, which have imperfectly developed areses. With the exception of Z. teluana, all species of subg. Zapoteca are characterized ly relatively fine, inconspicuous tertiary and higher-order veins that are difficult t0 discriminate even wmvder a dis speting microscope. By contrast, leaflets of Z. te uaa and al species of the other subgenera (Table 3) have thicker and conspicuous tertiary and qua ternary veine and froe-ending veinlets, The vera tion patterns of Z, sehuana (subg. Zapoteca) and Z acutewta subg, Acwleata) ace stikingly similar, ‘elleting taxonomic affinities between: the two sub lweoera, ‘hese two species share the basie brochi- Adodromous venation pattern of the genus; however, they differ in their marginal venation and ithe n of theie high-order veine and veinlts, Although the marginal venation of these two species is clearly looped, occasional free-ending margival veins are present. The vein cally oriented towards the adiecial-basal area of the lelletsm these two species, Zapoicea amazonica avd Z, microcephala have several characters in ther loves that depast fromm the rest of the genus. The tendency towards the ts concomitant with & rnurabor in these species is prob ably aseciatod with an increa2e in the degree of ‘organization oflealet venation. Hickey (1977) pr:PREV Volume 76, Number 3 Hernéindez 785, 1988 Zapoteca Tate 8. Characters of leaflet venation in Zepatren spec Telaive ser Ticks of Number Secondary Gy on Primary Marginal Tertiary Areoe ‘Taxon Venton Type Velo” Version Weim” evebprent Veils Sibaeous Neraone eucamprodronous 1 hoyed ‘ooderate incompletely 1-2 branched asaya roche eed iodromo di ‘aly Sutgenis Zapote. brochidckomens 1 doped hinike incompletely 1-2 Iranehed ce (exuding Z heed Ztehwava ——Vroehidedromaus ——1——looped wth Gne.emerate incompletely adel sig veine loeed raed Gernot Salyers dentate brockidodkommn 1 Ioped with frees mnderae snely ‘rd veins rai Genrod Subgenus Arnzon boelidodomeus 2 —_‘fibiated soodecate impertest deol “Material examined. Scbgemoe Nevcose:Z. neraoea, Bkmon 26 Suwonas Zapoteos Z formora, W939, H.210, Has H2I0 Traythonsks 5943 Zo posed a system of leaf clasifiation based on in creasing levels of regularity of venation (leaf ranks) tsbich provides a useful scale here. ‘The venation patterns of the two species of subg. Amazonia reflect an advanced consition beeanse their tertiary veins have relatively regular courses (third ranks see Fig. 48). In contrast, tho leaflet of the re raining species of Zapoteee fallin he second rank laecause their ertisry and higher-order veins are random in course apd are usually not wel differ. entiated from the lower orders. Of particule in lexest isthe presence of two coxprimary veins (Fig 48) in the leaflets of Z. amazonica and Z. micro cephala. This probably indicates leaflet fusion Hickey & Wolfe (1975) roported that loaflet fusion ‘sa common tendeney in the Leguminosae Leaf pubescence. With few exceptions, pu bescence type is nota reliable taxonomic charscter in Zapoteen, hecawse in most of the species i i clearly variable within and betwoen populations Zapoteca nervosa, Z. aculeata, Z, amazonica, and Z. microcephala appear wo he uniformly en lirely glabrous or nearly glabrous, whereas Z, mol lis is consistently pubescent throughout, In the remaining species of subg. Zepotera, pubescence ohuana, H.250% Z. alinar, H.952 Z. mols, Grayam 4201: Z poronwansss H-9A8, HI93 2, retagora, if 950, Subgenus devleato: Z. aeuleat, es rocephale, Hough 1711 (@), Calections in MO snless others tahested 1913999, Sobgeme sdmazonice: Z, amazonion, is variable reaching an extreme of ineonstancy in 2, formosa aud Z. portoricensis. A microscopic survey of cleared leaflets of 13 of the species re vealed three basic types of hairs (Figs. Ud). Thet taxonomic distribution is shoven in Table 4 ( rminology following Leclavathi & Ramayya, 1982) 2) Unicelllar conical hairs (Fig. 1) are the ‘most abundant and ate found atthe base of leaflets sand on the ruchille of Z. aouleata, along the leaflet margins of Z. microcephala, avd on all lea parts fof all the pubescent species of subg. Zapoteca. A very similar type of hairs, wicellular clavate (Fi 2).is occosionally found on the rachilla of Z. alinae able 4), (2) Mukicellular macroform conical hairs (Fi, 8) occur in all species of subg. Zapoteca except 2. formosa aud Z, ofinae, They are found, inter- mized with other types of sir, primarily on leaflet margins snd eachilas, (3) Filform capitate hnirs and ilform clavate Dies (Fig, 1) usually bave a unieriateeylindrical foot formed by a vanable numberof cell of varied Jengths, and 2 multiseriate (or rarely uniseriate?) hheadThese are usually restricted to leaflet margins aril rachills, and ace found in all species of subs, Znpoteen except Z. formosa.PREV 185 Annals of the Missouri Botanical Garden ‘Tams 4 Dasrbion of 9 lel trichomes in wleted secins of Zaporect. Ta = unieeule cosa Bb = nieollar clavate; If = mulch aeacroferm conic) 10) fife eassat or fiferm clorste, Relate abu dence tn indicated by: + ~ occasional. 4-4 ~ eegular, ++ = abundant. Allellecions in MO wes oernise fica H= aurhor's collections Type exo Calleeion i Te 1 Wi Sages Nereose Z. nervosa Bhmon 206 — - 8 - Subgenus Zapotec 2% olinae Hoa + + - + 2 formoss #210 ote — Z.formoss 1959 = ae - Z. porno Hour He — + Z. formosa H2%0 st Z fornen Seas 2562 ate — — Ztamboriana Brondlove $7412 34 + + Z. media H95T v4 = + te Z mols Grayum 4205 44 =o + Z, portoriconsis Hoe + oa " Z. pontoricensisssiap. fvida —H.709 ate - + . Z tehuene #950 44 + + Z teteogone 1986 + = ote + Subgeous Acwleate Ze aculeata Acosta 13399 + Sulgers omraconiea Z.amazonicn 2. microepats Worthowsts 5043, Haught 1710 (8) Stipes and possess a series of veins running parallel ta ‘the margine (Fig. 5, 6) in most spocies the stipules are conspicuous (ex, Z amazonica, 2, tetra zona), although in some they are very sal (8.5 Z. tehuana). Exceptional in this respest is Z Teaua, whic has spinoscent stipules (Fig. 46). and Z. nervosa, whose siipules are reduced to small (E mm) scales. Theenghout the genus the stipules ‘are adpcessed and probably have a protective func= tio during the early phases of leaf and inflores cence development, Zepoteca andiva is exeep: tional by having descersding or depressed stipules (Fig. 41). Moccovar all speries have stipals et the base of the distal pairs of pinnae and at the distal tip of every pinna, The stipules are persistent, leafy, INFLORESCENCES AND FLOWERS Inflorescences. Toe Bowers ace cansstently arranged in compact, homomorphicy «pherical capitate inflorescences. In the basic plan of intl essence organization, coltery or fasciculate pe- roles are axillary and subtended by a leaf, asin Z. alinae, Z media, avd Z. aculenta (Rig. 7A), There is, however, an evolutionary trond toward the progeessive reduction of the subtending leaves, which reaches its extreme in the two species of subg. Amazonica, where the leaves are always vestigiz|, and the inflorescence therefore appears tobe an elongate panicle (Fig. 7D, E). Transitional forms between the two extremes are found it several sposice of subs. Zapotece (Fig. 7B). For instance, in Z, mollis, Z. portoricensis, and od- bltonal species, the subsening leaves start deve- loping durmg or after anthesis. Sometimes, bov- ever, the subtending leaves never develop, as in viduals of Z. caracasana (Fig. 70) and Z. formose, in which the capivula are apparently aggregated in panilelke infloreseences. “The trend towards retardation and suppression of leaf de foliation aman the species of Zapatece has also of inflorescence orga ization. I interpret these types as steps in a con tinuum of inflorescence specialiration rather than ac diserote ond fundamentally diferent types. An indication of the derived eharacter of the “panicu late" inilorestenecs of Z. microcephala, Z. ania opment that results in differences in produced various pa NEXTVolume 76, Number 3 Hernéndez 787 1989 Zapotec PREV NEXT Ficunes 1-4, Leaf wichomes of Zopoteca—1. Uniilar conical Z,alinue Hl. Herndndes 952)—2. Unicel claete, Z alinae(H. Hernéndes 992).—3, Mulcellarmnecroferm conial,Z, rolls (Grayum 420.— 4. Fikform coptte, Z.telregona (H. Hernénde> 956). All olleetors at MEXU and MO. Seale bars = 100 ar onica, and other species in subg. Zapoteca is tho Tn summary, there has boon an evolutionary presence of persistent paired stipules at evory node trend of inflorescence modification in Zapotecd, long the flowering eystem. These inflorescence where retardation and suppression of the subtend- types, therefore, cannot be considered tritly pen- ing leaves has played a prominent role in changing culate, and for this reason the term “pseudopan- the overall aspect of the Mowering system. ‘The icle™ will be use. axillary capitula subtended by a developed leat Ficumts 5, 6. Stiules of Zapoteca.—5. Z. pertoricenis subse. portoricenss (H. Herndndes 948)—6. Z. teiragona (Bi. Biernéndes 956), Collectors at MEXU and MO. Seale bars = | em.PREV 788 Annals of the Missouri Botanical Garden Ficus 7. Patterns of ialbresconce organaaion in Zapotera.—A. Th eapitula at each noe are eubtened by aavelidoveloped lee Z alinae (Sous et al. 746, MO). -B. Tho cape azv subtended ty late developing kere Zporroricenss (Higgins 2578, MICH).—C-E. The subtending levee aro rest. (. 2. saracasona (lopli 15982, LL) DZ mcroceptate (Mats $800, US). E. Z.antasonica (Selaake 65), MO) NEXTPREV Volume 76, Number 3 1989 (leafy, axillary capitula) appear to represent the basic pattern. The paniclelike inflorescences of Z. imicrocephata and, especially, Z. amazonica (subs, Amazoniea) are highly modified and are cleatly Uerived from the basic type. Intermediate types exit ip many species of subg. Zapoteca, where & Single individual may have both leafy, axillary ca pita and lowering branches with vestigial or lato veloping subtonding Ioaves. Flowers, Aside from variation in size and in the relative proportions of their parts, the (lowers of Zapoteon species ace extremely constant. De tails of floral morphology and certain aspects of floral biology are in the generic and specific de- scriptions and in the section om seprodictive bie logy. The diversity of pollen types in subfamily Mi rmocoideae hat been studied extensively by severe) authors (Cuinet, 1969, 1981—a reviow: Sorsa, 1969; Niergods et a, 1983). One ofthe outstan- ing features inthis subfaonily isthe high frequeney of permanently compound grains, the2e eithor in totrads or more commonly in polyeds, Ae Guinot (1981) and others pointed out, the hosie polysd type in the Mimotoidese is the aealymmate type, in whieh the individual yoains are more or less free for one another. This form of polyad cohesion is in contrast to the ealyenm le typey where the ex temal layer of the exine i common to all cell Calyminate polyads are found in Calliandra AAs disewssed chewhore (Heminde2, 19863), several comprohonsive aureeya on tho pollen char actoristios of Calliandra (Guinet, 1965, 1969, 1981; Guinet & Barth, 1969: Sorsa, 1969: Nica soda otal, 1983) indicate the phyleticdistinetaess of Zapoteca, The kasi plan of polyad organization here is the same as that in the other genera of tribe Ingeae, but unlike that of Calliandra: sens stricto, All the species im the genus have aealym. rate, diccoid, radially symmetrical, 16-grained palyads (Figs. 8-13). The cight cantral grains, ‘which are distrbuced in two rows, are morpholog ically diferent from the peripheral grains (ie. the pallen grains exhibit heteromorphy). ‘The external Iyer of the exine hae an undiferentiated granular structure, and the grains have internal ‘apertures (see fig. 7 in Nievgeada et al., 1983). The sive of the polyads ranges from 76 to 141 at ‘Ono very striking diaencstic character, found no. where eke in Mimozoideae, is the occurrence of prominent lens-shaped structures in the eumtral fells on one side of the polyad. These ave found wiv Hernandez Zapoteca 789 in all species of Zapotcea excopt Z. nervosa. In conneetion with this, Cuinet (pers, comm, 1985) ‘observed that te rounded lens-shaped areas ve on both sides of the polyads (om eight gr daculesta snd Z amazonica; neveetheless, they less pronounced on ane side, Sich aroas some: times occur en some oral af the perigheral grains of Z alinae polyads (Fig. 9). Pod morphology is relatively constant oughout Zapotecs (Figs. 14, 15). The pods are usually straight, near, and flattened: they gen erally have nconstricted margins, visible seed chambers, and thickened sutures (terminology fol- owing Gunn, 1984), Pod size varies from 3.5. 0.7 ex in Z. formosa subsp. mollis to 24 x 1.2 em in Z. amazonica, There is some interspeilie variation in the texture (density) of the valves, ‘which is of taxonomic value, In this respect, the species may be arranged into thee groups: 1) Z. Lambertian and Z. flipes vith thin, membranes valves: 2)the remaining specins of subg. Zapotec, slong with Z. nervosa, with thick, membranous valves; and 3) Z. aculenta, Z. mierocephaia, and Z amazoniea, wily coriaceous valves, Wis i portant to mention that the terminology of pox Aexture use! here is relative, Wl as sch its we is not applicable to other stretuses (eg, Jealles) ‘The pads are glabrous except those of Z. cara. caxana whip. weberbaneri, Z. mollis, Z. porto: icensis subsp. pubicarpa, and Z. formosa subsp, saloadorensis, which are short pubescent, densely villous. puberulent, and pilose to densely pilose respectively Seods, In all species of subg. Zapoteea, the seeds ate widely ovoid to hornbesd and eompressed dnd lack ruptire ines, aris, or wings (Figs. 16, 17). In most members of this subgenus the testa is mntled, or dichrome, in which case the aroa encompassed by tho plourogram (ereola) has a lighter color than the outside area. There is con- siderable interpopulational variability in seul size land the coloration of the testa, especially in Z. formose, by contrast, Z. tetragona appears to be ‘ousistent in having dark, monochrome, ovate secs ther seed feature is the pleurogeain, common in subfamily Mimosoidese (Guan, 1984), Soeds of subg. Zapotec w 1. 90% plow rograme (sensu Gunn, 1984), although those of Z tetragona are always regular. The seads of Z ‘amazoniea difer fom those of subg. Zopotece in being narrowly rhomboid to nareowly elipti, larg br, ad in lacking the characteristic pleurogeam. NEXTPREV 790 ‘Annals of the Missouri Botanical Garden Ficunss 8-13. Unacetled grind plyad of Zapoteca speci —B, Z. media (R. Fersindes 3183) 4.2 alin i Herndndes 200) ake of pega ernst te lneshpedrvcure inact of he scsi gr ies 147)" thao Hl. Herndndes 950) 12.2 setragoms Gf Htemsndes 956, —13. 2. caracaranassbep. ueberbauer Johnson 2494, MO} bth sides of te payed are ‘Sewn: All colloctions at MEXU unlee oor inostd. Sale bars = 10 san Figures 6-10. 50 yn Figures 3. Seed dispersal. Zapoteca pods dehisce ex species of Zapateca over land and is probably plosvely with the valves splitting along both sutures effective over larger distances as a result of ac> from apex to bave (Fig. 15). This is probably the cumulative short-distance dispersal events over provalent short-range disperssl mechanism for many generations. NEXTPREV Volume 78, Number 3 1989 Ficunes 14, 18, Pods of Zapotcea tetragona (H. Mernindes 95 Seale bars = 3 em ‘This diepersol mechenism, however, doss not explain the presence of species such us 2. formes, Z. carucasana, ant Z, portoricensis on islands, Which they presumably reached by long-distance llispersal over water, Zapoteca formosa, for in stanee, oreurs throughout the West Indies and on some ilands off tho Mexican Pacific coast (Revilla tigedo Islands). Populations ofthis variable species frequently grow in cosstal areas. Although exper. iments have shoven that seeds snd mature pods of Z. formosa subsp. formosa taken from a coastal population of the Yueatin Peninsula floated for ‘nly two to four days in seawater, the sceds are highly resistant to saline water. As chown in Figure 16, seeds from this population (Herndndes 946) germinated after soaking in saline water (3.4% Sodium chloride) for up to 70 days. Most of the seeds germinated) normally when they were. me chanicallyscariied and transferred to petri dishes jn a medium moistened with fresh water. Ridley (1930)presented evidence that seeds and pods may Hornéndez Zapoteca | 14. Indeisced pod. —15. Dehisced po Aisporse for considerable distances floating on do bis, The great resistance to salinity of the seeds of Z. formosa, and presuroably of Z. carneasena and Z. portoricensis as well, suggest that these species dispersed by flotation from the continental ‘mainland. Farther diseussion about the distribution and disporsel of Z. formosa is provided below. ‘The taxonomic importance of seedling charac tors has been recognized by legume researchers {or a long time (De Candolle, 1825; Compton, 1912). Recently, such characters have buen ap. plied suecessfully in the construction of genesic and infragenerie clasifications in Leguminosae (Leonard, 1957; Vassal, 1969; Robbertse & van er Schiff, 1971: Kupicha, 1977; Guinet et al. 1980: and others). The dstintive seedlings of the species formerly assigned to ser. Lactevirentes in- dicated the heterogencous nature of Calliandra NEXTPREV 792 Annals of the ‘Missouri Botanical Garden Flows 16, 13 pororicensis (. Heraindes 948)—1 500 nam (sensu Bentham) and provided evidence for rec: copnition of Zapotece (Heminder, 1986s). Seedlings of 19 populations representing cight species of Zapatera, all belonging to subg. Ze poteca, were studied (Table 5). About 10 me cchanicallyscarified seeds per sample were germi- nated in peti dishes. After one to three days, when the radicles hed emerged, the germinated seeds were transferred to individual pots. ‘The seedlings were described following Duke & Polhill (1981), E10 3 6 an g 20 «40 =~]. days Ficume 18, Resitance of Zapetea formosa cube. formosa swede to saline water (3.4% sodium chloride) “The bars indicate percent of germinated seed. Ten sees yore ald ab coe Gr level Seeds of Zapotecs species showing the iregular pleurograms.—16. Z. portoicenss subsp. 7: formosa (Torres 4083). Calectons &t MEXU and MO. Seale bare = The principal characters used in the descriptions ‘wore! 1) position (whether epigeal, goal, oF hypo geal), morphology, texture, and degree of persis tence of the cotyledons, and 2) type (whether pin- nate or bipinnate) and sequence (whether opposite fr alternate of the eophyls ‘These samples were remarkably uniform, the seedings of each species examined being essentially Identical. Diagnostic characters atthe species level. ‘such as the square stems of Z. tetragona oF the ‘ovate leaflets of Z. formosa, beeame manifest above the third eophyll, 25 the fist metaphylls arose. A Drief description of the seedlings listed in Table 5 follows (Figs. 19, 20) Germination phaneroepigeal cataphylls absent Cotyledons sessile, elliptic to cliptic-ovate, foia ceous, ephemeral (remaining functional for ¢a. one week), Leaves stipulate, glabrous; frst two eophylls ‘opposite and pinnate, the pinnae with 4-12 lan cealate to narrowly oblong leaflets; thicd eophyll alternate, bipinnate, with I pair of pinnae; fourth cophyll alternate, bipinnate, with 1, rarely 2 pairs of pinnee, In summary, although the seedlings of subg. Aculeara, Amazonica, ond Nervosa remain un- known, the seedlings that have been observed so far, al in subg. Zapoteca, display a high degree ‘of inter- and intraspecific morphological homoge nity. NEXTPREV Volume 78, Number 3 1988 Flounts 19, 20. Seedlings of Zepoteca.—19. Fveday ld sending of 2. teiragona (H. Hernéndes 956) 20, Three-nerk old seeding of Z portricensissubop. portericensis (H. Herndades 948), Cllections at MEXU a MO. Seale bars = Ler. DIFFERENTIATION ‘There are several morphological features com: mon to all Zapotcea species. Among the most important are: brochidedromous venation patterns in leaflets; persistent, leafy stipules (exeluding Z. ‘aculeata); homomerphie, sphevica, capitate info rescences; cup-shaped sigras; 16-prained polyads with prominent leneshaped thickenings (exhiding Z. nervosa): and dry. linear, fattened pods with thickened margins and showing @ particular mode of dehiacence. The distribution of morphological features among the species of Zapoteca reflees four evolutionary Hernandez Zepoteca lines. which correspond to the four subgenera pro- posed in the systematic treatment. Subgenus Nervosa comprises Z. nervosa, which appears to be morphologically the most isolated species in the genus. Superfically, this species ap- pears out of place within Zepoteca; however, the morphology and the type of dehiscence of its pods and the overall features of its inflorescences justify its placement within thie gers Zapotece nervosa exhibits sovers] characters unique im the genus, the most important of which are: stems with fissured bark; large, coriaceous, grayish leaflets with a peculiar eucarptedromous/ brochidodromous venation: and 16-grained polyads ‘Taate 5. Species of Zapoteen in which seiings were ue H = author's elections Callecton Dat Dasriation Z clince Torres 3047 QO, MEXU) ‘Mesieo, Oaxaca 2. carncasana subsp. weberbaeré Jokason 2494-82 (MO) CColorti, Vale Z.formora aby. formosa Tohason 183-78 (MO) Mexico, Yueatén Z formosa ssp. formosa Torres 4262 (MO, MEXU) Mexico, Oaxaca Z formosa sop. formosa Johnson 490-78 (MO) Mexio, Yueatén Z foumona wep. ones okason 1270.79 (M0) Monica, Goereero Z formosa sep. rose Torres 6012 (MEXU) Merieo, Oaxaca Z formosa sip. sehoeti Starr & Birgy 169 (MEXU) Mexico, Sorora Z. formosa shop. schottis Felgor 84-124 (MEXU) Mexico, Sonos Z tambertiana deonte 134 (MO, XAL) Moric, Veracrur 2 tambertiana Martine: 0. 284 (MEXU) Mexico, Tamaulipas Z media Delgedo 53 (MEXU) Mexico, Hilal 2, pontricensis soba. portoricensis H1.154 (MO, MEXU) Merien, Veracrur 2. tehuay Torres 4167 (MO, MEXU) Mexico, Oaxaca 2, turagona 1-107 (MO, MEXU) Mesico, Veracruz 2, tevragona #7675 (0, MEXU) Conta Rice, Cargo Z tetagona 11.800 (MO, MEXU) Mexico, Verceur Z ttregona H.327 (MO, MEXU) Mexico, Veracruz Z.tetragona Johnson 1562 (MO) Menten, VeractuePREV 794 ‘without lenshaped structures, In this connection, it is reasonable to hypothesize that represents an evolutionary line that separated from the main stock cary in the evelution of the genus, and its peculiar morphological features would be explained hy divergent evolution. The laek of are late, lensshaped thickenings in the palyads of Z. nervosa, a character also absent in the vemeining genera of the Ingooe, appear to representa prim: itive condition within Zapotec and suggests that this species is phylogenetically basal in this genus Additional information ocr this critical species is needed in order to learn more about its phyloge- netic relationships, Subgenus Zaporceu i the most widespread and diversified subgenus in Zaporeea. Yet a number ‘of morphological features common tall the species ‘make this suhgenns easily recognizable. Al the species have pinnae with several to many pairs of thin, membranous leaflets, whieh show character: istic usually inconspicuous and simplified venation patterns. The highest diversity of ir types (three types) of the genus i found in sug. Zapoteras the species of the remaining subgeners are char. acterived by having only the unicellular conieal type (Table 4). Further, the valves in the gods in subg. Zapoteea, along with Z nervose, sre eon sistent thickemembranos, wheres those of subs, Aculeata and Amazonica are coriaceous. The pattorn of leaflet venation in Z. zchwana difers from those of the other species in subg, Zapoteca awd shows strong similarity with those Of Z aculeata (subg. Aculeuta). This may indicate fa phylogenetic link between the two subgenera. Subgenus Aewloasa contains only Zaporaea aculeata, wbich is characterized primaily by spi los (x characteristic that has elearly been derived within the genius), by an apparent Arhorescent habit, and by gine hoving fewer pairs of chariacoous-coriacoous eafits thon those of subg. Zapotoea. The curiaceans gods of Z, acc leata resemble those of subg. Amazonion beens Amazonice is made wp of Z. amas soniea and Z. microcephala, which are chatac- terized by having the most obvious combination of apomorphie features in Zapoteca. Their leaves are reduced to two or three large, coriaceous leaflets in each pinna. This inthe rent of Arend toward reduction ip the m associated with increased lellet size, This tend is also seen in other goncra of Mimoscideae (e.. Calliandra, Mimosa). Moreoxer, the leaflets in those species have « number of common architec: Annals of the Missouri Botanical Garden tural features reflecting an increased level of or: ganization, which s also interpreted as an advanced condition. A progressive trend toward suppression ‘of leaves subtending the inflaresoonce in Zepotcea has remlted in the highly specialized, elongate psewdepanicls ofthese species, Among other clearly apomorphic features of subg. Areazonice are the clongate sends of Z. amazonice, which lack the chavacteristic pleurogeam and are the largest in the gers. Biocroceapiy Zapoteca ranges from the southwestem United ‘States (southern Arizona and southwestern Texas) ‘and northern Mexico, from about latitude 32°30'N, southyeard through most of Mexieo, Central Amer. jen, and anuch of Sonth America (Venemuels, Co- lombia, Feuador, Peru, Dolivin, Guyana, Brazil Paraguay) to northern Argentina (ca. laisude 27°80'S). Zapoteea also occurs on most of the West Indies and in the Revillapigedo Islands (ie, Ila Sovorra and Iss Clarin). Zapoteca species aceur ina variety of environ iments, ranging from open, arid or seensrd,sceub: by vogetstion types (e.g. Z media, Z. allnac, forms subsp. mollcula, and Z. formosa subsp ‘schowti’)to the wet evergreen forests of Amazonian Peru (Z toricensis ssbip, flavida and Z, telragona. ave found in relatively wet, lowland habitats, genorelly fon disturbed sites, The greatest degree of diver Sifeation has ve ‘une-zonica). Most populations of Z. por el ny eas covered by sea sonal, tropical deciduous forests (sensu Breedlove, 1981). In addition ta the species found in clearly arid regions, most populations of cight species of Zapotcen\Z. nervosa, Z, lambertiana, Z.flipes, Z. mollis, Z caracasuna, Z.andina, Z. formosa, and Z celiwana) occur in topieal deciuous for: "eats, In addition, Z. portoricensis sulsp. portor: censis and subsp. pubirarpa primarily aceur in middle-elevation 02k, pine-oak, or subtropical montane moist forests The occurrence of three of the four subgenera ‘of Zapotecs in South America (Zapoteca, Act lata, and Amazontea), compared with only one Jn Mexico and Courral America, is probably « con. soquence of the mote ancient existence of Zapo ‘ecu in South America Although only subg. Za: al potece ix represented in Mexico and America, extensive diversification has taken glace at the specific and subepecie Levels, especially in Mexico. For instanee, there are seven subspecifie taxa in Mexico (.e., Z portoricensis susp, por toricensis, Z. partaricensis subsp. pubicarpa, Z. NEXTPREV Volume 78, Number 3 1989 portoricensis subsp. flavida, 2, formosa subsp. formost Z.jormosa aubep.rosvi, & formosa sobsp. ‘mollicala, Z. formosa subsp. schottil) four of them restricted to this country. In contrast, only four tmpecies of subg. Zapotera occur in all of South America 42. porloricensis subsp. flavida, Z. earacasana subsp. caracasana, Z. caracasana faubsp. tceberbauert, and Z, formosa subsp. for ‘mosa).and only one ofthese, Z.caracasana subsp. weberbawer, i endemic Viewed from a global perspective, there are no lear centers of species diversity in Zapoteca. The total numberof spevies andthe nimber of endemic species are reughly correlated wth the sizeof each region (en Mexico. 7 spciae/endomiosi Cantal America, 6/2: West Indies, /1: and South Amer. ica, 10/6. However, « more detailed analysis of the areas of species concentraion and the distr bution of endemic toxa revealed sev vith high diversity (Fig. 21) Southern Mexico Oaxaca and Chiapas, includ ing southern Puebla) has the largest concentration af specien and walapecis of Zapotec. Ths tele tively stall egion, characterized hye remarkable climatic and topographie diversity, contains seven species, and if subspecies aro taken inte account, the figure rises to eleven taxa (Z. media, Z.alinae, Z lamberiiana, Z. portoricensis subsp. porter censis,Z. portoricensis subsp. pubicarpa, Z. por- Toricensissubsp.flevida, Z. tetragona, £ formosa subsp. formas, Z. formosn subsp. roset, 2. fore ‘masa subsp. molicila, and Z, tehisana). Four of these are endemies (Z, alinue, 2, portoricensis sulsp. pubiearpa, Z. formosa subsp. mollicuia, and Z. teliuana), especialy wo dry areas of Oaxaca and the arid valleys of the Pueblan-Oaxacan bor- dee region. The number of species and subspecies ‘of Zapotec silenly decreases from this center in all direction: (Fig. 21). On the Pacific slope of Mexico, from Guerrero northivards, there are four taxa (Z. tetrogona, Z. formosa subsp. formosa, Z formosa subsp. rose, sad Z, formosa subsp. schti), deereasing tone in the northern portion. Zapoteen media and Z formosa subsp. formosa ‘are the only toxa that occur in tho Central Platea Along the entire Atlantic lope, from Tabuseo to ‘Tamaulipas, together with the Yucatan Peninsuls, there are only five taxs (Z. fambertiana, Z. por Loricensis subsp. portoricensis, Z. portoricensix slap. flovida, Z.tetrogona, and Z. formosa samp formosa). Zapoteen lambertinna is the only one ‘of these that cecontally endemie to this region In Central America, there are 4 total of eight taxa Zapoteca portoricensis subsp. portosicensis, Z portoricensismubsp.flavide, Z. mollis, Z care: Hernindez Zapoteca 798: oasana, Z, tetragona, 2, costariconsis, Z, for ‘mosa subsp. formosa, and Z. formosa subsp. sal tudorensis). Diversity is quite uniform among the tliferent countries (Fig. 21). Zapoteca ruolis snd Z. costaricensis are essentially endemic to Costa Rica, although a disjunct population of the former is knowa from Panama. Zapotece formnose 2ubsp. salvadoronsis is restricted to southern Guatemala and BI Salvador Six taxa of Zapotecs oecus in the West Indies (Z nervosa, Z. portoricensis subsp. portoricensis, Z. portoricensis subsp. flavida, Z. curacesana subsp. caracasana, Z. formosa subsp. jormosa, ‘and Z. formosa subsp. gracilis). Genecaily only a single species oveurs on a particular island: how: ver, in Hispaniola the four species, including the cendemic and morphologically distinct Z. nervosa focenr together. Howard (1973) pointed out that Hispaniola and Cuba are the most flonsticaly di- verse islands in the Weat Ines ard that they show arcat habitat diversity and highest percentage of cendemisin (Hisganiola with 93% of endemic gen: era). Zepoieca formosa subsp, gracilis isthe sec fal erdeanic entity im the West Todos, [kis marily restricted 9 enastal areas in Guba, the Bahamas, Cayman Islands, and sorthwestern Heit. Eleven taxa of Zeporera occur in South Amor” ica. The greatest diversity oceues ity the teopical ‘Andean countries, Venezuela, Colombia, Reuador, Peru, and Bolivia, where popiations are primarily foun in the foothills of the Andes, in the inter= Anlean valleys, and less commonly in the Arges Bosin. As illustrated in Figure 21, the number of taxa in cach of these countrice ranges from: four to six, with Colombia and Ecusdar most diverse. Perhaps the mast remarkable bingeographieal fea. tuce of this whole ares is the occurrence of five tendeanic taxa, which, excluding Z. carncasana aubsp. wcherbaveri and Z. andina, comprise the rect highly advanced epecios in Zepoteca, each with a large nunabier of derived morphological cha oters (he., Z. aculeata, Reuadorean Andes: Z rmicrocephala, Median Valley of the Magdalena River: and Z. amazonica, Amazonian Peru). Oute side this area, diversity is meh lower, with one oF two texa occurring within Guyana, Brel, Para guay. and northern Argentina (Fig. 21). Within this large region, however, there are two extra Andean enderic species: 2. flipes, restricted to the Brasiian states of GoiSs ancl Minas Gerais, and Z. sentelifera, fox in the states of Amazonas, ord and Rondénia, Brasil, and in the departments ‘of El Boni and Santa Cruz, Bolivia. Zapotece species are predominantly segregated engraphically and/or ecologically, although 3 NEXTPREV 798 ‘Annals of the Missouri Botanical Garden ous 21 treat a> seporao wit. Total amber af nurnber of cases of gross geographical overlap are known, However, oly « few cevords of true sya patey exist: in the District of Tlacolla, Oaxaca, plants of Z. alinae (H. Hlerniindes & Ramirez ‘942) have been foun iatermixed with plants of Z formowe wibap, rosei (Uh. Hermindes d Ramirez 43}, in this same ares of Onxace, Z. formosa subsp, formosa las occasionally bees Found grow= ing alongside Z_ formosa subsp. roset (IT. Her= andes & Torres 210, 217 and Magallanes 111. 113). Meo, in the tropical decichions forest of El Chorreadero, Chiapas, Z. lanbertiana (Breedtove 39604, H. Hiernindes & Ramire: 891) ani Z. onoricensis subsp. portoricensis (Breedione 11889) bave been collected; however, itis unclear cshether these are nasrowly sympatric. ‘There are some presumed hybrids, For example, in ether Gusternals amd in southern Mexico, a ‘number of collections provide evidenec for exten= sive hybridization between Z. portoricensts subsp. pportoricensis and Z. otragona:in southgen Puobla and northern Oaxaca, come plants with characters intermediate between Z. media and Z. formosa likewise suggest hybridization. In Guereero popu [Number of Zapotec species and subspovas per county. Mesiean tosh and Bralian states ere ‘endomie "ans lations of Z. formosa subse. formose and Z. for: ‘mosa aubep. rasei como into clo proximity. ond there area nurahor of collections of prabable yd origin, ‘The predominance of allopatrie distributions and the apparent rarity of hybrids in Zapotec suggest that lybeidization has played an insignificant role in the evolution of the genus. The few cases of siriet syimpatry may represent secondary contact of previously diferentiated populations. It could he hypothesized that speciation in Zapotec bas pr rmardy resulted from genotc differentiation of goo graphically isolated populations. There are wide disjunetions in the disttbution of Z. formosa subsp. formose in South America, from nocthern Colombia snd Venezuela to Bolivia, southeastern Beal, and Paraguay (Fig. 44). Av elaborated below, its distribution was probably con tinuous acres the Amszon Basin during the cool Airy phases of the Pleistocene, when a drier ond presumably more suitable environment for this sub spocies was widespread (Ab'Séber, 1982: Abey, 1982; Bigarella & Andrade-Lima, 1982; van de Hammen, 1962; Prance, 1982). Zapoteca scu NEXTPREV Volume 76, Number 3 1989 tellifera iz closely related and is known (rom faw disjunct populations inthe Amazon Basin (Fig. 43) Specialized characters in Z. seutellifera (e.g, sci telform extrafloral nectaries, scanvlent habit) and its presumod edaphic specialization suzgest that it ‘vas probably derived from 2, formosa subsp. for moss. Zapotece seutelifera could have been de. rived from populations of Z. formosa susp. for: nose that became isolated after the return to the present-day. warm-wet climatic condition ‘This patter, along with the patehy distributions of dry-foreat species like Z. lambertiana and Z. mollis, supports Gentry's (1979, 1982) theory of reverse refuues. This theary assomes that the pres. 1 discontinuities in the distribution of the dry Forests [dry forest refuges) couples with the exis: tence of geographically restricted, morphologically Uifferentiated dry-area populations, result from a shift towards more mesic conditions after the ei toatie deterioration in the Quaternary A second exarnple of possible allopatric specia- iow in Zapoteen involves the tre members of the highly spotalized subg. Amaconion: Z. mieroceph fla and Z umazonica, Zapoieea microcephale is restricted to the Middle Magdalena River Valley whereas Z. anazonia occurs in Araszanian Per (Fig. 47). These two species, shich are obviously ‘closely related, probably diferentiated a2 a con sequence ofa viearianee event eansed by the uplift of tho marthorn (nds, whieh rocched their prosent clevation atthe beginning of the Pleistocene some theee milion years ago (van dec Haminen, 1982) ‘Adtivional evidence suggesting an allopatric model of speciation for Zapoteea is the relatively common seeurrence of closely relied taxa dis playing contiguous oF nesely contiguous distribu tome, This distbution pattern is shown by closely related spevies, such at Z. tezvagona and Z. an dina (Figs. 40, 42), and by vietually all the subs species in the genus (Figs. 36, 39, 48) Rernooucnve Browony During the past few years a considerable body of data has accumulated on the diverse reproduc tive mechanisms of the Mimoscideae (S al, 1977: Kenrick & Knox, 1982: Kogtur, 1983, 1084: Knox & Kenrick, 1983: Hopkins, 1984: Bernhardt, 1988). Ensomophily was probably the original mode of polliaation in the sublaraily, as indicated by the presence of apparently insect, probably hymnenopteran, pollinated mimozod info: rescenees im the Middle Eocene Claibome For: ‘mation, Tennossee (Cropet & Dilchor, 1977), and in the Oligocene of the Texas Gulf Coast (Dughlian Hernandez Zapoteca 797 et al, 1980), and also by the wide distribution of this pollination sy subfamily. Ithas been suggested (Raven & Polhill, 1981; Polhill t al, 1981) that Mimosoideae were already. differen tiated fom the other subfamilies by the start of the Eocene (54 my. B.P.), Entomophily is wide spread in ihe Mimozeae (2.g., Desmanthus, Len. caena, Mimosa, Prosopis) which is generally con silered to be the basal tribe of the Mis Furthermore, most species af deacia (Acacieae) are insect pollinated (Bernhardt, 1983: pers. obs: however, bird pollination has hoon derived soeand arly in some Australian species ofthis genus (Fort & Forde, 1976; Beroharit, 1988; Kensick etal, 1983). In contrast, Parkin (Parkieae) is primarily batpollinated (Baker & Hervis, 1957; Hopkins, 1984), although 2 few specics are pollinated by ingeots (Ge divewssion in Hopkins, 1984). High-energy pollination systenss such a6 those involving hawkmoths, birds, snd bats are concen. trated among the genera of the more advanced tribe Ingeae (Cron et ol, 1976; Arrayc, 198: Koptur, 1983: Hernandez, 1986a; unpublishod be.) In the Ingeae, the transition from low-energy nation to high-energy systems, which ‘greater allocation to aectar and pollen, is te with inloreseenee muaifeations, For i= stance, eonnation ofthe stamina filaments t form ‘tube and increase in number and length of the filaments made it possible for the Aowors to hold larger quantities of nectar necessary to satisfy the ‘energy requirements of larger, homothermiv pol- linators. Similarly, changes in periodicities of nectar and aroma preduction, neetar quality, and timing, ‘of anthesis have enabled the plants to adjust to the physilogieal end behavioral characteristies of par ticular pollinators Mimosoideae have a number of ad productive traits that make them cwoidene. sting from an ecological and eve of view. Guinet (1969, 1981) discussed in detail that the most conspicuous feature of mimosoid pollen is the occurrence of permanent units of dlispeesal in the form of tetrad, oeteds, and polyads throughout the subfareily. Such units of pollination presumably provide the plants with a selective ad ‘vantage in oprovduetion. As has been demonsteate for Aeacia species (Kenriek & Knox, 1982: Knox & Kenrick, 1983), a single pollen unit is able to Feri all the ovules in the ovary. Another vide spread feature of the mimosoid inflorescences that is important in the bree is the occurrence of female sterile flowers. Viewed in an evolutionary perspective, theve reproductive phenomena have great potential as characters on systeti of the species NEXTPREV 738 which to base phylogenetic interpretations. Pub: lished reports concerning he reproductive biology of the species here assigned to Zapotec ate vir. tually ooneaistent, and the present chapter is in- tended to provide a Dolnervations, of some of the repreductive Features of thece plants account, based on my own Nectar volumes wore measured with calibrated ricropipets, and cugar concentrations were deter- mined using/acempersture.compensated Erma hand refractometer. In 4 few cases, when the (lowers were very stall, a syringe was used to extract the nectar from the sinninal tubes. All measarements sreve cle inthe greenhowse early in the morning, in order to estimate the total amount of nectar produced por lower. Between four and tom indi viduals per population were examined. Tei quite posible that the results in Table 6 might be lower volumes than: would be found in the wild, since it has been shown that nectar volumes and sugar concentrations may be significantly lower in plants roving in greenhouse pots compared with wild plants (Cruden & Hermann, 1983), However, they allow species comparisons. Field observations on flower vistors were carried ously ducing the frst three to five hours ince all Zapoteca species flawer at he ehavior of Fawer vistors was observed of anthesis, hi svith the aid of a head lamp. In most eases, visitors wero euceessfully collected with ineeet nets. Insect voucher specimens: are deposited in the Enton logical Collection at the Instituto de Bicloge UNAM ‘The number of ovules per gynoceinm was de termined by squashing fixed avaries in a drop of Alexander's stain (Alexander, 1969): the ovariee ‘wore examined with a compound Tight mioroseope Within 15~20 minutes after staining. Sex expres sion was characterized for each population by dis secting fixed flowers and counting the proportion lf hermaphrerte and feralesteile flowers ix each inflorescence, Selfscompatiblity was assessed on plants growing in the greenhouse using the tech ‘niques outlined by Bawa (1974) Pollinaions were accomplished using 2 fine brush within four hours ff the beginning of anthesis, once the anthers were ehiscens. Antber dehiscence was determined by touching the anthers with a clean microscope sides 4 large number of polyads on the slide revealed ‘open thecae. Given the small size af the Bowors, the inflorescence was taken a6 ¢ unit in the poll nation treatments. In every population, a deter mined number of inflorescences were artificially Annals of the Missouri Botanical Garden croct- and self-polinated, and several additional inflorescences were left untreated (apen) to test for ‘autogamy, Due 10 the dificulty iavaived in emas- culating the delicate lamers of Zapoteca, the pos lity of apominis was not tested. For the crass prllinations, mixtures of patyads from several in dividuale wore used whenever possible. For the self pollinated infloreseences the pollen donor vas taken from the same inflorescence oF from other inflorescences inthe same individual; therefore, a great number of thewe flowers were geitonoga- rmonsly pollingted. Treated inflorescences were marked with hanging labels, As the greenhouse was insect free, inflorescenee manipulation wat mini mized by avoiding the need to bag inflorescences. ‘Suunple sae varied fcom two to nine individuals per populition, Afior each treatment, brushes nd hands rete sterilized with ethanol. Since pod abortion ceuirs in Zapoteca species at several stages of pod developaent, the resulting pod> were counted only i they matured fully. Seed sot was estieaated jm every population by calculating the average inucber of fully developed seeds in mature pods Although i i difficult to eharacterine the phe: nological behavior of a given species throughout sts range, [shall attompt to draw some gecerali: ations cbout the flowering patterns in Zaporeea ‘This discussion is based primarily on species re strictad to Mexico and Centeal America. Tn wide Funging species such as Z tetraona, Howeving timex vary fron ane rogion to another. This species, inbieh usually grows in secondary vogetation de ved from losiand tropical wet forests, flowers testy during the dry seacons nevertheless, in the Mexican state of Morelos, where a more seasonal climate exists the populations flower primacily ing the rainy season. Zaporere portorivensis subsp “fasida, whieh grows in wet habitats, also Rowers ng the dry season. By contrast, the species growing in dricr habitats (thorn forests tropical ‘deciduous forests, and tropical semideciduous for sts) tend to lower predominantly during the sec: cod half of the rainy season (eg., 2. formosa, Z lemberiana, Z media, Z. alinae). The species fron drier areas have shorter, more synchronized Nlowesing, perils than species living in wetter hab itats e.g, Z. amazonica and Z. retragena), which continue to produce flowers for longer periods. Finally, Z. prtoricensissubsp, pevtoriconsis, which generally grows in montane mesophyllous forests (Geriinclogy following Reedowski, 1978), abe Bevee NEXTVolume 76, Number 3 1989 Flounes 22-26. ertncensis shop. porocenste(H. Herts Sat Gi isnt 1s : : A possible explanation for this diflerential flv ering pattern ie that in wot areas excessive rains may have destructive eects on the delicate info resconves of Zaporeea and may reduce the activity of potential pollinators (Janzen, 1967). I observed the devastating effects of rain on inflorescences of several Zapotece species in wet an dry forests. ‘These effects are clearly reduced in arcas of precipittion As far as I have observed, species of Zapoteca display two distinct flowering strategies. Most species (eas Z. formosn, Z. media, Z. portoricensis, Z letragona) have # very generalized flowering be havior. Here regular numbers of inflorescences open ‘every night over several months. By contrast, Z: ‘linae exhibits a more specialized strategy. Ind viduele of thie species intermittently produce re: atively large numbers of inflorescences on scattered Inforesences of Zeposca species et anthesis al photgrapis west taken at night — formosa ssh, formona ogi) on 2. formosa so. roe et) Ut. Herebndes 216, 212, ks 28:2 sesagona Ul. Hermie: 920) --28. F mda 2 tehuana (H. Heradndes 850) Al elections st MERU ad MO. Hernéndez Zepoteca 799, respectively) — nights throughout its flowering period, from July to September. This specialization is similar to the “multple-bang” strategy described by Gentry (1974) for tropical Bignoniaceae. Tt differs, how ‘ever, in thatthe flowering i not synchronized among the different individals that make up a given pop- ulation of Z, alinae. One possible adaptive value of this strategy is that these flowering bursts pro- ‘duce conspicuous visual and olfactory stimuli, thus increasing the chances of pollinator attraction Species of Zapoteca always have their flowers arranged in compzet, homomorphie, capitate in floreseences, which act as units of attraction Figs. 22-26). In each flower, the numerous elongatePREV ‘Annals of the Missouri Botanical Gardon Tame 6, Volume and agar density of nectar in seleted spies of Zapotera, All elections is MO. H = authors etons, Votame ia ‘Sopa desi OO) Taxon reion Data CSD.) 2(5.0) Zalinae Mevico, Oxsea, HM? 3~ 3 mm, brawa, dark brown, or black, the areola sometimes mottled: plenrogram irvegular, some: times regular. Somatic chrommosorne number 2 26. Distribution (hig. 86). Frequent in open sites seh a9 forest clearings, forest cdgos, roadsides. rivor banks, and wet thickets: in general in see ondary habitats derived from a variety of vege tation types. Gallections come from elevations of (0-2,100 m. Standley (1930) incorrectly included this species in the Flora of Yueavén Zapoteca portoricensis as circumscribed here, consists of three rather variable subspecies: por torieensis, pubicurpa, and avila. Foch has a fow distinguiching, variable charactors (Table 12), although they iatergeade and overlap. Consequent Hernandez Zapoteca ang ly. these subspecies are defined on the basis of ‘combinations of characters. Viewod ina loeal con text, each subspecies might be accorded specific rank; however, 2 synthetic analysis of all available material reflects the absence of strong morpholog ical discontinuities distinguishing ther. The three subspecies of Z. portoricensis have inet habitat prelerences: subsp. portoricensis ‘oceurs primarily in relatively wot, mid high altiude habitars; subsp. flasida is found in wer Toland areas; and subsp. pubicaspa oecurs in dey, inidealtitude areas. However, in the West Indies, subsp. portoricensis usually occurs in much drier, Tov, sometimes coastal, habitats. Lo, Sheubs seandonts eats 7-20. 26 rem: forolls 3-4 om longs marwe pods t 16.5 ex, long e. subep, avid Lb, Shrubs erect fais 4-13-15) > 1-3 bam corolla 4-U(-10) ra log: mature pods to 12 (14.5) om long 2a Pods glbrous, rarely icy: peduncles (1B-)85°12015) em kg. at athe fealys ghalzour to cist, the teeth rarely Sparel slows; coralla 4-2 mi loog. 62, subsp. portriconsis ede paberlens”peduncks 13-8.5(-6) fem long at anthess: calye teeth coms fendy sous; cools 7-24-10) ma long rn Ob sep pabearpe 2b. 60. Lapoteca portoricensis (Jacq) H. Hern. subsp. portoricensis. Mimosa guincensis Sum. & Thon, Bsks, Guin. PL 32s 182, facia linearis Does. ex Ham, Prode. PL. fo. Oecid, 59 1825. 4: hamilton Dose. ox am, Prod, Pl ea Osc. 59. 1323. A. alt Coll, Hor. Hipal, Appendix 2: 339. 1820, 4. olla. K. Pre, Reports Bot. Spat 183. 1835, 4 ospentina Macg., lars. 1-318. 1887. 4 1 legate Den Bot Ag. $268, 1854 velo: mcrae rnc. Bre WL 225, 1860, ‘re Janaien, Marchs Gukstype, GOET (a cea, sorygen GH. NY}. Caltandre nicarginensis Tab, & Lows x Loe, Box Tet, Syac 2 134, 189% tnaieaicaraguen fs Cauh. & Loos. ex Loes) Brkion 8 Row, N. Soe. #1. 25 G3. 1928, Te: Niseraguae Mata taps: Cofads Yasoa, 1,000 m, 4 Aug. 1843, Kethscih 204 (ectotyge, US no. 1260108 (ee tment and phot) here designated eletotype, NY {Gogmenth. The koto, wich nan 8 wre do stroyed in 194: (Pe Mepko, pore comm 1586 shuts ofthis sprcinen are at F, MEXU, M0, NY. Texts Calliandra portricensn var major Sprapse, Bo Mag, Yon B120. 1902. row Gul How, Reg Ker, ug. 1906 (ecurype bere designated, K). ts specimen’ was presumably calested by. Sprague NEXT820 Annals of the (Missouri Botanical Garden Ficuns $6. Dutrbution of Zapotesa portoricens auep, partaricensis (open squares Z. portorieensis sep pubteorpa starch and Z. portoicenstseabsp.fetida (aot) “pes met is dacs hat “he Kew plants were purclased in 1906 fom a nursery fi Bytcee as loge alba? and Rowered the Tem porate House in June and uly of tit Yor.” Ania sprague Beton & Rose, N. Amer. E25: 96 1928, Caliandra sprazuc (Briton & Rose) Lan dell, Lydia 2: 89, 1989, Ture: Mexico, Veracrs Xalapa, Schiede 660 Ueeweype here desgra, NY Keleeityen, BM, CAS, CCE, G, GH, Ky LE MEXU, MO, UC, US fragment. The specirers the only ence Inked o cleeone ""Sehtede 690," whereas the spaces st Ted anc Dey 690. “The specimens from BM and Gare loc "Selurde Tams 12. Comparnon of the sabspecies of Zopeteea pororicentis. Sulspecis poraricensts ‘Subspecies pubicarpa ‘Sabipecoe flavide inane ret she exec crab scandent Pais of pianee —(1-)2-20-8) 2 850-7) ‘Shape narroy oblong to anceste nariowly oblong to lanceolate oblong to abn davenaine lew 5-15 x 1-3 mm ate 2 mm 1-20'% 2-6 mm Pairs pina (1 )10- 15-30 18-36 Stile fog 10-17) am 1 me 1¢13) xa Peduncle length (18-185 121-19) em 1.3-3.5(-6) om 375 cm Calyx Tena Lamm 2-0(-8.5) aa 1-2 mm Induiment gla t liste tious wo demely villas gloom Cercle length 4-8 em 78-10) mm Sum Ped Lengsh 12-145) om 6 om 16.5 em Tndument___glabveus uberent aroPREV Volume 76, Numoer 3 1989 rd Dapge" (a) and the oe ot UC i abled ‘Sstiede™ a0) The specimen CAS, COB, GH, sat MEXU cot have any relerore fo olson {Ith seconde on thea fort nae el: Srauestotaspinenavcaecte by Shea, Wik {he eacepion f three HM, MENU, nd UC, all {he oles are had anata ae coe pre lovionss Wilk,” aed lng oon the boner. fq the eter dts proved, sd te spent of the flail shecesesimone amor eran a Fists ofthe same Sohede 090 collection. The Ipockmons ct G, GH, and NY havea nr, pro ‘hip mate w posterior, wh the pane “Sehioc" eves, tell brown “bat Selector da Sever aveed otal Western Europe. atrogh « Stan of eller fom Msi hve mistakenly ‘tn othr th Aevording to BBaracky (Ger. cot) Seueluemia dined Sede ar Callondra nocalonss Lande, Uys 2: 88, 1999, Tare: Maxie. Veracraz: Nose, Es Meru 1133, Peg 198 toleypes MIC trp, F, MEXU, Ny Caltianden sitepenis Lindel, Poyaogin 2c 1, 1961. TaPE Mesa, hayes: Barranea Hd, Siepee OGTR 184 Sasa 4040 Ck, IC Sey A, Fe MEXU, NY} Erect sheuhs, 2-3(-6) m tall: stems slender, sometimes to 12 6m dian. ot the haze; branchlets terete, sometimes dangled, glabrous to densely villous ith tawny Isirs, often glabrate. Pi (1-)8-7C-8),jugate; leaflets (12-16-67 pasts per pinns, narrowly oblong to lanceolate. rarely oban cealate, oblique at the ase, acute to rounded at the apex, usually mueronulate, 5-15 1-3 ma, labrous to villous, puberulent, or sericeous on both surfaces, usually ciate; stipules tang rowly triangular, sometimes slightly curved, 10 (17) mm long, glabrous to villous. Capitals axl lary; peduncles (1.8-)3.5-12{-15) em long a thesis, glabrous to densely villous. Celyx 1-3 mm long, the teeth usualy thiangular, sometimes lan ceolate to hinear-lancealate, usually glabrous or eil- tate, sometizes with a few hairs in the area of the teeth corolla 4-8 mm long, glabrous. Pods to 12(- 14.5) © 11.2) cm, straight or slightly curved labrous, Seeds widely rhomhoid to oveid, 5-7 4-5 mm, brown to black, the ereola sometimes mottled; pleurogeam usually irregular, sometimes regula. Somstie chromosome number 2n = 26. Distribution (Fig. 36). This widespread sub species has been collecte’ in sexeral relatively Io ‘alized areas: 1) Atlantic slopes of the Sierra Made Oricatal, in the states of Hilal, Puebla, and Sa Luis Potosi acl the eastern portion of the Trans vere Voleanic Belt in Veracruz: 2) Sierra Madre lel Sur, including parts of the Mixteea Region in Oaxaca: 3) Sierra Madre of Chiapas, and the Chia: Hernandez Zapoteca 82 ‘pas and Guatemala Highlandas and 4) West Indies, in Jamaica, Hispaniola, and Puerto Rico (including soine of the Virgin Islas). Disjunctions eecur i the Sicera de Tamaulipes, Tamaulipas: Temascal- tepec, State of Méxica: and in Hendurasy Nica agua, Costa Rico, ond Panaraé. Thore ie « col lection from Nova Friturgo, Rio de Janeiro, Brazil (Curran 655), which wan probably obtained fom cultivated glants. This subspecies has usually heen collected in mountain areas of moderate to rela- tively high elttue (800-2,100 m, in plant com munities such as oak, pine, pine-ouk, temperate deciduous, wet montane cloud, axl montane rai forests, especially in seconlary habitats. Never ‘holo, in come areas such as the Huastces in the states of Hidalgo and San Luis Potosi, Mexico, populations usually accur at lower elevations, reaching as low as 165 1. Siilaely, plants fron the Wert Indies usually mhabit low elevation areas, down to c0a level, with « much drier climate Common names, Barba de chiva (Niesragua), carboneillo banca (Costa Rica), granoline (Darvin can Republic). guseamaya montés (Bl Salvador), gunje cimerrén (Veracruz), guajilo (Veracru2), fuailo blanco (Veracrur),nightflowering acacia araaica), pelo de angel (Puebla, Sat Luis Potosi), famarindo de monte (Gustemals),timbrillo (Vera ‘emu, white tamarind (Gustemala, Jamaica). Additional specimens examined. Mexico. cases: 21 km aloo de Ins Logonas do Monto Bell, Cabrera 3600 (MEXU, MO} HL Chocreadero, 5.8 ‘t ‘Ghepa de Cores, Revedlore 17889 (CAS, LL, MICH, MOUS) Mn, of Metin de Mendeza, SW sie of Cerco Mosel, Breedlove 25917 (C85, MENU, MICH, MOL NY) Nipin ot Znacaotin, Pacaje_ Granade, Emughlin’ 2069 (CAS, MICTL, US), Ovando, Mesut 17925 (F, MEXU, NV Escuiala, 1 Triunfo, Metuda 13098 (CAS, MEXU, Ny, Siepec, Caealay Meru 18008 (CAS, MEXU, NY}. toate: Mp. de Tarehiol, 4.5 lm ESE de Tanehinel, Fores 295 (CAS, MEXU, MICH, NY: Rancho Vioj,Joeala, Fisher 16226 (CAS, Fe 13 km al sur de Tepemenac, Greer 591 (MENU Mpio. de Tenango de Dana, 10 koa NE of Tenango d= Doria, Hf Herududes ob Tinres 122(MEXU, MO} Sic Mertlin, I kor de Is densncin « Santa’ Miniea Hi. Herndnie: & Tirres 124 (MEXU, MO}: Mpio de iglksenge, 5 kan NW Temyultengo por carretera a fon, Hecadates & Torres 133 (MEXU, MO} Myo, sds Kesbiconliny 16 kia NW de angus, H. Her tadndae & Torres 120 (MEXU, MO Mpio. Molango, 1 na N de Molaro, Hevndndes & Torres 158 (MERU, MOk Mp. do. Ghapulhiassn, km $ de la frontera SUP Hos If Hernandes «Torres 149 (MEXU, MOR Mpio. Chapulloacsn. 9.6 kan SW de Chopolhmacén por ca. a Javals, 1 Herantes & Torres 150 (MERU, Moy, 29 kn SW Chapulhuacin por carretera a Jecala, He Worndases & Torres 13) (MEXU, MO}, Myo de Tiansustengo, 4 Laval E de Teangutenge, R. Herndn des 3909 (CH, MEXU, WIS), Nk de Tachi, 2 NEXTPREV Au al S de Tne, cia Apalaso, R. Heracles 53:2 (MEXU, Moi de Moyo, Lo, hon Nao Melange,R Herdndss S880 (MENU, NO Mi. de Juien, tacoyota, 15 kas al N do Juies, R. Her tuiades 6105 (MESU, MO he. 301 on highway between heal and Sata Ana, Moore 2680 (GH) camino ence Metepee y Tenango de Dora, entre 16'y 315 ka al NNE de Tuli, Rock 2444 (MEXUD B Cxpue 6 Tanya enangu dl boca 1 Pores cmt ox treeraJacala~Farazanchae, Tenorio & Romeo 2381 (MO) 42 ou. N of Fuoco Ignacio Ito. Dise, N of Yiwopin, Wanderlin ef at 1119 (MO) Mexico” Di Temeaitope. Bejacon, Hinton 207914. F. G MEXU, NY US). outa Pocus, Cervo Espino, Conci 3184 (US) 3 han RW de ls Gals, ear” Pockotls-Oan Seta de Misbunln, Sense of S610 (CAS, EXT) Dats de Jorlataca, a2 ke a SE de Copal, Sousa ot 41. 0929 (MEXU, TEX), Die, de Pula, a 22 km al NE de Pate de Goerrorm, Rouse etal 7088 (MENU: aut 4 Diat, Cuictlan a 9 kat N de Nacatepee, Souse et ti, 10406 (WM, MEXU. MO). Dist de Inga eo SW de Poekto Viejo ys 10 de San Tn Lach, Sus tat 12608 M0} 61 kn aN de Sa Gabel Mist Corr Porto Exconlio-Sia de Vega, Torres & Mort 012 (MO) musitac 7 kee a NE de Husuchinango, Rasurte & Duran 91 (CAS, ENCBE 8 ken al SW de Pate, Gasurto & Dara £07 (CAS, MO}: 8 ae al NE de Honey. lawtto & tard 128 (0M, MERU Myio de Venn Caran, 3 kin al W de Vila Cliissan Hasurin & Tard 386 (HEU, MO 2 hon 1 W de Chonan, Dasara Dard 485 (CAS MEXU, MOy Puente Apcleo, Booge 750. (MEXUY ttrededces de Xicotepec de Juires (vil Joie), Que tero 708 (CAS, MICH, TEX) Zoecopan, Ibarra eal (ENG, MO). sas 11 potost Myo, Huchutan, Ta tej Alone 2060 MENU), 21 i, 8 of Morelos, Kev yer & Chum 400914) teas Tacaniita Nem 4107 (Cli, NY, US) Lan Condon Pringle 175 (8M, DR, F, 6, CH, NEXU. MO, NY, UC, US} Tanasopo Canyon, Pringle 322A, BM, BR, F, ©, GH, MEXU, MO, Ut Mio Xi 3 hr al NE de Ej de Nii, thoes 10506 (MEXU. MICH, TEX® 2 ken a! SE de titel, Reedowsbi 20955 (GAS, TERE Mon: de Valle alederes de Ci. Valles Toren ne (MEU dg hry 85, 74 i N of Tatacinclle, 3.2 wi Sof Malipa Daickenousk ea 500 (ENCE, NY). ie acta aes Fis, 1 on SE Rencho el Galo Hs Hermindes Torres 140 MEXU.MOe Sierra A Contemola, Silvan 421 (ENCB, NY, TEX). Ve item, Moo. Tonayan, Congregocin de ate Cal ‘ede 2ITRIMENLIp Mp Times cereale esto, Calsada 3339(MEXU Myo. Hlpan, Cer de Sn Gist, Calne 622 (WES), Min. Yeruaa, Hrranea A Ceal, Chésero & Guérres 3665 (WIS, Tous 3 Lin el SW do Nico, Dorontes 85 (Fl 2k desputs da Gal HL Reereo, funbo 4 Paka Sola, Dorantes etal 713 (€, MENU: Cero dl lado ete de Bongo, Gaeta {HEX}. Tees ear Caan fhe de oe Reyes, Iahuseba des Heyes, @ Herne eal. 11 ENGB, MEXUy. 68 bas de Xalapa por ears 4 Nace os Horindes & Taree 254 (MENU, MO}, Mp, Nackineo, 1:8 km de Neolco por car. a Misarday 7. Hernanes & Tores 199 (MEXU, MO: Mpo. Rio Bas to, N de Rio Blan, por camino « Tenant, H Her taindes & lores To3 (MENU, MOR Mp. Rio Blanco, ST'Eade ie flesce por easing a Gero el Crt, Tesndnes & Chacin 463 (MENU, MO} ex. 5 E Annals of the Missouri Botanical Garden cf Hastaco slong road to Card, Higgins 2578 \ENCB, Dctih, Npio Taam, EL Choceo. Lara ct al 147 (MEXt, Boies! te Olno, Chaatla, Lunn ot ah 209 (MEXUS Terra Cokrads, ata de Andrés Tenexapa, ‘Medrono 1503 (NEXU Mpa. Chocamin, 8.5 km by oad Bcf Caoeamin, Nee 24257 (MEXUT Al Ede Pn fing & GimesPompo 1008 (GH, San Miguel, 2k al W de Ora, Neslig & Ginnes Pompe 2256 (CAS, F. OH NYE Xitdd Prine 01 Ts GH, TL, MERU, MO, NY, UC, US) evan dal Ro Mataingn, near Ors, Remon 307 (CAS, MEXU, NY; HL Esaulen, even de Xiocepo, Romer 348(CAS, MEXU, US, WIS} I Niet, care: tesa Oriaba-Zongolic, Motes 1460 (A, CAS, F, EXD, US). Spin Zovocolo, 25 km N do Zowosl, Tenorio of al £591 (MERU Mpio. Masala, Dag danas Lica 110 (Py Mio. Nogaes, Nogales, Licea 158 (P, Mio. Chocstan, Chocaman, Liven 192 (Fh pies de Xico,Temimi, Venera 862 (CAS, MICH, WIS} Moi. de Taeulér, Taewlir. Ventura 6857 QEXU} Mpio. do Bardeila, Benderla, Ventura 9949 (Li, MICH Mp. de Nlapa La Mesa de Chivers, Ventera 10534 (MEXU Bipinde Xibtopes, San Morin. Vor sure 19031 (MEXU). Gu aroun, A178 YERAPA? emre Siu Peto Carchéy Sueryo, Molina de Malina 12100 G, LU, N¥p caredera San juan Chanel, Motiva & Malina 22287 (CAS, , LL, NY}, 5 hin N of San Pedro Garcha AS°32N, O913°W), Wiltios oof $2226 (, MICH, US). cmguvets: Montaie Nenojs, 3-3 mi, E ok Canotén, Steyermart 31600 (F, USk Calap, Jie Stow 1093 fF, Pacaya, Johnston, TRIP UF. fecuINTL Fernecen San Vicente Facayas and Calieres, Moline Maine 27068 (OM, F. MICH, US), oTranata: Arvo soba Heyde Lies (GUL K, 0), ws Fol Gants Eig, Harmon 7340 {MO}, huwsverinana: bine Inedn and Finea Sen Rafe), Sera de le Guchuratenes, Stegermark 49470 nia: vey of Jutape, Stan dey 76371 (EUS) gueravrenasco: along old road heveen Finca Prineos and Potulin, Sioniley 6374 {Ep vsty of Sans Marta de Toni Steyermark 34356 [Ey Cotes Finca San Francisco, Sluerh 1049 (A, T. NY, US), sts fox: Goean Jumaytepeque, Hevde & {bus 4403 (G, GH, NY, US). Huson, tas: Men tals Pa, erre Gontin y Caabaros, Molina 1242 (. UTNYSUS):Yorocen- [ein fom Yorucha os Quobrade the Oo to Cnre Bilao, Holes £356 NY). FL SALVADOR, ‘tx suuvaons Velen de Sun Salen, Standley 22958 (Cit, WY, US). santa aves Certo Mone Casio ca 1 tr NE of Metapan slag toa to Punta Trio, Croat 122330 (Os La Majd betneee Hacionsa Momento Gnd Les Planes Molin eat. 15966 (E, NY, US). Nico fate, ESL Las Cruces, Lagan 33 (NOX Lag de Mirallores (3¢16N, B16. Morens 202 (MOK La Goayabie (139138, 86°11) Morena 17545 MOF 1a Cabatin (5°13, 86°13°), Moreno 17569 (MO) snort: Sota Laven, beewren Blaagalps. and Jin toga, Sivens 22562 (MOp Corlilera Cesieal de Nice Tagua E of Tcwtgs, Wilfans et a 24790 (F.C, NY). havacatra: Cerro Apaite en las fuera al B do Baca frp, Teles et 4275 (MENU, MO} Coots, Bes Ecnracs Cervo del Carpitera, Standley 34462 (US batweon Carta ard Tierra ca, Rowlee Sirk 910 QRY, US), Pasa chat above Bayete on ad to {Caco Horgueim Croat 2"029(MO, US! WIS). 3 mi N El Volean, Mewweell 3239 (MO vale ofthe woper Me Charsqt View, Whe 342 (M0, US). Downton Runt saraco: Nevarete, beat 1088 (GH, USK NEXTPREV Volume 76, Number 3 Hemandez, 923 1989 Zapoteca Ciatad Trujill, Alard 13568 (NY, US. a2ues Hepan- synonymy under Z. (Calliandra) portoricensis, fala Vallee San Joan on road to EL Cea, Fle 113376(4, E, GH, US Satigo, Cordless Septensina ae Laginac, et Arroso Herenqul, Fema T0080 (C, Git LE, NY, US). tatavonne El Hoy Freres 1080 C.F, 6 GH, K,NY, USk Boos Chea, Laracire 1023 (NYY Guayacanes, Farastre 1858 (A, NVR San Ce tobak even de Rajnye, Liegier & Jiméoee 6221 (NY) Saute Domingo Ci, tear Lo Tes fame, biowier 182) (NY, US E of Haury, pear Boos de Yuna, Liowier 12253 (GU, NY, US Ina Ioragua, Bisuey, Ceger 2549 (6 MEXU. NY, US 20 mi_W of Santazo, asp’ Pied, Ligier 18268 (NY, USk AO ea. B of ‘Senta Domingo Cry. Coayueanes, Ligier 15682 (NY. USh eae Las Mercede, Pedornaon, Liogior 26678 ( Gi, NY, US} fecal de Cumayass, Poger 19763 (AYR Selle, Bayajibe, Taylor 728 F, NY) Harn, ours DnINORD: St, Mibel do PAtalaye, Leonard 8020 (F, NY. Uc, US) Loogene, Pride 370 (WIS). Teale, Ringste Countan Spee, Briton #28 (NYY Se-Avsew Par Barétey 36307 (CH, Gordon Tom Har) 1316 (US Blue Mountinn Hitchcock sn. (MO). W of Malvern Howard & Procir 13681 (8) S of Cavers, Howard & Proctor 1405 Ak Mount Diatle, Mazon & Kip 161 [AE-GH NY, Us) neas Green River, betwoes Crchona ao Whiteld all, Menon & Kall) 1850 (A, , Gh NY, US} Port Royal Mountains Flamtend, Mazon 9728 (eli, USy Hollymcunt, Moxon J0s04 (NY. US Bala ‘lava, Oreutt 1469 (MO, UC} Troy, Perkins 1347 Gib, Chrstians, Holenwood, Proctor 10203 (4, NY, US St. Ainrew Porch, Craig Hill Port Royal Mountains, Proctor 23808 (A, F, LL, MICH, NY. US Poland Pris Stony River Geage, near Maung River jameson, Proc 28580 (LL), Fusnye Rico. Idand of Coleen Britton & Whevlor BOE, NY. US between Aiton and Ghana, Brion ef 0. 589% (NY} Cale, Bruton & Tisiton 7BENVh Lean Beton eal. 8220 , NY. US), Puente Alloa, Goll #1 ol. 481 (NY, USE neat Laguna de Tariguste, Ligier 10150 (GH, NY. US Ine Fajardo a Cabeza cde San Iu, Sinton 1643 (US yey, Monte Cedeo, Sierents 2090 (F. NY Corsica, tenis 544 (G, CH, Usk Peruelas, Simons 7018 (US Maunabo in Monts Melapascua, Singnis 3234 (G Chl, Laven, sin Pletos, Sinton 021 (G MO, NY USy Yeneo, Carnal a Griggs o34¢NY, US) Vinery Isrants. saisr rina: Bxge/s 210 (GH, NY). 1oxToL Ling Look, Direy 248A Edney’s, Bshlock 41(NY> rou to High Bash, Briton & Shefer 717 (B. NY, US). Chenvarto: USA, ussoumt Saint Lau, Missouri Do tauicel Garden, 1 Herudndes 948 (MEXU, BIO) caL- tron: Sana Bacar, all 8122(UO) Messen. oe MT ‘ook: Cded UniverstariaFacultal de Ciencia, Rice 245 (MENU), wontiors Coersavaca (GH), Beast. Ti fd lanes, Nove Heitrgo, Carran 055 (F, CH, US) Dowavtcan Kertinac. Seute Dorangn, Melo 49 (WY). Fraser, Paris (GH Java. Hoeto Hotanco Bogor Wartarg 2109 (NY). AAs shown above, there are several names that have boon inchued among the ist of synonyms of Z. portoricensis (eg, Benham, 1875, Britton & Rose, 1928). However, F have heen unsile to associate any relevant specimens with these epi thes. Acacia venuste Willd. (Enum, Hort. Berol 1052, 1809) has been traditionally included in Ushich i probably incorrect. The only meterial have been able to relate to this name is «speci at the Willdenow Herbarium (Cat. No. 19143), whieh comes from “Hort. Bot. Berol, W.." as indicated by a handwritten note in the lower right ‘margin. The briet description provided in the ori inal publication of this species ie a roplice of the specimen label, which probably was hundwetten by Willdenow kimcelf, judging by the style of hand ‘writing (Burdet, 1979). This specimen consists of to leaves, sith the pinnae willy separate [roe ‘each other, and eather few pairs of leaflets per pinnse. Because of the lel of flowers or pod, it fecnot possible to relate this specimen to Z. por ‘Zapoteca portoricensis subsp. portorieensis, as olinited here, i= an extremely variable, broadly tdofined subepeciee, the most variable within the species, tex be distinguished from clsely reloted subsp. pubicarpa by its glabrous pods, usually ‘snaller Rowers, glabrous to cihate calyx teeth, and rssally longer peduncles (Table 12), It differs from subsp. flavida by its erect habit, usually smoller fand more numerous leaflets, usually larger Dower parts, and smaller pods (Table 12). Perhaps the most highly variable character in subep. portoricensis is the degree of pubescence of its vegetative structures, Plants range from en= tiely glabrous or glabrate to having branchlts end leaf parts densely villous with Jong tawny hairs ‘This type of variation oceurs at both inter- and intrapopulationel levels, and, consequently. is of little taxonomic value. There are geographical pat tems in the expression of this character. One ex- n Veracruz, and to a lesser extent in Hidalgo and Chiapas, where there isa prevalence Df callections with densely villous branchlets and leaves. The other extreme occurs in San Luis Po tos, Tamaulipas, and Ganteal Amoriea, where gla- brows of sparsely villous loaves and stems predor inate, fa the West Indian plants, the leaflets are ‘consistently glabrous, but the branchlets may be villous or glabrous Other variable characters in Z. portoricensts subsp. portoricensis are the sige and number of poirs of leaite por pinna. Most plants from the Sierra Minteca, Sierra Madre del Sor, and Sierra Madre de Chispas, in the states of Oaxaca and Chiapas, Mexico, are characterized by pinnse with smaller, more numerous (10 67) pairs of leaflets ‘usually densely distributed along the rach, giving. the pine a feathery aspeet. Plants with small, muiterous pairs of lealets per pina aro also found in Central Arseriea. In Oaxaca. plants with leaves NEXTPREV 824 Annals of the ‘Missouri Botanical Garden of thie sort also tend to have larger lowers and usually larger stipules. However, these characters have bees found in oumerous clleetions From dif. ferent regions (es, Veracruz, Pucbla, Hidalgo), swhete they necar in plants that do not have feath: fry leaves like the ones in Oaxaca, eo that thors ia no definite geographical pattern and no bass for recogniving additional infraspeciic taxa. In eon tasty many of the plants from the West Indiew Ihave larger and less sumerous leallets, thereby resembling subsp. flavida ‘The vast majority of collections of Z. portort consis subsp. portorteensis have trate branchlets; however, plants with four-angled branchlets, a ‘charaoteristic diagnostic for Z. terrazona, are wot ‘uncommon in some regions. Som of these plants ay be the result of hybiization between subsp. portoricensis andl Z. trtragona, These two taxa fare genenlly ecologically igclated in Contral Amer: fea and southern Mexico, whore their gcographical ranges overlap. Ia these regions, populatians of subsp. portoricensis are found primarily in mon ‘ane cloud forest or similar habits, whereas those ff Z, terragone oecur in warner lonkaod habitats Populations with four-sngled branchlets are mast frequent in Guatemala, where these two taxa eome into clase proximity andin sore cazes ate probably stcict sympatric. A numbor of Guatemalan collec sions of subep. portoricensis have four-angled Lorarchlets,althoual som is characteristic is expressed only distally, or i less marked than in typical Z tetragona (e.g., Molina & Molina 12287, Stondiey 80574, 87132, Steyermark 34250, Willians et ai. 12100. and others). Sig nificantly. these plants are derived from areas where Z. portoricensis subsp. portoricensic and Z, tetra ‘gona approach each other, as for example, in the areas of San Juan Chasneleo-Cobin-San Pedro Corchi, Departamento de Altx Verapss: Colomha Santa Maria de Josie, Departomento de Quetzal tenango: IpalaComatan, Departamento. de Chi- (quirnuls; ond Fseuintla San Vicente Pacayaa. Departament de Kscuinta. Tadividuals with four tangled branchlets also occur in some other regions [eg., Oriashe-Cordobs, Verserus, Chiapas), snd feven in areas where Z. Zetragone does not oceur, sch as Hit ‘West Indian poptlatons often diffor from those fon the mainland in some charactors, possibly the reault of the emaller gene pools in insular popu lations and the selective effets of local climates, In many of the plants collected in the West Indies, the leaflets are comparatively larger, approxirnat- ing those of Z. portoricenss subsp. flavin, Ja- rmaican specimens are characterized by smaller (4.5-6 eu long) pods and by woually smaller, black seeds with cegularplerograms, as compared with the mainland populations. In addition, some of the plants from this sland have 1-2 jugote pinnae with fewer pairs of leat. Zopoteca portoricensis subsp. portoricensis and its close relative Z. caracasuna subsp. caracasena cooceur on Hispaniola, although it is unclear ssbether they realy grow together. Hore, as else- ‘shece in the West Indies, most plants of subsp. portoricenss dilfer [rom mainland populations 2 hhaving ealycos with the tecth lunecolate to linear lanceolate, a charaeteritic of Z. caracasana. In some ease the identification of herbariura speci: ‘mene is (roublesome if information concerning habit and filament color is not pravides, However, there are several diagnostic characters for each taxon (Cable 13, while comparing the two species, applies to the two subspecias discussed here). Zapoteca caracasana subsp. caracasana differs trom Z ‘portoricensis subsp, porioricensis in its prostrate’ Soandent habit, smaller flowers, white/red-purple filaments, and hy the feequent oecurrence of pe opaniculate inflorescences with more slender pe unos. The relationshiys between these two taxa fare discussed further under Z. caracasana. 6b. Zapotcea portoricensis subsp. pubicarpa H, Hern.. subsp. nov. ret: Mexico. Oaxaca: Dist. Husjuapan de Leda, a 6 km NW de San Jersnime Silscayoapilla, casi en la desvieid 2 Ciénega Zahuatln, veg. allerada ce Bursera om Lysiloma, 1,900 m, 4 Des, 1978, Sousa & Zarate 9822 (holotype, MEXU; iotype, cas). Pinae 2-4)oe; fle 19-20; npn utegue angutoadonge tl lanembte, Calpe A039) san fun dod alc cee ite vila; cena 7-410) fim lang spe pice as. Lopuine paberl Breet shrubs, to 3 m tall branehlets villous 10 densely villows with white.gray hairs. Pinnae 2. jupates leaflets 15-301 pais por pinma, narrowly bhlong tolancenlate, oblique at base, acute at apex. trucronalt, the median ones 4-7 x 1-2 mm, ciliate, raraly sparsely vilous: stipules nareowly triangular, 11 mm long, usually densely vllows Capitula axillary; peduneles costae, 1-3-8.54-6) fem long at anthesis, glabrous to densely villous Calyx 2-8(-8.5) nm long, the teeth triangular, the area of the teeth villous to densely villous: corolla 7-8-10) mm long. usually vilows apically: ‘ovary glabrous, Pods ca. 6-7 om long, brown, puberulen. NEXTPREV Volume 76, Number £ 1989 Hornindez Zapotec 825 Distribution Fig. 36). Endemic oa woaticted area in southera Puehla and northern Oaxaca, ‘whore it has boon collected in disturbed places in the transition zone between oak and tropical de cduous forests ard in thorn forests. Collections fcome from 1,850-2,120m, Plants of this subs species grow in a generally drier cnvigonment than ‘other rnainland populations of the specie. Aialtionol specimens esomined, Mexico. reir 259 kn Sof Menica Cy, Berea. (2.709 (MERU vicky of Sun Lab, Darasea de Tacuosto, Purpur sn (IMF, CHL MO, NY, UC, USs Carve de Yerba, San Ls Tulsa, Sioms Mintoea, Parpus 2658(, UOr 14, mi. SW of Avotepee, Senior et al. #646 (US) 4.7 hm al SW de Acairper, Sousa 7780 (MO), Mpio ‘& Zapesili de las Sain, & 8 kn al SW de Artec Sousa « Rico [0322 (MO) Fl Coro, 10 km NW te Caltepee,o ra SE de Acazapes, Tenorio 6859(MEXU}, Mpie.Caltepee. Riot de a Mrbe, Masa Chica al W de Cahepoe, Tenors & Romero 7504 (MEXU, Zapotecs portoricensis subsp. pubicerpa is closely related to subsp. portoricenss, from which it can bo distinguished primarily by having puber. tent pods, villous calyx teeth, larger overs, an usually shorter peduncles (oe Table 12). The ova ‘ies are glsheous, hut pubescence becomes event soon aftr the pods start to develop. The charac lerstiee used to define this cubspecies are not di contimious wth those of subsp. portoricenis However, the particular combination of morpho logical characters defining sulap. pubiearpa is ds Aipeive and. appears to be relatively constant Uhroughout its range. This ac ite biageogeaphic conaisteney jlify its secopption an ep suhepecies The pubescence of the pads i the main dig nostic charactor for thie subspecns. Nevertheless, tairy pode have boen found in two colctons of Z. portoricensissulmp. portoricensis fom Nica pa (Laguna 333, Moreno 8272),"Thes pants, However, have sualler flowers and longer peduncles than ote characterise of sulep. pcbicarpa they cleanly fll within the hits of sibep. portricensts Flower size is variable in subsp. portoricenss, wth largo (calyx ~ 2-3 mm long. coralla ~ 6-8 mm long) overs found ia plants throughout the range of this subspecies; such flowers, however, are rarely as large as thowe of Z.portoricensis subnp. publ arpa si the plants on which they occur never have the hairy pods or short pedicles character ine of subsp. pubicarpa Ge. Zapoteca portorieensis subsp. david (Urban) H. Hern, comb, et stat. nov. Cat Giandra flavida Urben, Ark, Bot, 24A: 4. 1981; Britton & Kilhp, Amn. New York Aced. Sei, 35: 137, 1986. TvP#: Grenada, ad Bel: vedere, 1,600 ft, Dec. 1889, agers 6226 ectotype, US 938738, here designated is slectotype, GI), Seandent shrubs, 2-3-7) mm tal stems slender branchlets terete, usually spreading and descend ing, elabrous to sparsely villous, sometimes densely villous in the youngor parts. Pinngo (2-)8- 5-7} jugatej leafles 8-16(-36) pairs per pinna, the me ddan ones obloug to oblong-obovate, sometimes ssubfaleate, oblique at buse, rounded 10 acute at eronulate, 7-20 % 2-6 mm, finely villous ‘on both surfaces: stipules trangular-anceoate, oc casionally lineae-triangular, to 11-13) mam long, labrowe, ciate, Capitula commonly axillary, oo casionally in psouopanicles to 20 em long: pe dancles slender, 3~7.5 er9 long at anthesis, Calyx 1-2 mm long, glabrous, the teeth tiangular to deltate, carely narrowly oblong, acute, ceasioally inregular; corolla 3-4 rama long. Pes 0 16.5 % J.2 cm at maturity, thickly membranous, usvally black ot maturity. Seeds ovoid to very widely ovoid, 2.6 = 4-5 mm, brown to dark brown, the urcola usually motled, pleurogram irregular. Somatic chromosome nutnber 2n = 26, apex mm Distribution (Pig. 36). Zapotece portorivene sis subsp. lavida primarily oecars in lowland trop ical forests, where its commonly found along river hanks, in inundatod orcas, and in Yt thickets Tt range extends from coutheastern Veracraz and northern Oaxaca. Mexico to Venezuela and Pera, ‘where it has heen collected in several disianes pop ‘ulations. Additonal eolleotions have heen mace in Grenada (Lesser Antilles), Guyana. inthe states of Para, Cears, Permambuca, Goiis, Malo Grosso, Trail, aod in the department of Senta Cruz, eo livia, In Mexico and Gentral America itis primarily ‘a lawsland subspecies. extonding ahitudinally from 0:0 670 melevation in Venszuela it reaches 1,100 rm. This subspecies has been introduced to Afriea, ‘where it has been collected froin naturalized pop lations in Chana (Ertl 275, Vigne 4011) and from cultivated individuals in Nigeria (Guile 702) Additional specimens examined. Mexico, cusps Mpls. Palenque, « 0 om W dal onzonque Palenque, FH Hernande= & Torres 805 (MERU, NOs Borah Hanser 282 (CH, MO, US Mp, Oceeingor 3 kis 8 Fromtors Fcheveria, Hlartines 9120 (MEXU Molo cocingo, 2 an W Cruces Corsa, camino Pal Toco Lacsntan. Mert'aes 10136 (MEXUk Mpio. Ovo congo, 4m $ Bido Beneménito de ae Arserea% camino Flor de Creso. Martinee 10/79 (MERU: Mp. de Paleo 2 8 kn al W de Cataaj Sona ef a 2277 (BM MEK, Mn, de Ocosrgo, ea Booth, Valdicin NEXTPREV 826 2201 (KAL). ovxsess Chitopes, Martin Tantepee, Chieges. Mvtnes Calderon (A, Lh ETC, TS) aansco. Pactaljes Rib Taco, Conta 2731 (LL, MERU Mato, Reviroca 052 0X US: Hull, San Mig, 1 im al W oe Cerone, Souce et al 12208 (BM, MEXUL Verscntz: Chaca. font, La Iota, pre sn, AL Mpo Caton avn, Fb Coxcoapa que dewnbocs en Laguna Sontecomap at BT Heonénate: & Torres 793 (MEXU, MO}, Miser, “Mei! im (NY, US Laguna do Sonsscomepen, seb kz ‘here del Rio Caseagpan, Catemaco, Menende: 157 MEXU, MO) Giiestata. atta vonca2 Lagan Spel, ai. SW of Sct, Steyermark 44420 (Fy. Calduie Tuarekheim 7737 (GH. K. MEXU, NY, US). 124as Jocala, H. Joksson 1075 (F, VSe boween Dartaoath tad Moraes towando Logo Uebel, Moneana del Mico, Steyermark 39108 (F, Gh Lago label, opponte Sar Felipe, betwen Son Fale ara mou cf Ro Ja View, Stevehmark 39685 (Fost: Lacan, on El Carbs boat 6 Lan SSW, Contreras 3937 (F, Lly MEXU edo, Cadkense Road Controras 9480 (F, Ll. MENU}. minute Retaliein Berni Corio 1252 1k). int: Mobo River, Peck 752 (GM), Wio Grane, Seape 1200(A, BMP, C,H, K, MICH MO, NY, UC}, Jecate Groak, Setupp 1216 (A, BM, FG, GH, MICH, MO, N¥, Lich Heenrmas sara nama’ Linens, Standley & Lindetie 7540), Neeanaen 414Y 6 Coro La Fa (3°40 1O°N, 84°56°2 5°), Ons 1131 MO) Pads lias vaste Rio Are, Due & Miarson 12706 (MO), {Yostoonns, ons aactnor sess of Moron end Tome he to mouth of Rin Cay, Seyermarl 12702 (Fy MICH, NY). caRanei: between Puerto Cael and Sat Felipe, Pitter 153 (GH) vanact¥ Loe Caan, Pier 3759 (CH, NY, US) Couoseis, coeds Rio Teuanss, Itwcen Ri Slo and La Narva, uke 9825 (NY). a Store de Lb Macatevs. Plea Boni, Pllspsoa oof Tait (US sueranoen: Care, andré 204 (hGH, K. NY, US) Ecusoor. cuayac: Caper, betveoe Cuay al ai Daule, Doduon & Gentry 12579 (MO). Le RLS Between Qhevede an Santa Domingo, Dadson & Gentry 5295 (MO). Pom MUten: Pachites, Schunbe 2571 , GUS) sine Ror Paveattamnbe Valles Kilip ‘Stull 25272 NY, US Puerto Yoon, Killip & Smith, 26220 (NY, Usy Paste Termidr. Kilip & Sn, ‘eas (NY. US). Lon Yariniogins, C26 8729 (Fete, Ki sex nme Chava, io Hulaga, King $015 (i (AS. F GH, MO, NY" UC, U5) Tian Ji, Alto Bis Hiuiaga, King 4298 (A, BM, F, MO, NY, UC, US) Morea! Céotres, Tocacho Nusvo, Seats 4798 (CAS, "GH, MICH, MO. NY 4 Paco Guados 37808) dds Tin, Hatchback 34270 (NY, Sera do Cas 30 in Sof Capris, nen eval. 17903 NY) nacocons: Arps, Projet in, Sia & Roadrio 4841 (8) ns ade Baterre, Duche 1517 (BM. US) Mente Slee. Florin Haha. Duce T6031 (BM, lis Monte Nk ge, Serra di Lay Pri 30205 (US), rensauve, ‘eptra, Berto 242048, CH, MICH, US, WIS Visenia, Mate do Hag. Jundis, Pedra da Nogra, Tavares 2S4NY oust sits e402, Prov. Teli, Bae Vac Seine frac 199 CH, MO, NV, US, WIS Proy- Del Sart, Geen del Sur, Cat fkenasisty Seinech 2456 a. ‘This subspecies can be easily distinguished from subsp. portorieensis and pushiewrpa by itsscandent Annals of the Missouri Botanical Garden habit larger, sly Jess merous Teall, snd vival Targer poss. In addition, subap. avid primarily groves in habitats aseociated seth lownd tropical forests, whereas the tno other subspecies have been collected chiefly i aubtropical, mid: to relatively high-altitude habitats. Zapotece porta: ricensis subs. flasda can be distinguished fom Z caracasina by baviow larger leas, Sower tus contanly glarous, dltate-wiangulr calyx teeth, white flames, and lrger pods, The capi tla are arranged in pewlopanicles only accesion- ally in Z. portorivensis sikep. flavidsy this is a very common condition in Z. caracasora Zopotecn portoricensis subsp. lavidi is rele tively constant morphologically throughout its wide range; however, a number of collections from South ‘America havo « combination of charactors that fer from those ofthe Mexican and Central Ame: iean plants, Characteristics of a collection from Colombia (Philipson et al. 1418) suggest mor phological intergradation between Z, portoricensis subsp. flavide and Z caracesana: narcovly tei= angular to linear-triangular supules, leaflets 8 2 ram, nmmerons (to 36) pairs of leaflets per pina, capitila acranged in elongate peeudopanicles aly with linear-lanceolate teeth, and white filaments ‘An slimost identice] situation is dieplayed by moet of tho Bradlian ellotions af Z. portoricensis sokep. favida (Andrede-Liww 70.5777, Ducke 1517, 16051, Frées 30205, Guedes 378, Hasschbach 34270, Pickel 2130, Tavares 751). Here. several of the vegetative and reproductive characters di agnostic of either Z, porvoricensis subsp, flavida. ox, carecasona itera in pric po Although it is not Known whether Z. portori consis subnp. flavidte and Z. earacasane ever 26+ tually grow together in specific habitats, they have been collected in comparable mierohabitats (eg. raver banks, shady sites, and thickets), which raises the possibilty of hybridization. Populations of Z. caracazana, however, cocur sore commonly in rier habitats than do those of Z. portoricensis subsp. flavida. ‘The distributions of these two taxa overlap only in the area of the Andes, where there is much loeal miceohsbitat differentiation Callections from Bolivia have soane characters that io not correspond with those commonly found in individuals of Z. portoricensts subsp. flacide. Forinstaner, the filaments are reported to he whit pink, instoad of white,in Steinbach #22. The eslyx in almost all the Bolivian eollesons has lanceolate teeth (Steinbach 5412, Weddell 4237, Williams 267). Tn addition, the mature pods are always smaller than typical for this entity (to 10 em Tong) NEXTPREV Volume 78, Number 3 Heméndez 827 1989 Zapotecs Fromme 3 and the valves are thicker (este Pierce sus, Troll 1738, Williams 244), 7. Zapoteca mollis (Standley) H. Hern.. Ann Miszouri Bot. Gard. 73: 755-763. 1986. Cal Yaniira mollis Standley, Conte. US. Nat. Herb, 17: 431, 191d; Stasdley, Publ. eld Mus. Nat. Hist Bot. Ser. 18: 492. 1937, Annestia rvllis (Sasiley) Briton & Rose IN. Amer. F123: 65. 1928. TyPP: Costa Rie: boissons Nicoya, Jun. 1900, Tondie 13536 holotype, US no. 578114: iorypes, BM (2), F, 6 @), CH, K (2), LE, NY @), US (2), et shrubs to 3 tate under magnifieation, abundantly villous with tawny hairs. Pinnae (3-)4-Fjagates petioles eelin- lar, terete, with a conspicuous channel in tho adaxial surface, 1.2-1.5 em long, abundantly sl lous; rachis $.5-9.5 om long, abindantly villous achillae 4.5-12.5 cm log in welldeveloped leaves; leaflets 15-26 pairs per pina, narrowly oblong, oblique at base, acute to rounded and mucromlate at apos, the median leafiot: 7-18 x 2-5 mm, the proximal onos smaller, all membranous, drying a light green in the abasial surface, demvely pilose wth white hairs on both surfisees; leaflet venation inconspicuous; stipules leafy, triangular-lancealate, acute, €4, 6-14 mm long, parallel-veine, pilose, ciliate. Capitula in ciaple. rarely eampound, pew Adopanisles 3.5-15 era longs peduncles waualy fa= ciculate, rarely solitary, 2.5- em long at anthesis, usually densely villous with tawny hairs, Calyx ea. ram long, glabrous, the teeth lanceolate, seule, fea. 2 mea lang; corolla infondibuliform, ca. 3-4 tall; branchlets tevete, cos Disribiion of Zepatica mals (ts) and Z. coataricenss (rings), Cots Rien aod Pana, ‘nm long, glabrous, the lobes ovateJenceolate, seuter Slamments about 2.5 em long, the basal half white ang the distal half oehrcleucous, the staminl be cca. 2 rum longs ovary ca, 2-3 mi long. when fertile, glabrous, sometimes with fine translucent hairy sessile, Pods to 9 * 0.7 em hen mature, with the apex rounded, rarely truncate, rostellae, thickly membranous, densely elous with long tay hairs. Seeds widely rhomboid, fattened, ea. 4 3 mun, dark colored, the areols lighter, pleurogeaim ievegulae. Distribution (Fig. 87). This species has heen collected im tropical dry forests near Nicoya (Pen insula of Nicoya), in San Lucas Island (Gulf of Nicoya), and in Rfo Lagarw, all in Prov. Puntare nas, Casta Rica. Plants from a presumably disjunct population have been collected i Panama Caty, Panama. Gusta Ries. Additional specimens examined, PUNTARENA: Idand of San Lucas, Gull of Nicaya, Bar toy 2750 (BM, MO, US): Ile de Son Lees, Coll de Nicoye, slong Quebrada Hacienda. Vieja. 957'N, SaPAW, Groyuin er ab £708 (MEXU, MO) ble de San Lucas, Golfo da Nioya, between Playa Tunbabots aia Passa Bazeaona, Grayam 4201 (MEXU, MO® bl 4eSan Lica, Golfo de Nonya, along tale Pim Girl, Grayam 4290 (MEXU, MO} Rie Lager, extronque a Monteverde y Cnucimal, Teles 4307 (MERU). Passen Sta Cru, Aorayn 7 (MO) ‘This species clsoly resembles Z. portoriconsis subsp. davida in general characteristics of leavo= nd inflorescences, but itcan be distinguished reodily hy having an eneet habit, which eomtrasts with the scandent habit of the subspecies, and by being villous or pilose throughout, including the pods, NEXTPREV 828 Annals of the uti Botanical Garden which are densely covered with long tawny hairs. Further, the filaments are white in Z. portoricensis subsp. flavida, bat white in the basal half and relirolencous inthe distal halfim Z. metlis(Grayum 4162, 4290)-(It would be desirable to chock more living individuals for thie feature.) Contrasted with 2. portoricensis subsp. flava, whieh is commonly found in thickets in wet lowland habitats (5. along, water courses and arse pords), populations of Z. rollisusually oceurin relatively dry habitats. With the exceptions of Z. formosa subsp. saluadorensis, Z. caracasane swap. weberbauert, and Z. por toviconsis subsp. pubiearpa, this isthe only species in the genus with hairy pods. In addition to tho few collections of this species from Costa Riea, there is & collection from Santa Cruz, near Panama Cay (trroyo 37), which ine dicates a disjunetion in its eange. This collection, with immature pede, matches the plants collected Costa Riea well, Recent attempts (1984) t9 20 collec from this population have been unsuecess ful, prosumably due to widespread destruction of natural habitats inthis part of Panama, 8, Zapotwen earacasana (Jacq,) H. Hern. Ann Missouri Bot, Gard. 73: 755-763. 1986. Me ‘mova earacasana Jacq. Colloctanea 42 1.42 1791. Acacia caracasana (Jacq) Willd. Spee. 4: 1068. 1806; DC, Prod. 2: 467. 1825: G. Don, Gen Hist. 2° 419. 1882. Cal. Bandra caracasana (Jacq.) Benth., London J. Bot. $: 99. 18444; Bouth,, Trans. Linn. Soe. Landon 30: 543. 1875; Briton & Killip, Ana, New York Acad. Sei. 35: 137. 1936; J. F. Macbr., Fieldiana, Bot. 13: 69. 1943: Wood. son & Schery, Ann. Missouri Bot. Gard. 3 2601. 1950: L. Cérdenas, Revista Fac. Agron. (Maracay) 7: 181. 1974, Feuilleve caraca sana (Jaeq,) Kuntze, Revis. Gen. Pl. 184. 1891. nmeslia carueasana (Jacq.) Brision & Roe, N, Amer. Fl. 23: 65. 1928. TPE Facges Keon, PL Rar. 3: 20, 4, 632. 1793, (Gwolype lore designated, capies at MO, NY, U), Thave designated this excallontilustration as nootypo (Fig. 38) hocauso no other element ‘on which the description of Mimosa caraca ‘sana was based eppasrs (0 have survived ‘Seamdent, subprostrate or prostrate shrubs; stems usually branching shortly above soil level, ca. 2 fem diam; branchlets flexible, elongate, usually to '3C-5) m long, terete, sometimes angulate, costae. slabrous to densely villous. Pinnse (1-}2-7-jugate: leaf primordia usually densely villous or pilose petioles glandular, terote, with a conspicuous ‘channel adaxially, 0.9-5 cm long, usually villous ‘or pilose; rachis (0.4-)1-7.5 cm long; racillae 2 66-75) em long; leaflets 10-26(-37) pairs per one, narsonly oblong, oblique at base, rounded to noute at apex, the median ones (1-15 LOC11) %()LS-2.5(-8)mm, the proximal ones salle, all membranous, glabrous 10 short-villous, usually ciliate: leaflet venation usually conspicuous alax ially the primary aud secondary veins prominent stipules leafy, narrowly triangular to linear. tla, sometimes subauriculate a hase, acum parallel veined, to 7(-12) mm long, glabrous to villous, ssvally kate, Capitola axillary or in simple ‘or, lesa froquently. compound pseudopanieles to 20(-26) om long: peduncles fasciculate at distant nodes, 1.5-6.5 em long at anthesis. Flowers pe samerous, sometimes hesamerous: calyx cup: shaped, deeply divided tothe base, 1.52 menlong, altbrows to deasely covered vith relatively long wavy hits in bud, glabrous, villous or ciate at anthesis, the teeth lanceolate to linear Taneoolates corolla campenulate, 3-4 mm long, glabrous, the Jobes lanceolate: fiaiments ©2. 2.5 em long, white in the bacal half, pueple-red ia the distal half, the staminal tube ea, 1.5 sam longs ovary ea. Um loug when fertile, glabrous of necasionally wih a few hyaline hairs, shortly stpitate. Pods straight, sometimes cghtly curved, apex usually eurvedly fostrate, LO(-11) % 0.7(-0.8) em, thiekly mom branous, glabrous, glabrate, or caneccent, becom: ing densely short-pubescent, Seeds ovoid to shom boid-ovoid, 4-6 % 3A mm, mowed oF browns plourogramn irsegular. Somatic chromosome num ber 2n = 26. Distribution (Big. 29). Populations of thie species occur primarily in disturbed habitats de- rived frvm a wile variety of vegetation types, rang. ing from dry serubby vegetation to wet evergreen {eoroste, [ts range extends from northern Venezuela (including Isla Margarita) southwest Colombia, Eruador, and Peru, where it has been collected in scattered localities. I has also been found in His ppaniols and ia dist dduras, Perhaps this i sist population in Hons the speries within Zapereca Fico 38, Minow earacasane” 3: 20,4632. 1798. (- Zapotecacaracesana sep. caracasana) (rom Jecgun's Fon, Pl: Rat NEXTPREV Volume 76, Number 3 1989 Hernandez ZapotecaPREV Frown 39. Annals of the Missoun Botanical Garden Disrtion of Zapoteea caracasana subnp.earacnsana (ts, 2. earacasana subsp. weberbaueri (tra) and 2. carncasana of unkrown sulmpece ants open sat). Jations occupy the wiles! altitdinal 10 m. Woodson & Schery (1950) incorrectly included this species in the Flora of Panama lated to Z. portoticensis, and these twa species have been merged at times. For instance, Bentham (1844) considered Z. (Calliandra) caracasana as 4 syn ‘onvin of C. portoriconsis, although subsequent, in his cevision of Mimosoideae (Bentham, 1873)” he recognized them ax distinct species. Later, Woodson & Schery (1950) treated Calliandra portoricensis as synonym of C. caracasaza, The ‘ost important characters distinguishing these 10 species are outlinad in Table 13. Although iden tification of hesbarium specimens is admittedly dif fieult in some cases, Z. caracazana can be distin guished from Z. portoricens by ita presumably ‘coustant scandent or prostrate habit, ually small: cr leaflets, and capitula with more slender pedun les, whicly are usually arranged in pseudopsni. ccalate inflorescences, a condition rarely found in Z. portoricensis. Observations on evitivated ma. ‘erfal and information from herbarium specimens suggest that the seandent/ prostrate habit of Z. Zapotece earacasana ix closely caracasina & widespread throughout its ranges however, its uncertain whether itis characteristic ofall the populations or if eect plants occasionally ‘occur, a2 indicated by labo! information om a fest specimens. Within Z. portoricensis, tho seandeat habit oceuts only in subsp. avila, There are, portant differences in growth habit be tween that entity avd Z, carecasama: cultivated reeimens of of Z. caracasana (H. Hernando: 961) had extremely short primary stome, uruel siving rise to drooping, very elongate branchlets 10 em above the soil level, while those of Z. however, portoricensis subsp. flavida develop very long, {exible primary stems, sometimes resehing several meters before giving rise to secendary branches. ‘The Bowers of Z caracasana are generally smaller than those of Z, portoricensis, and the calyces are consistently decply divided tothe base, with the teeth lanceolate to lineardanceslate. In addition, the two species are casily distinguished by thei filament colors (Table 13) and by the usualy smaller pods in Z. earacasana, Pod size, however, is highly variable in Z. portoricensis, Although the degre of pubescence of stems an lcavea is extremely variable in Z. caracistiaa and in Z. porioricensis, the branchlets of tho former NEXTPREV Volume 76, Number 3 1989 Taos 13. Heréndoz: Zapoteca 831 Comparion of Zope caracasene with 7. porervensic Z. caracasane Z. ponoricenss Bate ‘eandent, prosaate Tait 1 Samm Gopi onary arranged in pemdopaniles, oF volary aly Aseply eve wich the tut lareecate to sear lancenate Corll engi S-1mn Fisnent color wwite‘redpurgle M en cn are never as densely villous as commonly in the latter. Similarly, the indumentum of the calyx is Highly variable in Z. eerecasana, most of the cok. Tections with completely elabrous calyess being concentrated in Ecuador, However, the peculiar vestitute observed in the ler buds of ths specias has never heen: observed in Z. porioriconsis, Bar. ther discussion of the relationships betwers these neo species is under Z- portoriensis. Zapoteca caracasanc consists of two closely related subspecies: earacasana (with glabrous pods) and weberbaveri (with eaneseent or pubescent pods). As io the other taxa of Zarpoteca with pr escent pods (Z. mollis, Z. portaricensis subsp. pubicarpa, Z, formosa subsp, saleadorensis). the ‘ovaries of Z. caracasana subsp. weberbaueri are flabroce, and che haire of the pode presumebly develop after f ‘his subspecies occasionally have a few hrs. tization. However, the ovaries in te, Pose slabrous or nesay 20 - és 1th, Badr sanascont or pubsocort 1b, eulap. vcberbaeri ibep. earacasana 8a. Zapoteca caracasana (Javy.) Ho Her. subsp. earacanana. Branchlets villous to glabrate, usually glabrate, rarely densely villous, Calyx densely covered with usually wavy, long sins ot glabrous in bud, the tenth usually ciate at anthesis; avary glabrous Mature pods to 10{- U1) ¥ 0.7(-0.8) em, glabrous, rarely elabrate, Sends 5-6 % 4 mm, Diseribution (Fig. 39). Relatively common slong roudedes, in cultivated fields and thickets, smd in shady ston. I Hispanials. On the mainland, it has been collec: in northern Venezuela (inchiding Isla Margarita), ina fow presumahly disjunct localities in the de. the West Ines it occurs on Fearon, eost 20% 16mm tesally axillary, rately in pamadopsn ‘cee teeth trang of date, rarely La aloe to linear lanceolate 3-10) site partments of Magdalena and Mets, Colombia, and the Department of Ayacucho, Per. Zaporeca ca racasana sulup. caracasnne appears to be alla patric with Z. caracasana subsp. weberbaueri, although their ranges come into proximity in Co: lombia and Peru, Collections come from altitdes ‘of 200-1,800(-2,500) m. This subspecies hes also Trees eollocted from cultivated individuals at the Botanical Garden of Para, Brazil (Huber 2078) Given the fact that the only collection from Ho: dduras lacks pods, it ie not possible to assign it to a subspecies, Common name lie, Caja (Dominican Repub Addltinalspocimens examined. Hart: Massif des (aos, Lee Gonaives, Ehman 9081 (GH, &, MO. NY, S US) Dexnucas ReruBtic. Movre cust! Sarre Domingo, Contlera Central, Moers, Ekman 12012 (8). 420 io del Meso, Elman 13723 (GH. 8, US). aanionoco: letween Fl Agasente ad Saponin, Liogier 10782 (F NY} suntacts Hato dil Yar, Vale del Cle, Eka As Fy C,GH,$, US); Corrode a Caleb, Jingne= ‘Sto (US) Cortana, B Chor, Linger 12230 (NY rear Lae Meroedos, Podornals,‘Liegier 16087 (NYE. fear Sapo Liegior 18190 NY}, VENERLELA, ARACCA Coren Wiliams 20415 (B, US), crstanio Tend: Gentry & Morila 10326 (MOk near Caracas, 107(G, CHP. US): Lon, Tamayo OC, US} between Sabana Grande and Barta, SrtA, Wiliams 10592 P, US) wasps: eae Bain BSN, WPs0'W), Brevelor 3149(M0, NY. 3p Tyevpore Fore Rowrve (15'R, 70415°W), Brotlor 2445 (US). aut snare B Vall, Miler Johnston 20 4F) romTucutse. Ghanare, 17 hm NW of Tacupiie, Devidee ot al 21420 (MO). Couch. ete: Vilovcenso, Cuatresnsas 4001 LFS) Sierra dele Merarenss Ii Guaguyes Phin ff al. 1826 {fy US) Macbatiuss Manure, Seer Peis ahi, Caatrensune & Casters 25077 US) inv, sracucnor Earl, bewnen Harts sn Bio Apa imac, Kilip 4 Smith 23007 (NY. US), HoNths UnPin(?e Quebrada Nf Copin, Poole & Waison 1034 (ub. Bb. Zapoteca earacasana subsp. weber baueri (Harms) H. Hem. comb. et stat. novPREV 932 Calliandra weberbawert Haroy Repert. Spe. Now. Rega Veg. 17: 88, 1921. rvve: Pert, Huancavélics: Prov. Tayaeaja, Valley of the Mantaro, orthosst of Pampas, 1,600 an, Mar. 1918, Weborhauer 651 (leetotype, GH, her designated: isolectotypes, F, (fragm.), S. US). Branchlets densely villous to usually glbeate Calyx very densely covered with usually wavy lang Innis, or glabrous in band, the teeth slate ow gl Dbroms at anthesis ovary slab «fo, translucent hairs. Mature pode to 10 * 0.7 usually eaneseent when young, densely ahort- pubescent at maturity. Seeds 4-5 > 3-3 Somatic chromosome aumber 20 = 26, Distribution (ig. 29). "This subspecies occurs primarily in relatively dry, scrubby vegetation. It has boon colloetod from scattored locates in Co- Jombia, in the departments of Antioquia, Cauca, Cupdinamares, Santander, Tolima, and Valles the provinces of Chimborazo, Guayas land Loja: and in Peru, i the departments of Apts rimae, Cuzco, Huancavélica, Lambayeque, snd Turbos. Callections come from elevations of 2,500 m. Addivional specimens examined. Couns, swoguia: veiity of Medalin, Toro 787 (NY. US) [GUNDINASIARAYelededores de Pueate de Sea Ato de Tena, Cuatrocasas S284 (P, US) Lx Esper, rede 4674 US) Sunanderio, ride da? (IS), satan Rio Surat valley, between Fl aboneily nt Urat, Kip Snich 16853 (CH, NY, US bervoun Bl Rollo end Tonay Klip & Smith 19392 (A, CH. NY. US). TOU Chicora, Haugh 042 (US), VALLE 30-35 kin Es Pat rico, Johnson 249442 (MO). Beton einen leimeen Hulgea ara Neranjupata, Asplund 15508 (, TL, NY, S) canyon of the Ro Clanchas, near Huara, Comp E3107 (F, GH, MO, NY, US} Veriur, Rose & Rose 23517 (CH, NY, US). cl eyss: between Crongin an Ceres, Asplund 15985 (B, NY, Sk Coayaga, Asplund [5982 0. Go ML NY, Ps, US) he 19 Wl Pangaals on Gaya Diets e ul. 19590 (MOy, Capers, kn 2 e Das, Dedson & Genary 13862 (MO) tou3" 35k W of Loja, bein Sun Pedro and Chinchar, Espinosa 1295 (US) Pet nia Casein it Rio Pachachaca Valley, 33 SW of Abanoay, Hs et al 838 (NY, WIS): Ahareny Fargas 9154 (MO} Haceeda Luce. Oropes Voy Vargas 9773 (F, ©, UC) 61210: roy. Coorencie, Ro saris Mayo. Alfiro oJ (MO); Prov. Canvencin, Bek ter Fargat 10435 (F, C, GH, UC} Prov. Convencion, lrodederes de Quilebama, Vargat 249 F Prov. Con Veneida, Cheuea Mayo, Verges 5472 (MO), Husscnteuce: Teyaeaja, Huachocolpa, Quisabamle Tooar 4145 (UISk Tayacaja, between Che}ya) sal Su cubsorbs, Torar 4579 (US), Laxmay aut: ke 28 Bot Gino, bane Olson and Ines utchivon a Wight 3417 (NY, UC) suMib~ betanen Quebrada Seca y Ma Isp, Fetrayra etal. 10085 (BO, US). Annals of the Missouri Botanical Garden ‘This epithet hoa traditionally been associated with Z, portoriconsis (Mackride, 1943), porhaps NEXT Deeause of the superficial similarities between thie species and Z. caracasana. ‘The type material of “Calliandra weberbueri,” bomever, has # nt ber of features suggesting ite inclusion within Z ‘caracasana, For instance, the calyx is deeply a vided, and the filaments are “whito/inkish:rod™ as indicated in Weberbauer's collection, In add tion, the “prominent” leaflet veins in the type collection, ax seen under magnification, are like those observed in many other collections of Z Acacia fragrans Ten. nor. nv. Cat, Ort, Nap. 1 was apparently never described. Thore i, how: ever, one specimen at K, cleaely ideniified ae Z. caracasina subsp. weberbaueri, whiel was pre sumably collected by Michele Tenore in the Orto Botanico di Napoli ts original source i unknown, 9. Zapoteca tetragona (Willd) H, Hern. Ann. Misouri Bot. Gard, 73: 755-763. 1986. Acacia tetragona Will., Sp. PL. 4(2): 1069. 1806. Calliandra tetragona (Wild) Benth, J. Bot. (Hooker) 2: 139. 1840; Benth, Lon don J. Bot. 3: 99. 1844; Benth., Lanvlon 1 Bot. 5: 102. 1846; Benth., Trans. Linn. Soe. London 80: 543. 1875; S. Watson, Proc. Amor. Acad. Aets 22: 410. 1687; Britton & Killip, Ano. New York Acad. Sei. 35: 137 1056; Standley, Publ. Field Mus, Nat. Hist., Bot. Ser. 18: 493, 19.7; Stanley & S, Cade. rn, Fl, Salvadore 1941; Standley & Stoyerm., Fieldiona, Bot. 24: 27, 1946; Woodson & Schery. Ann. Missouri Bot. Gard. 37: 261. 1950; L. Cirdenas, Revisia Fae Agron. (Maracay) 7 131. 1974. Fouilleew fetragone (Willd) Kuntze, Revis. Gea. PL 189. 1891, Anaesta tetragona (Will). D. Smith, Enum. P?. Guat. |: 10. 1889. Britten & Roso, N. Amer. Fl. 23: 64. 1928, Mimosa ‘quadrangular Poizctin Lam. Bneyel. Suppl. 1: 72. 1810, nom, leg. (This cuperfuous epithet was based on Willdenow’s deavia te tragora.) TYPE: Venenuela. Caracas: in rips arenosis Ruvii Guainite, Bredemayer 17 (lec totype designated by Hernindee (19860), B. Weat.no. 19147, not seen: photos at MEXU, Mo). Callin portoreenss var. malige M. Micheli Bes Gaz. (Ceaufrdorl) 20: 285. 1895, 1v0E: Grote real, Senta Rosa: Son Tuan Utaps, 4.300 fe, Oe. 1895, Heme & Lax 0090 (letoype here ess sind, G rlectorypey Gl US ( Caltandr tonona tom & Roe wy Beton 8 Kili,PREV ‘Volume 76, Numbar 3 1989 FicuRe 40. Dsirbution of Zapoteea tetragons Ann. New York Ae. Sol 35: 197, 1936. rH Calorbin Astiegsc: vain of Modlin, 18 Avg 1920, Toro 483 oletypes NY phot, Ch Mostly shrubs to 6 m tall, rarely trees to 12m. tall; stems 2-4(-207) em diam; branchlets coe: spicuously 4-angled, sometimes herbaceous in the younger parts, wit the surface costare under mag rifction, mostly densoly villous with tawny hairs ‘when young, boeoming glabrate. Pinna (3-)1-7 jugate: petioles eglandular, with a conspicuous ‘adaxial chansel, angled, 1.7-6 em long, glabeour todensely villous rachis(1-5-)8-9(-11.5)em long, labroas 10 dencely villous; racillae (3.5-)1.5- 10-15) em long: leaflets 10-35 pairs per pina, narrowly oblong, oblique at base, rounded wo acute at apex, the median leaflets (58-18 > 1-4 rom, the proxieal smaller, all membranous, gous oF ciliate marginally, always glabrous on both sure Facer: elle renation inconspicuous; stipules leafy, alpressed, triangularlinear to triangularsanceo- Tate, acute at apex, straight or curved parallel veined, (0.5-)0.8-2.5 em long, elate, villous oF lobrous. Capitula axillary, rarely arranged im pscw doparicles 6-22 om langs peduncles fascieate, (25-95-12 cm long at anthesis. Calyx 2-3 mm Tong, the teeth delta, eliate: corolla furinelform, (598 ram long, glabrous, tho lobo= Tenceolate, sometimes with tulte of hits apically: Blaronts ‘ueually €2. 40 mm long, white, tho s1aminal rubs ‘a, 3-4+ram long: ovary 2-2.5 mm long, glabrous, shortly stipitate, Pode to 16.5 % 14 em, at apex truncate ro obtuse, rstellate, thickly membranous. glabrous. Seeds usually ovoid, 5-6 « 3-5 mm, rmetly black or dark brown; plourogears regular, obscure, Somatic chromosome number 2n = 26. Distribusion (Big, $0), This widespread species hhas boon collected primarily in moist thickets, along streams or river banks, aad, in general in ruderal Iabitats derived from topical and subtropicsl ev. cergrvon forests, Its range extends from Mexico (Nayarit and northern Veracruz) southeast through, Geatral America along the Andeaa Cordillera in which it reaches southem Ecusdor. Popubstions develop primarily at moderate and less frequently a low altitudes (0-1,500 s), slthough in some parts of Central and South America they ean reach Alitudes of 2.300 Common names. Angel Veracrn,stzalaguio (Puchla), harba de chivo (Weraerue), cabellito blan- +0 (Oaxaca), cabellito de angel (Michoaein, Guer- rero, Pucbla}, cabeza de vie)o (Oaxaca, Veracruz) ccarhoncre blanco (Colombia), earboncila (Costa Rica), carboncillo de for blanea (Costa Rica) cola de iguana ([alise9}, guaje silvestre (Oaxaca). gua jile Gaksco), guajilotsneo (Oaxaca), lig iu (Oa. xxuca, palo cosilo (Guerrero, pelo do angel (Pueb- 1a), porosina (alisco), quebracho de montata (Honduras) caqie’alib (Quecehi nume, Guatemala). Aditonal specimens exomined Mxico.cuiaesss Mow. Tene Menhen Chav, Breedlove 2601 (8 ERC, MICA US), Bae Oeamesasthy 1220 bo N Gevacouils aking the ef. te Malpena, reediove Thorne 20995 (DS, LL, MEXU, MICH, MO Mio. NEXTPREV Mapsctpec along Rio Testecapa 10 km SE Mapestepes, Brevdiowe & Trorae 0739 MENU, Mpio. Frontera Gomalsp, slong rd, 19 Cad Grauhtemony Breedlove 39086 (0, MEXUh Mpio. Paalcigo, Abk'ulbal Nab bore Poltleigo, Breadiove 50102 (MEXU H Sal ator, near Slowilizps, Collins && Doyle 229 (US) Lago Sopa, Rewrss Montes Accs, Gove rol 220 (MEXU) Iahustan-N, MacDongott 76 (NY, US}, Ex fins, Tura, Metude 27074 (DS, F, MENU, NY) Guaesushd, SE de Mopactece, Miranda 6270 (MEXUe oar Yajakin, Nelsou 3417 (NY, USF near Tape Kelton MH (NY. US: slededoves de Reyéin Ramos € Fink 501 (MO), Mio, Venustzue Cartan, above Soyattan, slong rd fom Las Rosas to Pug, Ton 3135, (DS. ENGEL F 1, MICH. une Mp Carla 0 5 Ne F1 Neca, Siagallans Eat 2820 MEX MICH), cernar: Bs Cabra Awa Hen 10842 (BM, F, 6, GH, MO, NY, USp Dic Caleams, Fon de Curran, Hinion 11059 (DS, F. GH, MICH, US): Det Montes de Qos, Vallocites, ffnton 12780 (DS, GH, NY. Usk Atovoe, Kerier 760 (HM AR. K. MICH,” US). Jasco Me pes of Coren as Mig steal gen Go Serva de Macaotli, Ilan eft 470 (MICH Mp, Pibuato, El Agoraders cance 2 Pihuarn, Marines «eh 1404 (MENU 16 oh, SW Auten, MCVaueh oak 10858 (ICH) Goacalaco, Palmer 65 (RM, GH, NY. US) bso: Dat, Tenaratepre, Nejc, Minion 2029 (MENU, Dis Temasesiepes, Teale, Hinton 2207 (A, BM, F.C, Dit Temacealioper, Acailén, Hinton Tor (8, BMG F, G. MICH, US). cuoscsy: Dit, Cock sornin,Zarmnra, Hunton 12244 (DS, Gil, MICH, NY. US, Dis Cosloomsin, Gonkomin, Hinton 12381 (DS F, GH, MICH, NY, T, US). Dat Coaleorin, Serre ‘Terra, Hinton 15278 (GH, US Puente Alto, Li Jem Aro do Rosle, Souca & Soto 8008 CAS). NonEL0s Alpuseca, Amar 17 (MEXW}: Cunt. Las Taaas Levy 1S (ENCIiL Hoajiain, Lyomuer 374 (BN, MO, US} Yavtepse, Pringle 9727 (fy C, Cll, K, MO, RY, US) avant carretera Boers VallneJakonba, Page 22 GENGBE Sen Blas, Hult 3027 MERU, US). ost Moo. Pochtl, 2b NE Chacala ota hm Pune de San Juom, Delgads 604 (CAS, BO, UC Dit lel tn, Mylo, Sea, Maria Chinslpa, Matias Rosner, Co Tonia Cauhtemce, Delgado 923(ENCB, F, MERU, MO, Yi Dut. Putla 1.5 hom NE de Putin por carta esis fe Hercvlex & forves 43 (MEXU, MO} Sie de Sin Pedro Noles, Tales, Jargensen 702(BM, ines Calderin 297 (ALL. UC, USy, Choapsm, Vave, near Hie Yoreo, Mesia 9270 (B. CAS, F, G, GH, MO, RY.S"UG US) Ranchers Tohepee, sominn a Plime Hidaigy, 19 kom N Pocula, Soune eal 338 (CAS, US, WIS Dist gulls, 8 Ero NW de Sex Gabriel Mice epee, Soura ot al 4375 (MEXU, MICH, MOR Dit {Tenn 2 kr al Node Fuente de Fiero, cn le cor. Sta. Ma: Galchri, o sda 4 10 km al NE de Toni Sac eal 8259 (BR, Dit. Julep, 1) ko SE de San Sotastin Tncspa, Sousa 8450 ENCB, MENU): Dist Pulls 2 Lin Puta de Guorero, Sousa etal. 280 (ENCE, MENU, MICH, Dist Juchtan, 10k NW de Palomares, Sons & Teles B77 (MO Dis. Tastee, Bs. Lar Boni, San Pele de lo Pofa, 9 bee NW cel init Osases-Veccorsn act. Matias Remora Tuxtepe, Souse € Toles 8800 (MOF Dit. Texiopes, Mote ers, 5 ka NE de Valle Nacional, Sousa & Rien 9255 (CAS, ENCH). Dist Jarutepec, 12 kt N de dewitci al Vvero Gap tl hae Sam Asin Chasen, Torres 1a. 1636 (MO) GE, 6, KO Taps Goes tn vice Ma. ‘Annals of the Missouri Botanical Garden vet Mio, Pantepee Lm N de Mecapaap, Basurto 8 Dusdu 118 (CAS ENC, MOy Mj Mae te Ides 1h de Vals Al Camschn, Basra Dorm {194 CAS. ENCB, MO} SE Assets, Mirando £108 COAEXU) Nennen in carr, MizailasNacinon, Aven doo 117 NCH, F, RY Vale eCard, Boareran 1s (G Gli, NY, P, 8) 1 ken NShuapln, Calera 495 (CAS, ENGB, F, MO, NV} San Ards Test, Rancho Palzogiapan do Don Arsto 18°27 N,95715°6), Chlsada 42200F WIS) mde Papal, Crh © Johrsion 5223 (TER), Mp de a Palo, Le Pala Poiveroy Belg ot ol 896 (CAS, ENCD, MO}, 16 del camino Ceilo-Ls Lagu, Hilaslin, Dorantes ot a 407908, MO. NYie Maacia, cairo & Tacha, € Heradndes ef ab 203 , NYX bipin Goemaca, T nde Souecrsapen pon caw a Onaga, Hom Indes & Torres T0r (MENU, MO} Mp Cabals, 1216 kom NW de Santiago Tuna poe cart « Leo de ‘jada HL Hirndndes & Torres 0 (WEXU MOS Mp. Catewaco, Rancho la Pina, 25 kin de Catena forsumin » Montp, ete Bean de Rises on Tatas y Sooecinpany 2. Herndndes & Tues 799 (EXT. MO} Mio. Catonaco, 5.7 km de Catemaco por sui Moi i Homies & Dir 0 MEN, MO Mio. Crdsha, sobre te Fortin-Cindobe, 1H. Herninde: & Tres 27 (MENU, MO, La igus tence Sn Toné Avatena y Husytanaloo, Mrques et al 1357 iy lana dee Lave, Marta Calderén 1540 (3, BM, CAS, FLL, MO, HY, US Mpio. Ctemac, Pie Ar rita Caron 1808 UCAS, ENR, Pisin RYU Mpi‘Totoyuen 1 on SE. Tetons Slang Hing 108 EN, SHDGW), Nr xa 1590 (3 kan Eu Angel Cabos, Neil 5388 (MO% Barrance Sal Fortin, naar San Martin Tacatpoc, Purpus 16439 (A. EUS Minsrla and Papania, Schiede & Deppe 691 (OM, G, GH, K, MO, NYX Zonghea, tain cum 2 Zamalapan: Neca (18°407N, SOSOW), Page? hou NG Fs Ko Sloe, cme Cole» LE und, Hsalgetilin (7™16°N, 93°37 Wt ft 1334 (BM, MO}, Uaslo Galvin, Lows, de Sa foe: Femara 583(CA8 ENCH LD), Ati, Kose, Fengura 4976 (CAS, ENCB, Mictl, TEO. Goateanee {sth vines Ci, Tarcein 691 (GHB, US) ses Sea Chace, Wis 4004) cavguitx Wel i IplaseAmaly Stroma JB ene Dole siver ban of Bio Sapo Cuntoras 10580 LL, MEX, §)eactese Sun Jos Matin, Donne Sith 2263 (K, USh nr Escala, Stomsley 05903 Wha Ted butmoon Ercole snd Finca Zapato, Stoney 64910 (A, Peas fio Secapa SE Exu Stuntley 89266 (9) vzaat: El Eston, Contreon TET GULL MENUS) gencuurensoe® Intooeh Coiba fa Moreedes, Stondley 87992 (8), nstmanutre Be thal Bersoall & Caro 1193 (Ky laa Vetoes, Hon 240 (Gy sinty of Lax Deleas, © Retake, Stintey Reb07 [E. sexterbgies mene ae La Standley 56102 (Fy sux wan Finca Hl Posen, Potrero Tos, Voleén Tomko, Seeyermark 37648 (7) sat ROSK Cail, Heyde & Ls 4148 (6, OB. USK regin of Petenren, btwoen Toxo ad Caacapon, Stoniley 79085 Gh non EtMotinn, Ste 70241 (US) secumuesgitz near Maaaovango, Standley ‘bone (Pesan Peencieo Zep. Steyrmare 32143 (Boz. 2 Car El Coy, Gran Se Oro to Blo {ale some lanier rll ayer P0894 NY Nae, Gente 2188 (A, MMTCH 37 on cto, Haring Ninoy, Gente 8998, 1, MERU. MICH, Sh 6 em NEXTPREV Volume 76, Number 3 1989 B Cave's Board, Seuss of af 12189 (MEXUD; Masel iver Inuons at Ue ed of Kinloch Ra Spetlma 1424 (MO), Barron Ceoek, Whieford 2097 (DAI). BL Sut Don. AMUACiLDGN vicky of Abuoehapn, Standley 108. AGH, NY, USp near Sako Atchucci, Soudley & Padilla 2873 F, NY). ta unerTaD. San’ Ansrox” Rio Saco, Clausen & Rover 6208 (MERU, BY). sat seivacons vicinity of San Salvador, Standley" 79206 (GH, NY, US) Sevalavse Sof Santa a, len & Armour 7069 (F, TL NY, US) Hono nas. sruatinss vcnity of Tes, Standtey 55161 (A, Fy NY, US). consrscen: 8 fan 6 Son Jor Cemayasuny tina & Melina #1472 (€, M0, ora: ietween Copin suns and Santa Ria, Motive & Moline 24661 (Fs MO, NY. US) conte’ gear Lake Yojon, Wilioms € Wars 16700 (P GH, US} sax sigue: Rio Ssearpa, Molina 978 (P) ouancio. le tween Jutcalpa aod Carpaneoto, along fio Juelpe, Blackmare & Heath 95” (BM, MOR Dukes Nombre de Cals near Les Moris, Blockare & Heath 1605 0M: senity of Cataczmas, Stanley 18146 (F). YOR: Poe Sulitana, so Hagen & tom Hagen 120908, NY). Ne aRscLA, anorace etre Wisi) El Carmen, rairgen We Cove (15°37, BS'SD'W), Aeaguisnin & Maren 1587 (MEXU, MO} 20144: Blueelde, Cra, Long 100, Py Le Poctorla, 5 hn W de Wark carrtera Wasa 2 EL Tums, Mereno 29135 (BIO carzeere nueva. Wank, El orrnguor, Orie 1225 (MOL Amparo, Ort: 1304 (MO), Ma, Sona, Sannes atria, Orvis 1313 (MO), seer de Nogee Was, extre El Exile » Rona UsHIS N, 8:°25°W, Graz 2184 (MOY Sato, slong Rio Pis PE: and surrosnkrg illu. (LAOU'N, 83°38), Pipoly 033 (M0 Caso nom, near Rin Karsh, Ser bet & Morena 197 11 (MO) Coste Ics. MLATCHLASSa Pode So Reo, Brenes 14474 (NV onde el Rs Busi, flodnes 1709 (F NY, US); Alfa Ruz, Plea, Smith 1202 (F, NY), antco: vein of Turraic, Cojrey 66192 {MOp. Foes Perla, este Reserva do Vide Svesue Tapansi y Push - Heradndes et ab 073 IMEXU, MOx 2 kia de Cervaties por cars San Jose HW Heraindes et al. 730 (MEXU. MO) cowrcasTe Finca la Paces, Opter 858(P ence Liberia y Ques Grande, Poroda 1137 (F, MO} Quebrada Zopibte, Par ‘que Nevional Rinebn dela Vieja Souse ot al 12750 (Moy vicinity of Tihrin, Seondley oe Valerio 45021 INY, US) TeRsDIM: Booka, Bio Segundo, Lean 82 (P) tae Los Diacaites (US ruber sei) on 9 Gae piles, Carlson 3429 (P, BIEL, US), san soste Rio He Fradiva,iiburars Rio Chris det Pacifico, NW Canain, Ganeral Val (O729"N, 83°37 W), Burger & Lecsear 7083 (BM. F, MO, NY, Usk Sex Jose, Tendus 1505 GBM, BR, G, USK The Torres, Fonds 2800 (BML BR. F, GH, HMO NV. US, slime Mo leo, Eines Ei Qazanta, Wittams er. 28404 (P, NY, US) Pawan, Sokighi! Cerro Horquets, Blam & Dueyer 2630 CL Dit. Boguete, above SI Gansinro, D'irey 9825 (MOK Det. Bogue. Bsja Mono, Dacidion 920 1 Fy MOK along flighvay te Concepeién at Cuesta de Piedra, King 5300 (TEX, UC. US): between Ho del lob ard Ceres Vaca, Pion 5423 (Sh i$ El Yolean Tyson 73, (MO), coeaé: El Valle, upper Rio Astin, Alon 3819 (6, CH, MO, Salons road 2 ri ahve EI'Valle on cod 19 a Mesa, Cros: 28297 (MO, NY). vautse Rio Puewre, Ietween Crem Malrand Coro Taearcana, Gentry & Mor: 1.389 (MO). vERactas 3, W Santo Fe tr pst veurlsAgricls Ale Pied, Croat 23708 (CAS, FBO. N¥p Vesteunta.sicrano reocast: Corilera de Avi Ary von facing open alone Carus, Steyermark $5147 Hernandez Zapoteca (MO). sésany: Depin, Tovar, aleededoces de Tova Baulie 5ah2 (F). tuna: Los Guayaliton, see de Barut, srivoguiete 5952 (NY) Peace and visi, near Caracar, Pitter 7471 (CH, US) ‘remnas entre Fle Grande y Labetera, Lo Llaneda, Cardenas 1243 Py borween Las Dele ad Paras along Rio ach fa, Sieyermart 57420 (Fy, along highway. N eee Delci, Steyermark & Liesnor 118750 MO), ru Dept. Carache, suse El Paraiso y El Balin, Benftes de Rojas 1980 (MO>, San Jack, Varaayo. 1607 (US) racinionts dl Ro Momboy, La Puerta, Tamayo 1811 (US) Cotouals. snout: Birbe, vemty of Bloelin, Toro 577 (NY, US ROvAcN Corlera Oriental verter ental, entre Guetoguc y Cunyats, rgenes del Kis ‘Sinmun, Cueireeuses 9702 (F, US). cata gran eae. tera Chibi, Dagee Jaramillo 4017 (Fy NY, US) atca: Popasin, Loken 9468 (6, GH, 8, US sroon (Cal, W ee of Cauea Valley, Pater 926 (US). cunoa ates: ene Chipogue 7 Cann olededres de Puente Servien Gurefoserriga & Yaraimille 20550 (F). x= tints: Secru Nevala de Senta Masia Quebrada, La ‘Siena 10°50N, 13°59°W), Kieelvide 2165 (US) Nonr hie waraspe weniy of Chinkeots, Killip & Sith 120051 (A, Cll, NY, US} Calaga Valley, uoat Tepaté N of Toledo, Kulp & Smith 27141 (A, GH, NY. US) Tous: El Libeno, Gaseia Barriga 12227 (US). Vale Cord. Occietal, verte er Sdetal Haya del Rio Sr fui, La Laguna, Cuatrecases 15674 (F, US Car Ailere Oosienta, Hoya Sel Rio Albin, El Zaeudo, Cus trecoxas 22100 (F.S, USk Dita Johason 2460 (B10), [Ecvioor. azvay: along Rio Pra beeen Hasienda Yuba tnd Tne San José de Coit, gion of Sone fain, Steyemmark 52737 (F, MO). wolivan: Cord. Oc ‘dental, Lanse, acorn 6343 (R), came a for CChieal to Maldonado, long Rio San Juan, Carry & Shipp 26621 (MO), cininon sor vicinity of Vents, Rove & Rose 25527 (GH, NY, US). errupext: Quevede. tac oa areas ang Rio Pek (29E10'W, 1535), HolrNistwen'e! wl 3075 (P, MO, NY). Colo USA mort Sins Lous, Mlssour Boteniel Card, I Hernindes 956 MEXU, MO), wv, Mosmaingarien ‘Todas Mt. Cede), ean Oostrrocm 13707 (BR Zaporeca tetragona is closely aed to Z. por Joricensis bat can be readily distingaished pri marily by having conspicuously tetragonal branch _ lets larger corollas, and longer filaments. Despite tho opinion of Weadion & Schery (1950), who suggested that Z. tetragona (Calliandra tetra- zona) is distinguished “unzatually” by its tetra. onal branchlets, implying that itis not difleren- tiated from what portoricensis complex,” this entity is maintained here as 0 separate apécies. Discusion of the re= lationship between Z. tetragona and Z, portors ‘ens is provided under tho latter species. Zapotvea tetragona is probably also related to Z. aniina, from vhieh i ean seadily be distine fguishes! by having adpressed, usually larger, ti ngular-linear to triangular-lanceolate stipes, dele tate calyx teeth, and totally white filements. Discussion of the relationship between these tira species is provided under Z. andina. hey call the “ NEXTPREV 838 Acacin quadrangularis Link (Enwsn. Hot, Bee rol, Alt, 2: 445, 1822.) has traditionally been ne clued in tho list of synonyms of Calliandra tetra gona, but I have been unable to eonfiem this synonymy. Link did not cite Willdenow's sloncia tesragonn, nor Poiver's Mimosa quadrangutaris, snd given the Fact that almost all af Link's personal hherbarivm was destroyed during World War I, thore i no materi] avilable with whieh to evaluate this hypothesis 10. Zapotecs Beuador. Amey: valley of the Kio Tarquis few kr south of Cuenca, 2,500 m, 1-2 Feb. 1945, M. A. Guiler 47 (holotype, US no, 2106703: isotype, NY). Figure 4 lina H. Hern. sp. nov. PE: Fratices re presser jurenitius conspiou er orks. Prac, 3-Cjgay fobobe 15-32ge fn plone ‘Sau, argue blog tplaevadonoanarglnee best tno dnbor descendentns ye) depres praediae- Calyx ca. 2-mm longo derihor angts-blog Fs rents bos ala ad pice purpurea Shrubs to 2 m tall; branchlets eonpicuously Acangled, especially the younger ones, costate one der magnification, glabrous to sparsely villous, Pine nae 3 6,jugates petioles eglandular, with a ecm: spicuous adaxial channel, 1.3-4 em long, glabrous to villous: rachis 2-4 cm long; rachillae to 8 em long: leaflets 15-32 paits per pinna, narrowly ob- Tong, oblique st base, acute at apex, the median leaflets 5-10. < 1-3 mim, all membranous, glae brows oF ciliate; leaflet venation inconspicuous ste ples leafy, shallowly trangular, with 2 basal ox Fieles, descending or depressed, parallel-veined, to 0.6 cm long, globrous or ciliate, Copitula axillary: pevdoneles faseiulate, $.5~7 ern long at antes. Calyx ca. 2 mm long, the teeth narrowly oblong acute; corollt funnelform, ea. 4.5 mm ng, las brows, the lobes lanceolate; filaments white in the taal half, red-prple in the distal half the tari tube ea, 2 mm long: ovary glabrous. Povls ca. 7 Lem, aeute at apex, rostellate, thiekly mem- branous, glabrous, rticulate.ceined. Seeds oveid, ea, 5 3 mm, motled; pleurvgram irregular. Dissribution (Fig. 42). Zopoteen andina is fenvemie to restricted areas of the Andes, in the provinces of Azuay and Loja, Eeuaror; the de- partments of Cajamarca, La Libertad, and Piura {io northern Porus and the department of La Paz, Bolivia. This specie is allopatric with Z, etragonas taken together, the two species oceupy a nearly continous area of the northern Andes (Figs. 40, 42); 1,400-2,650 m, ‘Annals of the Missouri Botanical Garcon Additional specimens examined. Rewsnon. 104s: Hac. HusicosPani, 35 han Se aja, Espinosa 2229 QUICK) Pesc, conrusazs: Depo. Cejaunren, La Mow: {aia (CumargoCantumaei}, Segdsiegus et al 930) (010): Youn Cumuango, Seadsicen 10821 MOR Dept. Cajmareas Quecrade Heyyia antigo Yond, Saree eg Tapes 1062? (MO) La tine ao: Prow. Ota Wiseanebsh Laps et al, 2648 05). pti: Poor Can shaguey Ferreyra 3135 (MO} ice Pala y Fuique, ferent Canchaque, Ferayra 10920 (Us), Bouts. 922 Sorts, Holway’& Helcay 520 (CH, US Sorts, propo Expada, Mandon 759 (HDC, BR. GH. K. NY ‘thot locality, Bang 7812 (OM, F, GH, MO, NY, US. Zopoteca andina rosorbles Z. reeragena in most of its vegetative Teatures, although the Deanehlets are sometimes visibly four-angled anly in the youngest parts. I differs from Z.tetragona primarily by having smaller, triangular, descending for depressed, auricnlate stipules and by having narrowly ablong calyx testh, mailer floral dimen sions, and white/red-purple filaments. Available collections of Z. andine do oot eeveal the habit, aod it is uncertain whether plants of Z, andina fare erect shrubs or low, subprostrate or prostrate shrubs, as suggested by the aspect of some of the Bolivian collections (eg., Bang 1812, Mandon 759) 11. Zapoteca eostaricensis (Brivion & Ross} H. Hern,, camb. nav, Annestia costaricensis Britton & Rowe, N. Amer, Fl. 28: 67. 1028, Calliandra costaricensis (Britton & Rose) Standley, Publ. Field Mus. Nat. Hist, Bot Ser. 18: 492. 1987. rvs: Coste Rica: borde {4u Rio Convento, Jan. 1897, Pittier 10643 ectorype, US no. 577350 hore designate: isolectorypes, BR (2), US (2), Shrubs; branchlets slender, terete, smooth, gla brous. Pinnso (2-)3-jugates petioles very slender, ete tetragonal, with « conspicuous ad chaneel, 4.3-7 em long, glabrous; rachis 2-2.5, frm Tong in 2jugate leaves, 4.3-6.2 ein long in Bejugate ones, glabrous or glabrate: eachillae 7~ 13 cm long; leaflets 10-13 pairs per pinns, the median ones narrowly oblong, the distal ones ob Tong-obovaw, oblique at base, acute at apex, 7 x 0.5-0.7 om, the proximal pairs smaller rnarcower, all olve-green adaxially, palegreen ahaxially, membranes, glabrous; lealet venation inconspicuous stipules leafy, avate to wiely ovate, rounded to cordate at base, acute at apex, to 13 % 8 mm, glabroue, palmately veined. Capitula ox lary: poduneles 4.5-6.5 em Tong in fruit. Pods to ces. 15 ea ong when mature, acute, rostate, thick. Ty enernbeaens, glabrons. Soeds evil, 7% 4 ane, Irown; pleurogeam ierogular NEXT837 Hernandez Zapotecs 1989 Volume 78, Number 3PREV 898 Proune 42. Distribution of Zapotrew odin rally due to the paucity of material. On the basis tf leaflet morphology and obscure leaflet venation patterns, Z. costaricensis is probably related to Z, portoricensis subsp. flavida. Zapotec costart consis ean be easily distinguished from that entity, however, by its muck larger lellets ani) charac teristic ovate/wilely ovate, cordate stipules. The narrowly oblong leaflets of Z trast with those of Z. formosa, which are almost always obovate, 12. Zapotecs formosa (Kunth) H. Hern. Ann. Missouri Bot. Gard, 73: 755-763, 1986, Acacia formosa Kunth, Minwoes 102, 1. 32 1822; Kunth, Nov. Gen. Sp. 6: 278, 1824, DG. Prode. 2: 466. 1825. Callendra for ‘mosa (Kunth) Benth., London J. Bot. 3: 98. 1845: Bonth., London J. Bot. 5: 102. 1846: Bently Tran, Lina. Soe, London 30: 542, 1875: Beruh. ia C. Martasy PL. Bras. 5:41 1876; Loon, Contr. Ocas. Mus. Hist. Nat Colegio “De la Salle” 10: 237. 1951. Fruit Annals of the Missouri Botanical Garden deca formosa (Kunth) Kuntze, Revis. Gen. Pl 187. 1891, Annestia formosa (K unth) Britton & Milkp., Bahama Fl. 159. 1920; Briton & Roce, N. Amor. Fl 23: 67. 1928, rvve: Mex feo, Guanajuato: Bonpland 4288 (holotype, P. seen in mierofiele; type, P). The Label information on there spec vate the locality. but according to the de ition, it was collected in Gnanajuato 1.068 hes (1,922 m) elevation. The holotype i3 ox wemely similar to and presumably served a5 ‘the model for the illustration in Kunth’s work (ig. 43). Erect shrubs ta 5 tall; stems erect, sometimes drooping usualy slenler, sometimes to endian: branchlets terete to angular, Leaves eglendl the pinnae 1-3(-4).jugates petioles terete, with & ‘conspicuous adaxial channel, 0.3-7.8 om long, slabrous to villous; achis 0.3-6.8 em longs rach We 00-125 em long: leaflets (2-)3-15 (17) pairs per pinna, oblong to oblong obovate oF very widely obovate, the proximal ones sometimes Praune 48, “Acacia formas” (= Zapotecs formosa sep, frmoss, fom Kanth's Minones 32,1822. This slau acs sly asl or Bonpland #068 1 coach wal a Conran, Ma. NEXTVolume 76, Number 3 Hernandez 1989PREV BA icune #4 Annals of the Missouri Botanical Garden esibutions of Zapotece scuelfera stats, Z. formana wip formosa (los), Z. formas sp. soleadoronsis aah ioes). Z formosa svbep. gruel: (pen iaaghs}, 2 formar sap. rose (pen eitees. 2 formosa soba, mallicula (id eangles), an becoming elliptic or widely obovate, all oblique at the hate or the proximal ones sometimes bevorning seule, rounded to acute at apex, usually mucre nate, the distal lealets O.4°3.9 * 0.23.6 em, the proximal ones smaller, mostly membranous, glabrous to pubescent, pilose villas; leaflet ve ‘ation mostly inconspienou, primary and secon ary veins occasionally conspicuo.s; stipes leafy, ‘riangulartolineae-triongular, or lanceolate to ovate- Janceolete, to 13 mm long, glabrous to willows, pitula axillary oF in simple peoudopaniculat in orescences; peduncles usually fscieulate at dis tant nodes, 1-11 em long at anthasis. Flowers peotamerous; calyx cup-shaped, LSA mei long, flabrous, the teeth triangalar to oblong lanceolate, corolla caropanafate, 3-5 mo long, gaeous, the Totes elliptic to lanceolate; filaments ca. 2.5-8 Tong. white, greenish white, red-purple, oF white io ‘the hacal hall and red-purple oF pink in the half, the staminal tube ea. 1-3 ram longs ovary cea. 1-3 mm long, glabrous. Pods to 130-15) exo Jong when mature, at apex truncate to rounded oF acute, usually restate, thickly membranousy glo Ibrous, glabrate or densely pilose. Seads rhomboid oruoss bsp. set on squares). ‘ovoid, sometimes vory widely 20, 1-8 * 3-6 rom, brown or gray, the areola usualy lighter: pleuro luram irogular, sometimes somirogulae, Somatic ‘chromosome number 2r = 26, Distribution (Pia. 44). Usually common in disturbed or undisturbed habitats derived from arid pica deciduous forests, and lens frequent- green forests This species ranges from AAvizons to Argentina and into the West l Gallections come from 02,000 m. Zapotece formosa os circumscribed here con sists of six highly vanable aubspeciess formosm, salvadorensis, gractlés, rose, mallicela, and sschotii, This infracpecifc clesification is based primarily on leaf and flowor characters (Table 14), Varying degrees of morphologieal intergradation cccur between diferent pais of subspecies. A total of 24 specie epithets have been published for plants that fall within Z. formosa. Most of these rams really designate regional variations i leaflet characters. Ie light of the enormons degrer of morphological variability within this widespread species and the relatively common occurrence of NEXTast Homéndoz, Zapoteca Volume 76, Number 3 1989 NEXT or ee oR eae a od 2 wer "6 “TT = wm SEL sand ora si lage ayddepasaimin apn, oo one esta spayed axomowed oq poond ames apm amps geno souamiag mt ¢ 6 ¢ ec ad 0g ensign kame snore? 9006 aa ae 2-)0-E LHP SEX Tek MO LL-G'E « ZE—B UM BI-L x 9G-OL tela) ee 208) oF wo or-8't eo sti-se wm osesceT we er-2T set wer rea et & We Be Tape apo Ta pe ssbedens oedens aces molars PREV row vaned07 jo eonateqs v9 J une ‘PU SILPREV ane intermediates, 2 conservative approach has been ‘chosen im this treatment. The following sequence of the subspecive reflects their putative relation ships. Poe pe: plant by thought — 12h tap. salbediresia 1, Pode abou ar Rabat; pant glbroas to heey lament ste or aresrich white a Leas blog. laeows sul ma iy tongue on kg ae sul gris 1b, Eze ebbngalavatey glares to vlibos pu needa to ote ole, ws hing Ta. sulap formosa Fisnaicvvedpigew siecle tial redgarpe ork aly So Toake acids 2-0 num; capita ‘ayy Stamens white bony and Sites pak ovr purple aly Sa Pecos $11 mn lng Tae tbsp. meine sb, Pavlos 12°95 mm lone ip chen a ob ae Lents 831° 3°31 ame ape cailry or preudopaniulee lo nents er wooly eed paper wie sland respagl fo pk dally Ba Sep ese ab 12a. Zapoteen formosa (Kunth) H. Hern, ‘subsp. formosa, Cltiandrseupillae Benth, Lonon J. Bot 3:98, 1844 Bent. ‘Trans. Lin. Soe. London 30: 542. 1875, Aanesia capita Ber‘) Briton & Rose, N- Aer 125 68. 1998. tare: Mexico, Oaxaca ad rcose Mois San Febpo, Jab, ndvieus 404 (ectotype, IK, here designated welectoypes, F (rasrent), KP) Calliendrs matacopiyila Benth. London J Bot. 3: 100, 144: Benth, Teens, Linn Soc. Lordan 30: 544 1875. tyre: Menico, Ossaca avatiees Monts San Felipe, Jao, dries 401 lototype, Ky here de ignned:icectoypes, fragments G, K.P, US (Gragwenty, Calta vinifera Bevuh., Landon 1. Bot, 3: 98 td, Mert, Trans Lixn, Sa. Lorn 80: 542 1875. rare: fea: Tueedie sr. (ectety, K, he Aesgnated boleesotype, K) Caltiandre grails Klotz ex Raker, Refug, Bot 5 ‘204, 1878, non Crea, 1861, nov Stacey, 1929, te Reavlae, 1981: Benth, Trane, Linn, See, Lon Sk 30: 542. 1875; 1 Candoras, Revise Fac. Agron (Maeacey) 7: 181. 197%. Calltendra jormose vor dracilis(Klotsch cx Baler) J. Macht Con Gray Heeb. 59: 4. 1919. Callandra moritsions T. Cirdenss, Fema 25° 40. 1983. (This name seas pullished an omens noeutn for Callender gracilis Kieth ex Baker. The designation of frits 8B? (UM) es the lectotype made by Ci ‘Annals of tho Missouri Botanical Garden devas is inorzects the ype eollction s Otto 404, ‘which Klotact xpliiy sited fn the orignal de Seiption of © gratis) reves Colonia: Orin 404 (etotye here designated, LE) The holaype, which realy wae at Ty wis probably dated {o4s (P. Hiepio, pers comm, 1986) addons, duplicates ofthis collection ero ot fourd at CORT not HBC. ealinavioa pendiflora Rose, Cone. US. Natl, Herb. 5: 1038. 1809. vinnela pendula (Bowe) Briton & Ross, N. Arar. Fi. 23:69, 1928. 119e Mexeo, Jalsec: Balaios, 10°19 Sop, 1897. J. 9. Rese ‘2024 thestotype here designated, US vo. 301875: fncectospes, GH, K, NY). Calandra maraivata Grech, ex R. O. Willams, EL “Trin. & Tob, 12200. 1; Briton & Kili, Av Nem York Acad. Sei 35% TA7, 1926; LCSrdenas, Revsts Fac Agion(Mavacny} 7. 121 107. ts esa marginate (Gis ox RO. Willa) Mold, Revit Stasmer Bat 6 177. 1940, TF Bek ‘Wot Indios Tina, Cranger 326 uctatype hae designate, Ky eseeotypee, NY, US). To specie Pay ein ore ee Fr 710, 1801 Colluanva gentry! Stanley, Fad Mos. Net. Hist, Bot Ser. 17: 201. 1937; Gentry, Publ. Casnogs Inet West, 527: 122 L042, tyhe Mexia, Chiuabas: Gussaremo:, Rio Maso, Canyon Chanato, wopicel Sonora, 5 Aig. 1980, 4.5. Gentry 2903 holotype, Fe wowypes, A; MENU, MO, 5, Uc, US) fnnesia chapaderoana Bron & Reve, N. Amer, Fl 21 69. 1920 “Tyre: Gamtemala, Santa Rosas Chae ppdero, 5,000 f, Aug. 1H fleyde & Lue 3740 Aicctotyge eve designated, US no. 942577 isle topes). Ancesit roa Mein & Rone, N- Amer. FL 23% 68, 1928. Calligndracooki (Briton & Tose) Sarde, Pull, Field Calunbian Bu, Bot, Se, 8: 277, 1950, yee: Guateuul,, Pei between Yaxiusi, and Yaxhi 9 Ape, 1922, Cool & Martin 209 (ec types hero designated, US no, 1084668, Ameslieizoleoensis Beton & Reso, N. Am Hl. 23: 1997 1928: Loin, Conte, Oeas, Mu. Hit Nat Cle fp "De la Sole” [0:287. (951, 7¥Pt= Bl Soeader ensonate: weit of lao, n thicket, 19-24 Nc. 1922, Standley 21010 (lectotype here designated, NYreleetnypes, CH, US. Anneli rientess Titi 8 Rose, N. Ames: Fh 2: 69, 1928. Calliandra orients Briton & Roe) Ted, Cntr. Oca. Mus Hist. No Cao “De la See" 97, 1950, tyre: Cabs 137 Aug. 1906, Taylor Ui Annestin saberbicutars Msican & Rose, N, Amo. Fl 25: 69. 1928. tare: Mesic, Verureuz Baton del Cerrina, Aug. 1912, Purpus 6084 (leottype here Aesanated, NY; bobetotypee, F, CH, UC. US) Anneilia yoroesisrison, N. Ase, HL 23: 198. 1928, Callandra voroona (Briton) Standley, Publ Fill ohumbion Mids, Bot, Ser. 42 309. 1929. 110 Honduras. Yoror near Progroso, 30m, 13 thice'y 24'lan, 1928, Standley 58093 lecttype here de. ignated, NY, olesttypos, AF, USI. rect shrubs to 3(-5) m tall: stems usally slen der, sometimes reaching 5 cm diam; branchlets NEXTPREV Volume 76, Number 3 1988 Hemandez: Zepoteca 843 generally ascending, sometimes drooping. Pinnse (1-92-34) jugate: patole 2.2-7.8 cm longs rach is 1.76.8 eon long rachilae 2.6-8(-12) om longs leaflets 3-8(-11) pairs per pina, the distal leaflets 1.1-5.9 x 05-36 em, oblong-obuvate to very widely obovate, sometimes faleate, the proximal ones elliptic to widely obovate, the distal mms oblique at ase, usually hecoming acute proxisally, round cd to acute at apex, usually mucronate, the prox lial leaflets smaller. al glabrous to short-pubescent or finely villous; stipules lanceolate to ovate-lan: ‘eolate, somtimes lightly curved, 10(-13) mm. long. glabrous to pubescent or villous. Capitle axillary or in psoudopanicles 40 22 cm long: pe- long at anthesi:. Calyx 2-4 angular 10 oblong. Lneeolate, deeply divided: corolla 4-6 mm long: flaroents whites sometimes greenish white. Pods 10 13(-15) % 11.2) cm, plabrows when enature, but recasinally villous or sparsely villous i easly de- velopmental stages. Seods rhombrid-ovoid ta wily so, 6-7 * 3-6 mm, brown, the aceols usally lighter: pleurogeam strongly’ irregular. Somatic chromosome number 2n = 26. Distribution (Fig. AA). ‘This cubspocies has the widest Uistrinution within Z, formosa, Its range extends from northemn Mexico, in the states of Tamaulipas, Chihushua, and Sinaloa, south and southeast through Central America vo cenral Costa Rica. ln South America it oecurs in northern Vert emuela and northern Colombie, and also in Brazil (Minas Gerais), Bolivia, Paraguay, aud northern Argentina, Ithas been collected ors Cuba, Trinidad, ‘and other smaller islands in the West Indies. This subspecies has not been collected in Panarna, most ‘of Colombia, Feuadar, or Pert. It is relatively ‘common in open sites in eea:onal, dey ensironments such as tropical dry forests coastal thickets, and avid thorn serub, although cecasionaly ite found in wotter disturbed or undistusbed habitats in en evergreen of evergreen tropical or subtropical fr fests It has also been collected occasionally in pine and pine-oak forests in Mexico, Collections come from altitudes of 0-2,000 m. Cativated individale in Guanajuato, Mesieo yiekled the collection Puges Common names. Bellotica (Atlintien, Colom: bia), caba pixoy (Yucatan), cabellera (Oaxaca), chiplin de tote (vaternals), chapuli (Guerrero), claveyino (Chihualua), barb’ montaies (FI Sale vvador), huaje (Sinaloa), ibehe (Yneotin), taman- ché (Yucatén), topetveje (Guanajuato). verancra (Bolivar, Colombia). yeg civ (Oaxaca), yaque (Su cre, Venezuela), ditional specimens eseonined. Mexico, ea¥ Yat alededones de ln Grate de Xvgcunbibeupemy S kn We Dolonetta de Rején, Cabrera & Cabrera {9514 (MEXU® 8 kay SE do Connuas sel camino (Canto Regional Cala en hm 99 dee te Ese eau Chetumal, Cabrera & Cabrera 10979 (MEXU} etorr, Lundell 825 (CAS, F, GH, MICH, MO, RY, Uc, US, Wiss Tuspets, Lundell i072 (F ciyry Mplo. of Tula Gutierren (5 an N ol Tusa Guterer tong roed to HI Sumer, Brendlove 10085 (LL. MO, USk Mo. of Og de Giro, Mexican Livy. 190 athe Zonwcantsn guraje of Multa, Breedlove 36531 (MO, INYj Bip. of Ocorocosuia de Bspinoce, 13-18 ken S of Oconocostla, Breedlove 37244 (MENU, MO}: Myo, a Teiteria, 6 nal S de La Teiitar por eat Fontes Comslepa, J Hemdndes & Chovdn 548 (MEXU, MO Rancho Lindovea, 24 km Ede Vila Force, Miranda 5977 (MEXUx read between Tenia ‘an Vaal, Nelson 3264 (GH, US. Mp. of Oooo, Fear Naneln Muruntk ear Gconogoy Ton 3401 (EL: MO. N¥.'US) camcake Wikgo, Rio Mayo, Gent 2360 (A! MO}: Mpo. Balls, ence Kirte ¥ la Buf Toorio & Romers 9585 (MEXU}. cosh: SW foothill ef the Nevado do Cain. [1.5 ri, of Haciende Se Aerio MeVangh 16] FH(MICH) boeayets Fore 2982 (WS) Clana ato: Moi. Guansjusi, Catade de Pasta, Areilono sn. (ENCB} 10 ken SW of Las road te San Frencisee del Rinoin, Mevaveh 24272 (MIL ‘SUEAREO: Li. SW of Garrapat, Howell 107 (MICH 2.5 mi Nof Acalniztla, Herold ® Clarl $49 (MENU, MICH: Dis. Motes de Oe, Valesios, Hinton 11626 (CAS, GH, MEXU, MICH, NY, US): Teepan El Verde, Hlinaen 14335 (CAS, GH, MEXU, MIGHL, NY. UC, US), Mezatlin, Krnse 1121 4ENCB): La Cor, Langlasee 27TH. G, GH, K, US Dist Aldana, Serre Made de Sur, N of Rio Balas, Temicen, Barrance. de la Jule, Mesto 8922 (B CAS, . G, GH, K. LL. MO. NY, UC US near Lala Rose eal 9617 (USk: Mp, de Tespan de Galeane, EI Pasulmiebe 3 Kn del wetronqe Freche» Coriao am la caer. Tacpan do alee Ze huatanej, Tonore et a. 1337 (MOk Mp. de Zina raj I kan l NE de Zikustanea, por oer. Zn {apele- Ca. Allaminaro, Tenoruo rf 0798 MOK 15 kr AUN se Chilpancingoy Tores 1 ul 1205 (MO 24 1)N de Zunypango del Rio basis Iga, Torres 12 (0) suo! above Lake Chapala, Barkly oa, 7015 (F, TEXY Hacwanca de [a Geum, La Gris, Tonal Eo 5485 (MENU soyes, {12 wi. SW of Al 1583 (MICH, 16 ni, SW Sf Aula, MeVauah 10857 (MICH, 5-8 mi N of Le Cuosts below the pass 19 Talpa de Alene, MeVaue! 120532 (MICH nose Gesdalsjra, Roses Painter #64 USK eer Eatin, Kove fe Painier P526 (NY. US) micsapacan vit af Morea, Arne 5506 (A, BM GH, MG, NY, US} Aula, La Mina, Cueroro ot al 454 MEXUy, Dit, Gakorsin, Sexe Nara, Minton ef ab 15131 (GH. Ly NY, US between San fox de lie Mitanon sac) Amati (Sts, Ans), McVaugh 17950 (MICH aroued Lake Cute, Prvgle 416. (BM, BR, Pc, CH, MEXU, MICH, MO, NY. B, UG, US; Mow tehén, Pringle 2725 (F, ll, SHCH, BIO, NY, US) 1? ri E'af Iiglpan, Waterjali 16404 (US). womens: 6 fra Woof Vautepe, Hunt99 (MEXU, MICH Cuer- Tavace, ose a Roce 17073 (US), Severs Laguna de Stu Blas del Oro, Luna 9084 (MENU) 10 mi, SE of Akuscatln onthe rood to Barranes de Oro aa Aas UUs, MeVangh & Korte 759 (MICH) oxxsen: Date NEXTPREV Annals of the ‘Missoun Botanical Garden ‘Theol, eyfeente de la Vigou Jui, 12 km B de ‘Totelapan, He Hernindee & Torres 210 (MEX. MO, Dist. Poche, of km NE de Checalapa por eas» Fince Monte Gitta, 1 Hernandes € Totes 34 QUEXL (MOL Dist Putle, 20k NE e Pas poe car. a Tesco, Hi. Sernindes & Torres 443 (MERU, MO} Tools el Valle, 10 ti. Ef ety, Kenoyer 1629 (AX, viciiy ‘of Mila, Messer 227 (MICH vrrityof Vlas, Neo 45 (USi near Onaca, Prengle 400! (A MG BI, E CH, MEXU, MICI’'MO, AY, P, UC, US 410 SINE de Tovidin del Camino, Rico of al 251 (RM, MEXU, MOy Moase Albin, Srath 119 (GH, USh Mw be Tunirepes, 27 ke al Edo Rio Vere, Sos orl S2[ 1 OIENU ha Toye, Me a ta, San Agate, on aS de Mao Masques, Sous ef fl 7288 {MO} Des» Babs de Santa Chun 3 he de Santa Crag, Sousa ofa 759Y(CAS, MEXU Dit, Tuntepe a3 kanal N de San Mateo Yell, Souse 7955 CAS. MEXUs Die. se Thecus Row 6 9 hin 1 Ede Toelspan, Sousa & Tees 8501 (MOy Dist, do Mautepec. Mpa Asinesin Tlaollita, Los Tune, 2 hen Al ENE de La Reforma, Souse & Telex 0 (MEXU Dist Zimatlin, Sart Cruz Micepee, Soran eto, 10454 (CAS, MEXU, TEX Lon al SO) te Se Joa de Gri, Tlie: & Magllones 111 (CAS, MENU, MO, NY}, Dist ‘Teiuameyee, ladors W dal Caxro Guingla, Ly Torres 595 (MENU, Formas Acct Miranda 3059 (MENU) Iehind Cerro Colorado between Naceeeslen aad San Ane tovie Cae, ole the Burcanen dee Manos, Sindh ft al 40736, GH, MEXU. NY, TEX, US) quantal Mex hwy. 57,21" N of Queritaro on rod to Son Is Pola, Anderson & Anderson 041 (ENCD, Gil. MICH, NY} Hasina Rivas, sear 1006] (I F,MO, NY, US, FRtin, dreuclies 19 (CAS, MEXU), Penjamo, Magallenes 2672 (MEXU) yorvtena Roo: Meio. Carle Perio, Rancho El Ramona Dura. & Ginsied 694 (MENU: 22 kanal ENE ce Felipe Carilo Puerte, camino « Vigia Chen, Grether ot ale 1977 GHEXU Like al SW le Xela, Grother ev ul. 1327 (HEXU, MO} Gobi, along Da ell Cadell & Lan tell 7846 (A, CL, MIC, US: Capsmaly lan 1? dele fare cone W, Péres & Reamoe 108d (Ul. MUXU. MO). 18 iy al'S do ies Ruins se Tul, Quore & Greer 924 MHEXU, MO}: 3 anal Se Cnt. Sousa tal. 10804 MENU, NY); LagaroCérdensn ene canine Hetco de Kanturdkin sa San Arcofo. Coen & Chan 2926 (NAL) saw tins Pink 13 mis W of Nove Me teks, Keser d Crim 395514). aie nals ety 4959 (CAs, CH, MEXU, MICil. MO, NY_UCy Lae Mess Ser Suiutate, Genny 6356 (6H, MICH, MO, NY}. tanasio: San Paleo Balsme, limite N de le a Ue Hiserva Federal, 8 edo Neranite, Garcia ee Pela 2279 (XAL). rasuuivass Mpia. Gaines Farias, 3 kan NW de Gémee Farias por eamin conto al Remcho (ie) Golo, 1. Herminds € Torses P45 (MENU, MO} 10 gs NW of EI Frogreo which Tk NW ef Ocarsps (EEN, 99BIW), Snort etal 10951 CAS, CH. MENU. MO, NY, UC), Yneuch Buna Visoy Donates eu 1430 (Li, MEXU Mpio, do Ialeomalco, seeder de Iorslee, Gandara € Doranter 100 (MERU, Paces pin, Purgus 8401 (GH, UC Mpa. Zapata, ta baja fre Coral Fab Pialtepee, dav 16 kin al Eola, Sousa & Ramos 4890 (EXO). cats en fos alee shres dee Grutas de Balansonehs 36 hen We Vallis, Cabrera © Cabrera 91) (MENU), Cana aayecy Lk despues de Canlamayecsesnino a Pio, (Chan 1814 (ALR Hurueta, ks 12 carr Homcente & Sil, Chan 2827 (AL Chkiaonat, Neos, Chan 3590 (XAL 9 ko Bo Mou, Comeay & fonsan 490 28 (MO, NY) Chasis, Enriguee 090 (MEXU, US) Progreso, Gaumer 1173 (As Bal, Bt K MA MICH, MO. NY. UC, US). Siti, Ganrser ef el 25410 (FG, GH, MA, RO, US. Kenly Gannon al 23431 (4, FG, Up Donal NW de Tis Siro Tica, Grathor 525 (MENU) 7 han al N de Colona YYoeatin roche de Colonia Yueatin El Cayo, Grether 557 (MEXUs.5 kay ol We Chotian, (ether 109 (MEXU, MO} 3 in W of Valladlid: Johan a Comeoy 441-78 MO} Osesacab, Xu May 74 (KAU) Chichen la, Sveere 1438 00M, F, MEXU): Usa, Soere 2073 (MICH): una, Sioere 2752 (MICH Dees, Scwant 255 (GH Valldshy Ploy, Cane $0 (XAL) Moi. ‘do Yeucahs, al Edo Yovenls, Fare et ul 380 (0, MO, NY). Guvaotia. 4174 taharae:hotwous San Cristal YVerapaz and Caisoy, Stevermark 42011 (US) ci urtesanco: Alemed, Jonaston 925 (Fh San Mastin, Morates 1302 (Fi cngumguts: Cer Brn, vicinity [i Negro, below Montes Momieneprn,Steyermend 30054 9) ct stents: 25 ka Sf Austin, Harmon & Duyer 2987 (F, GH, LL, MO, NY. US} Isiah ‘ein of Jala, Standley 7735048, NY) notes Teck, fntrada de Bajo de Santa Fe, Contreras 197 ty LL MERU}: Dos Lagones, om Ixcwra Rod, km NE of the vila, Contreras 144 (FLL, MEX) Uasgesin, Contreras’ 3505 (F, LL, MEXU Nbilpa. Gronde, Ma anché. Coniroras 5487 F, MO. NY Guayaca, Pha, borkenng Legura Goayacin, Contras 7409 ( LL, MENUS roud tn Fires Tony hlaeor do Meneon 1 ‘hn W ofthe Belie-Castemaa border, Rortos Pak 579 (MO) sicerartyuce ear Tris, Bace de San Lo ene, Standley 59800 (A, F).saNts ROSA at Calpe, Standley °8533 (F)botnean Sata Ans This tal merle of Rancho Licas, Sierra de ln (hichutataney. Srey. frmarh 513570 (F) Betas Inn Cure, e004 to So tal “trnason & Eambore 1762 (MOs El Cayoy Charch T08(MEXU?). HonDvnas, oaaacte:nesr ° Heasleu1979 (MO). cots han se So Montaia Ln Zana, ean det Rio Pras, Male 9423 (F.GHL US) fads de Sac, Juan Lindo” crea de St. Sa Melina 376 (F, GH. US), NowA24s: rear Sach Franco, Rio Yoguro alloy. Williams Motine 1 482 (FGA. UL, MEXU, MICH, MO, UG) otane carr Sen Frarciacn La Pax, Mort Uval, Molin 1333 (B- LLL NY, TS). Fa Savanon, sitacarive Pala 8 (US). cafwansasout La Relea, Caldoran 2408 (8, US). 1a Libs TAb: Santa Tela, epee of Velen Stn Salone, Taian & Molina 19225(0GH) Niesbactn. wouc Las Pas C2e2N, A535"), Moreno 10642 (MO), Santa (Cron 12924'N, 85°49°W}, Moreno 16619 (MO) ans ras a La Poa de Oriente, 1 him doe care S(11S49', 86°08 W), Moreno 10710 MIO} Gonarco El Aguseate; 4 kin of inet, Neu! 1058 (MO), cman: SE bt Cosguina, Forces 146 (MO}, Volein Sus Cratbl, Finca Les Rojos, 20 ken de Chinandega (12°42°N, B102°W), H. Hevadades o al 982(MEXU. MO} fla ‘SE del Vand Casiay sm aniea de Avge (12°70)N, 86°57 Wi, Moreno 1972 M10} La Sua 13 kan al NE eChinandege (124'N, BOW), Moreno 250¢0 MOK kin 136.5 face. Goataule, cemino al Volein Chonco GEUN, 8704'W), Sendino 1457 (MO). coven Puerto Monota N of Aocvapa (11SEN, 85106), Sr ‘eons 10042 IMO} Hda. Say Martin, nar cnuense af io B Jordin and Wo a Pradera (12°17°N, 25°19"), ‘Stevens 22620 (MO) este Sato de Etarzuela, 5 km NEXTPREV Volume 76, Number 3 1989 S Gal de Ese, H. Herniudes oat. 624 (MEXU, MO} 0 kes de Pusble Nuovo cart. « Limay Gacro San Remon (S°20'N, 86°31"), Moreno 3140 (Oy La Gariune (3°08'N. 86"19°W), Morena 2148 (MO), ‘k: Coyaega 5 hav al NE de Conadopa (197228, 1'20'W), Moreas 253359 (MO). chan: Ila Apaver, Cero BI Hane (lalla nore), Grijaloe. G08 (BO) Sa ‘Agustin, camino a Currey 425 km de Granade QGSASEN, 11°S5'W), Bhneno 2639 (MO Volrin Mom, Iirodo por ln nex Lae Debeiax (LI°S1'N, 85°50), More 4089 MO): Ltrs aN de Ca de Toa (11°50, BoOV'W), Moreno 5404 (MO). nurs" 7 han & Is entrada. del camino vio)9 a Tintaga (19H 36°N, BSSHRO'W), Ten & Rollete Joh (MO). a eat. vieja Le hn 68 debajo del pent del Ro Tarn GETEN, SOUNW), Gren er al. 1025 (MOY Co laviera a Bl Trinsto San Pablo (1205, 86841°) Moreno 4841 (MOL sapui: Cerro Queues [13°41N. ‘BOPBI'W), Stevens 22008 (MO waren: Curse de Goyal 12°30°N, 86°08 Wi, Aragitetatn 36 14|MO) San ebro 1205-168, AGL SW), dra qusrata 3054 (MOR Gene sur (12%00-08'N, 85°15 196). draqusstia 3749 (MO Managua, Punta Chitepe, coatado W de Laguna Apoveque, Grpoiea etal 105 QMO) ear. entre ae Conchiae'y Msachapa, Giceidn Casto 1067 (MO} Sen Francaos de Bolan, Méndez $4 (MO car, 4 San Fanwiseo Libre, $k despus dela San Antonio (1228, BoUTW)! Morene 707 (MO S of Villa Franca, Sandivo 34% (MO). along hwy. B en 24 kin SW of intersection nih y= 2 bee 28 (11°57, B620°W), Stvent SOE (MO) txracaton: Ermita do Exqupulag, 20° km NE de BL Tuma (1380°20°N, BSSOSOW), Cryale & Gryatee 1100 (MOE Cane ‘era Panamericana, Cuesta El Verado, 3 kr al E Lan Dalcias L2MHI'N, 8709"), Moreno’ 18366 (MO 1 val E de le Macorda Lin Arden (112, 85°42°W), Moreno 19004 (0k eae. v2) &Jnoteg, as Troncas IIZS9'N, 8558°W), Moreno 22927 (O10, El Tempe furs 3 kam al Ede Poeries Vics (L2935°N, 560076), Moreno 24831 (MO); along hy. 1 at enteance w La (Crom, Bhan SE of Las Calta (1°30 N, BOTW), Stevens & Monciol 18611 (MOP Macao de Peis Blan (os, SE sie, drainage of Quebrada Bl Quebradin, sores Nard W of Hide. Sap Seteniin (13P14-15°N, 65°90"), Stevens et al 2/271 QMO} wuss suconu’ care, & Dk pito'S km al N de Ocoxal Senta Mors (13°59 N, Borde Wy," Moreno 25253 QMO. ZeLavic La Lut CISH'N, a4°47'W), Oris 1589 MO Sina, Seymant 1290 (MO). Costa Ries. ctraacaste: Babi EY Coc, Babie Plays Hermosa (1032°N, a5%0'W), Burger © Baker 042 (Fy. Parque Nacional Sante Rees, Hh Her héndes et af. 098 (MENU, MO} FUNTAAESe Monte Yerda, Haber 392 (MO). oR oxen Rin Vii, Sa Sine, ones 20565 INV} Plots Hcolearice Boa Vil Colin, Poveda 34 (Fy La Caja, Faerio 1315 0) Cup, swanico oe cubs: Seno Bay, Clemanie 2270 {8 Sens Cee 0702 (GHD OS) Fn Be Todo, Clemente 6260 (GH, US); Fines Calimanes, ‘mente 6369 (GH Loma Gilad, Clemente 2030 (Ci Tieopud Hench, Clrmonie 7439 (CH, NYY Bavate, Ek ‘man 0432 (NY en la tona ens el Pace de La Mine 1? la Fxacin de Rocko cre de Punta Gord, Lipes (US bordel amine en Santa Maria se Lovey Lip & Arun 340 (US) Vita cust fas Savon Ean tw Lama de Caen, Zin 3999 NY, US) ist lad of Huevos, Fen 173 (NY), Vveautun, anzanth Tsk Chimana Sepun, Nof Canta L0P18'N, 66°46 M6). Hernénde2 Zapoteca Sterermark & Manara 107998 (US. anycuis dei da Instat de Honea Agel, Facood de Agroronin UCN: Maceeay, Crdents 1929 (F, Naglaleno, Hee NEXT Fans 10803 UF, G, US) anlvaee headwaters of Rio Sace Windack & Guppy 60 (E, NY, US). casa sheledoes de Marra; Rojas 1091 (Pro. Cabelo Garvan de Haman 1144 (A. F. GH, MO, RY. US) ache de Gara, ear Son Josqin, Puser 7902 (F GHLUS) Us Gabor ao Lake Valencia, Pier 12218 CAS, 6, US) onstnaro sina sts between Carscas and ancy road to La Guairs Putor 11019 (A. MO, NY, US), Memos Posinesla Psroguans, La’ Caines, tween Puctlo Nuevo ond Sante na Hretler A363 (MO, NV. UC. US), Charan Valley ioe Estee ial Ta Vitra Breteler 4562 (Fy MO, UC, US} ‘hawoa! Parque Nocona! Gusto, 22 hen N of Alagresa Ae Ontuco, N of te border with Exada de Guat, Yor 17855 (NV Gonrenas Valley, Pitter 12003 (A NY. US) Nuva tsranar le Margatay leg, 2429 (SY). ues: Poero de Hier hae el valle serraias de Paido, drvegutva 4719 (NY, UG, US Pentsala de ‘Araye, 20 km NW of Caio, 2 kan W of corner where ond fom Gavaco to Crocopata see oro (LORIN, G840'W), Licsner & Goncles 12017 (MO) racine Sof La Molata near Venwavelen-Colosbisnboedes (150°30°N, T2379°W), Stoyermark eal 120205 (O10), vanacus: Hacionde Thon near Guame, Pier 51260 (all, NY. US) rou San Martin on Rio dl ator, Piier 10532 (GH, US} Couns. erase: Consevin, Dogan 678 (F), Les Perdis, Hacienda Nioduce, Dogand & Jaranilo 140% (USh Cos del Go, ana de Salat, Dugan 421 (03 Coste del GGarbe, car. Puerto Canin, kv 8, Dagan 6574 GB atari Pos 304 (AY, US) neue Ca Pops, near Cartgina, Killip © Sth 14068 (A, BN, G.'CHUNY, Use Toeeellay near Turbeco, Kili & ‘Sinth 1438 (4, GH, NY, US) oun between Ces teeta sini Caraga, tio Casa, Chatreoas & Costa Fat 35595 (U3) nacpsurne: Bocrry, Hough 3689 (4, USk all 1. Pasbito, Parque Nacional Tayrona G19N, 75°98'W, Kirhnede 3919 MO, NY. Us 1.3 km B of Aesop Batvar and 14 ko of Santa Maria Q1°07'N, 2815°W) Kirkbride 2606 (NO, NY, US}, Mo. de Sums Marte, Don Jaca, Romero 19995 €, MO, NY) wontt be saniasbet Vale dt Ro Poralons! en ios alrededor de Sontiago, Moline Barkley 16 (US)- Bhat. untv0 eno: Corns, Rol e175 (K). Bouin tonsa Signa, C2rdeen 5739 (US). sues unves Prov. Cones, Cai Ca deus 4775 (US Alto Parape, de Michel 125 (NY. Panaciy. Fuerte Olrpo Chars. Rojas 13013 (MO TER) Zniscien Bio Ape, un Bio Anuabon, rel, Fob S308 HM, GH) Ancerins tacos Uae Venn 20298 (A). suv: Due Ls Citegs, Krapont. cher & Criatdoal 17540 (LL, Ve, WIS) sath: Josqun Gonales, anita 252 (NY Dost. do Orde, Crap ‘radar 3202 (GH), Ceo 20 te Fever Fp 255 (U8 10 kn Ede Taraga, camino a Tone Kree porickos & Schinins 39283 (WIS) M4 in Ee Ee Litcaci, Kroponichas € Schinin’ 30871 (Gk. Dpto San Mortn, Gra. Balls, prox. El Slade, Logeame Medina © Cueszo 20149 (Wisi: Panaje dl Ture. trent. Lavente & Heronyris 275 (BR G, NY. UO} Depta’ Ord, Rote 30, Quelead El Caer, Mavuaiak tl 504 (MO, Py Depo, Callers, Mojo, Meyer 135760(GH, NY} El Tebocal a Rio Santa Maria, Meyer 4521 (UUs Dole, Orin, Urinal Rio San France,PREV 846 ‘Annals of the Missouri Botanical Garden Meyer 813 (8, NY Depo. Orin, Bella Vet, Perot: 206 (NY, Cy" Depte. Oran, Tartogal e Las Tabiliss, Setceter 1052 (8-4: Deo, Gonchipas, Aleman Vorsast 9612(A, Ni. GH. MO, NY.US): Deo. Antias, Goro Negro, Pencard 10390 (BM, NY), sunriace Dot tho: Serta de Cumayn, Rata’ 68, 70 Ln SW de Seo dol Batre, Krapovickas ot af 37300 (MEXU) Ficus: Son Pedro Colac « Chulose, Casillin 69572 (ng Dipto,Trancan Vipos, Sehreter 69573 (8, Fh De. to. Borrayaco, Tans, Veanurt 1549 (BB, CAS, Fy CH Mo, NY, 1, Us Dept Tapa, Venturi 924 (4, C45, US): Dept, Borruynen, Cee del Campo, Fen furt 2525{4, BM, CAS, GIL, US) Cuurvarn. USA scouhis Se Les, Masour Botanical Garden, Her Dnéndes 946, 0593, 954 (MEXU, MO) Mexico. Guana justo, Carden of the Alameda, Dugée 1 (GH, NY, US) Specimens not exsigned to sulspecific stats, pre bly intermediate batiscen Z formon subsp. fine fend 2. formona subsp. tose, Mexico: cvanexo: Di Gives, Pangarabate, Hinton 6876 (A. BM, F, CH, MENU, NY, US} Db of Mina, Plaeres Camerin, Haton 9293 (ENCB, Ky LL, MICH, NY, UC 18 kre NE st topic, Jako 1270.80 (MO. NY}: Bite. de 2am fpango del Rio, ky al N oe Mili, cazeIguela~ Epanemgo, Lac ot of, 120 (MERU. MO} Moto. de ‘Ghiparango de lox Brevin, sabre a draw.» Chehalen, 2 Bk al NW se Clponcing, Ea ot af 76 (MEX, MOy Disrito Alama, N of Tl Bsus, Tense arvana de Jola, Mesia 8237 (CAS. GH, MO, NY. UG, Usp t mis We Zarpango dl i, ico & Punk 207 (CAS, MENU}, Zaporoea formosa subsp. formosa is the most 14 and morphologically variable subspe- © species. Fourteen names have bee: published char fal within the boundaries of this Subspecies as [have defined it; most of them have teen based primarily on regional variations in leaf let morghology and vestituro, These characters, along with the size of the lower parts and pods are highly vatiable st an inter- ane intrapopulation level, especially in the Mexican and Central Amoe ican populations, and thus they aro not usful tax ‘onomicelly. Perhaps the most variable character is Teale size, Lut the variation does not correspon twa geographical pattern A sinilar situation necurs with variation in leaflet vestits plants wit ahort pubescent or villous Feaflts cur in the same areas fs plants with glabrous leaflets throughout most of the range. In addition there ix coutinuous variation {a leaflet apex morphology. ranging from rounsed to acute, Plants from the southern hits of this subspecies are comparatively consistont morpho logically. In this area, they usualy have mest to relatively small leallets, nd the nuraber of pairs ‘of lealets per pina ix quite cegula. The entity most closely related t0 Z. formosa subsp. formosa apiears 10 be the Contral American subsp. salvadorene!s, whieh is distinguishes by its densely pile pods. Zapotec formosa subsp. for- moses consistent in having glabrous pods, al though in El Salvador and Venezuela, plants vith clabrate pode have bean collected. In a fow eul- lections of subsp. formosa trom Oaxaca, Jalisco Honduras, Veursuela, and Cuba, the pods are pi- lone or sparsely pilase then very young, but soon become glabrous. Zapoteca formosa subsp. formosa uo appears lovely allied to subsp. rosei, from whieh it differs hy having white or whitesgreen filaments (Table 1), Plants of subsp. formosa ako usually have larger leaflets and aro goncrally more robwst than those of subsp, resei, Ia a number of instances, however, the ilentification of these two subspecies is troublesome if fowers ace not available or if filament color is not know. In some areas of the Mexican Paeifie slope, especially in the state of Guertera, where those two subspecies prosuinably fsceur syrpateeally, hybridization probably takes place, resulting in morphological intergradation. Ia this area, many eolletion® of subep. rosci have larger leaflets thao normal in this entity, thus ro sembling those of subsp. formosa Sach areas of jurailation are in Teepan de Galeana (e.g, Hie ton T4335, 10894, Lark etal, 200) and Zivsota ngja (e.g. Tenorio ot al. 398, Ladd et al. 210, Sousa 6159). Similarly, in sora localities of the District of Yawtepre in Oaxaca, plants fro the so subspecies have been found growing together fespe HL Herndnder & Torres 210, 217, Telles Magallanes 111, 113). The collections belonging cither to Z. formosa subsp. formosa or to subsp. rasel, bot of ambiguous ideatity, are listed sepa- rately under 2. formosa subsp. formosa. Zapoteca formosa subsp. formosais distinguish able from the West Indian subsp. gracales by hov- Ing ablona-okevate leaflets, usually larger, lanceo- late 10 ovate-lanceolate stipules, and. glabrous, pubescent, or villous leaflets, which contrast seth the consistently elabrans ones of subsp. gractls Subspecies formosa differs from subyp. moliwla and subsp. schotti by its more robust habit, white ‘laments and larger petioles, ballets, stipules, Hower parts and pods (Table 4). There is a wile disjunction in the South Amer ican distribution of Z, formosa subsp. formosa from northern Colombia and Venezuela to Bolivig, southeastern Brazil and Paraguay (Fig. 44). Mor hologieally, the populations from the southern fegion clearly belong to the subspecies. Perhaps this geographical fragmentation ie related 10 the ‘carrent scarcity of relatively dry, seasonal, lowland hhabitas in the intervening region, As reviewed by Haller (L982, and ceferences therein}, climatic, fiuetuations doring the Pleistocene had a great NEXTPREV Volume 78, Number 3 1989 Horndndez Zapoteca 847 effect in tropical areas. In lowland areas of the Neattopis, alternating periods of cooler and drier climates probably caused extinetion or displace rent of species adapted 40 the warm-wet forest ‘conditions ding the interglacial pericds, and only 4 few areas (forest refugia) remain relatively we ‘changed. Consequently, ureat expanses of wet for cess were replaced by drier, more open vegetation inthe Amazon Basis (Ab'Saber, 1982; Absy, 1982, Bigarclla & Andrnde-Lima, 1982; van der Hass men, 1982; Peanee, 1982: and others). In light of thi body of idoas, it ean be postulated that the populations of Z. formosa subep. formosa found ‘ore suitable envionment ducing the dry phases of the Pleistocene, and then were continuously dlstriuted across the Aenazon Basin: and that th preseat-day disjunction in South America is the recut of fragmentation of @ formerly moro contin ‘uous range afer the warm, Wer present-day con: dltions were established 12b. Zapoteca Formosa subsp. salvadorensis (Britton & Rose) H. Hern, comb, et stat nov Anneslia satvadoronsis Britton & Rose, N. ‘Amer. Fl. 23: 68. 1928, rvve: El Salvador: Mieinity of Sun Salvador, 650-850 m, 2-7 Feb, 1922, Standley 20452 (eetorype, NY, hore designated: isolectotypes, GH, MO, US). Ereet shrubs to 4m tal, usualy finely to densely villous throughout, except for the dowers. Pimae 2-3,jugate: petioles (1.8-)8.1-.5 om longs rachis (0.9-)1.6-4 ex longs rachillae (1.8-)8.8-3.6 em longs lenllets 4-6 pies per pina, ablong-obovate, oblaque at base, rounded to acute at apex, usually ‘mucronate, the distal leaflets 1.6-2.6 % 0.7 1.9 ‘em, the prosimel ones smaller; stipules lancealate, sometimes curved, w 1.2 em long, Capital eaally auilary, cecasionslly in simple pseudopanicles 10 7 ern long; peduncles 2.2-8,2 em long at anthesis, Calyx 3-4 mam long, the teeth ablongslancealnte, glabrous corolla ca, 5 micalong, glabrous: filannents ‘white. Pods 19 9 = 0.8 em. pilose to densely pilose with sehite hairs, Seeds chorboid-oveud, o. 5. 4 ram, sometimes red when immature, b he arcols lighter: pleurogra Distribution (Fig. 43). Endemie to southern Guatemala and EI Salvador, whoro it has boon colleted in the departments of El Progreso, Chi quimula, Quiché, Solo, and Zacapa, Guatemala, fand near San Salvador, EL Salvador. I has been, collected primarily im disturbed, brushy habitats at altitudes of 300-1,800 m. Additional specimens exomined. Guarewn ‘euguinuc: bscwoon Chiguila and Sta, Roa’, Mele ne & Moline 25136 (F, MO, US) nese divide om cea fom Zacapa te Chisuimla, Stendley 71743 (FB macatsos Motu Valey. Becton, & Breohon 270, NEXT (ENGR, MEXU, NY). quent: near vilage of io Blanco, Walliams ct a! 22456 (BM, F, Ui, NY, US). sovol vilage of Son Pedco, Voleia Sen Pedro, ootitaeing Slopes cowande Lago de Aili, Stevermark 47203 (Pi {scars Bib Hood, base of Sera do las Minas, Standley 73079 (Fy Rio Hondo, Steyermark 25694 Zapoteca formosa subsp. salvaitarensisis closely related to subep. formosa, from which it can be distinguished by having pilose or densely pilose pods and by being villous on all vegetative parts. By contrat, presance or absence of vostture in subsp. formosa is expressed ierogulaely. throughout its range. Geographically, these wo subspecies are allopatric 126. Zapoteca formosa subsp. gracilis (Criseb.) H. Hem. comb. ot stat. nov. Cal andra gracilis Griseb., Mera. Amer. Acad. Arts & 180, 1861, nos Klotasch, 1871, nce Standley, 1929, nee Renvoize, 1981. Anas: fia grisrbachii Wetton 8 Rowe, N. Annes. FL 23:07. 1928, This was published as a nomen, ovum, bocause th name dmnoslia gracilis was preoccupied by Britton & Roce for a species based on Acaeia gracilis M. Martens & Galeotti Gee synonymy under Z formosa subsp. rose). Callianuira grisebachit (Briton, & Rose) Standley, Publ. Field Columbian Mus. Bot, Ser. 3: 277. 1930. €. grisehachii (Brn. ton & Rose) Leén, Conte. Ooas. Mus. Hist. Nat, Colegio “De La Salle” 10: 237. 1951, Liogiee, Fl. Cuba (Supl). 72. 1969, rye: Prope Cubs Oriental, Sep. 1859-Jan. 1860, Wright 151 pro parte (lectotype here desig: nated, CH. isolectotypes, CGE, F,G, LE, MO). As Len (1951; soe synonymy Below) pointed out, Wright 151 includes two elements under the same number. In addition to the species ronsered here to represent the type of this taxon, there are five collections, also num: berod as 152, at G, GH, LE, NY (lragmont) and 5, that presumably came from another locality and belong to Z, formosa subsp. for ‘mose. One additional specimen at G inclines om both mater Caltiandra comata Gresb, Cat. Pl, Gab, 83, 1866, non eth. 1846. Calandra jormesa var. cubensis F.Dfarbe. Cone. Geay Heb 59. 4.1919. (Based Tlandra coms) Annesin (sce) Briton & Hine, Ne Amer FL 24:68, 1999, Calindra cubensis Qc. Mactr) Leb, Conte. Ocso, Maw His, Nat Colegio "De Le Si" 9:7. 1950; Lain, Conte, Osas Ms, Hit Nat Coleg "De La Sais” 10: 287-198 1: Linge, fxbensis JFPREV PL Gaba Sup). 72, 1969; Proctor, Kew Bull 11 ABI 1904, Type: Cubs Wright 3406 (lstoype here designated, GH). Erect shrubs to 2-5) m tall, glabrous through cout, Pinnae 2-B(-H)jugates petioles 1.3-5, Tongs rachis 0.7-5 em long: rachillae 3.5 Jong: leaflets (5-I7—15 pairs per pinna, the median ‘nes eliptc to elipticablong, the distal anes oblique at the base, the proximal oes becoraing acute, roused to subacute, rarely acute at ape, som times macronate, the medivn leaflets 9-22 % 3.5— 11 nm, all usually drying Hight green stipules Tinear-triangular, to 5 ram long. Capitulo axillary: peduncles 3.5-19.5 em long at anthesis: eal) 2,5-3 mm long: corolla 5-6 ram long; laments “white, Pods to 11 x 1,2 em, glabrous. Seeds ovoid, 5-6 x 3-4 mm, usually mottled, pleurogram se nicegula. Distribution (Pig. 44). Restricted to several islands of the West Indies: Cuba, Bahasas, Cay- rman Islands, and Haiti, where it primarily aceurs im rocky, caleareous arcas in dente thickets, and, in genoral, in secondary, arid hebitats at elevations ‘of 0-600 m. Additisnal specimens examined Baas Ane etycn, anwens Fight Mie fay, race 1854 (F, NY) ‘scum sano: south ea, Titan 80254, NY) sso: fod to Merpu's Bi, Brace 6631 (F, Vs Behring Poin, Codirey 41424 (SVs Soutn Andros, 8 of Litto Grech, Correll 13431 (NY Bm. Keak Andeos Town sirport, FAIl 3005 (NY: Mengvove Gay. ewe Lisi rec, Smal! Carter 8450 (F. GH, NV. US). ener Iiawis Wha Cay, Briton & Milopaugh 2189 (8, RYp: Goat Cay, Bruton & Bilypaueh 2288 , NYY Anerson Cay, Grou! Harbour Cv, Correll 13087 (F, RV. must uahpe South Bimini Correll 2174 ( RY: fhmin, Cat Gay, Corarll 45614 (NY Soath Ba NE corner of bland, Gill 17205 (F), exons ibasbe Tong Cap, Neni, Mllopameh & Milspmugh 9255 €, AS, NY: Ling Gay, Sond Milopaue & Milopaveh GEE, NY, USy South Caices ison 7090 (F. CH, K. NY) ca BiAND: Hennes Harton, Bye 138 (A WIS) Zone Hill Creel! 46005 (NY) wast: Glas Windsn to Harbor Ian, Brivon & Milspough 3302 (F NY North outhers nace Watkin Hill, Correll ‘Hlo60 (MOY 6 mie N of Jain Coton, Correlt Hil “#5244 (NY) EM ca Lat Gast Cay, Briton & Milypagh 2643 WF. GH, NY. US) 6acv: Great fps Rocky Bont, Croll 4747 1(M0, NY). ao et Eitvence Town, Briton & Mltypangh 6257 (F, NY) sen enoibenc: near Nason Curtis 83 (A, FG, GH, MO, NY. UBy Bloe Bile, betwen Lake Cunningham and ke Kilarney (1725.5, 25085}. Webster et a 103104 (GH, US): Hog Ward, Wilson 8272 UF, NY). gece ani: Great Ragaed Ian, ison 7000 (P, NY) ross saws Bruton Milipeagh 2149(F, NY) ayia Beant pear Tibet's Tuer Procior 29355 (LL 08-1 km inna from Spot Boy, Proctor 35139 (US) Ging: canantey: Cayo Satnal, between Carte F ‘ote and El Fueste, Pena £5339 (US), ln Gari, near ‘Annals of the Missouri Botanical Garden Laguna Grande Shafer 802 (GH, NY, US Santa Lea, Shafer Wi4 Gil, NY. US Cayon Canal Nuevo, Cas Tagun, Shajer 2660 (F, NY, US), Cayo Guajobo, he NW ead, Shayor 2855 (FH, NY, US} vikcty of Tiini, Shafer 2001(F, NY, US) curve: Pama Peds, Dige Bay, Briton etal 12471 (NY, US: Rin del Moe, Iauco region, Leon 12426 (GH. USk Lane de Tabs, Mavi hy, ee & Alain 18007 (CHL US) Hin to Myabe, Shafer 1410 (NY, US worhnand of Rainn, Nive Bay, Shofer 1702 (NY). eat ous lo: Pas de Glnjabn, La Mulata, Barker 18313 (NY Pen de Cus fmm sen & ewa 2005 (NY. Surta de Ne. Rio Pikte, Ekmn 5780 (Gj: Bones, prope Porto Ries, rain 607 (NV). Hart le de NordOues, Port de Pin, Floor 2856 (NY, US Ful Tt Vallee Torte Isa, Leonard & Leonard 11312 (Ay GH, NY, US). Corre ‘arto: US.4. ruontDe: Biems FacchiTropicel Carden, Gis 3072 0, Zapoteca formosa subsp. gracilis is relatively uniform moxphologially compared with the other subspecies of Z. formosa, [vis velated to subsp. formosa, from which i ix distinguished by having ‘elliptic to ellptiewoblong leaflets and nayrowly tr angula stipules and by boing glabrows (Table 1). ‘The soods of subsp. gracilis differ from those of the remaining subspecies of Z formosa by having seminegular, instead of irregular, pleurograms Subspecies gravis and subsp. formosa aro essen tially allopatric, though there are a fes collections ‘of both from Oriente Proiner, Cabs. tionship is discussed further under Z, formosa subsp. forms Although leaflet morphology i# quite constant among populations of Z formosa subsp. gracilis, 4 fow plants from the Bahamus (e.g. Correll & Correll 45614, Curtis 82, Gillis 11298, Hill 3003) have unusually narrow leallots, These plants coukl Deconfused with Z. portoricensis but ean be read Iy distinguished by thee smaller and narrower tp tes and relatively thicker leaflets (also sce Table 14). In addition, Z. formosa and Z. portoricensis ace strongly segregated geographically v0 the West adi. 12d, Zapotecs formosa subsp. roxei (Wig fins) H. Hern. comb. nov. Calbinndea rosei Wiggins, Contr. Dudley Herb. 417. 1950. Calltandira sehotti subsp. reset (Wiggins) Felger & Lowa, J. Ariz. Acad. Sei, 0: 83. 1970, Acacia gracilis M. Martens & Gal ott, Bull. Aead. Roy, Sei. Bruxellos 10: 311 1843, snneslia gracilis (M. Martens & Ga Teott) Briton & Rose, N. Amer. Fl. 25: 66. 1928. Calliandra gracilis (M. Martens & Galeot Standley, Publ. Field Columbian Mus Bot, Ser. 4: 309. 1929, non Grisch 1861, neo Klotzsch, 1871, nes Ronvoizo, 1981. H. NEXTPREV Volume 76, Number 3 1989 S. Gentey, Publ. Carnesie Inet. Wash. 527) 122, 1942 type: Mexico. Oaxsea: bord del POcéan Pacifique, Oct. 1840, Geleorti 3190 holotype, BR (nt seen); boty pes, G, K, LE}. Galine socorrente Mobs, Cts. ray Her 70: 71-1926. drasslia socerrensi (1-8 haat) Bacon 8 Rose, N- Amer F233, 1928. Te Meni. Colima: la Socoro, Mar Jone 1897, We anihony so. thltype, UC a. 88544). Ins Inter ore Hoban Proc Call Acad. Se, 20°62 TOS.) pated out that Aalny made two urn Ive ales on Socorro skin. Sine he mpd shit they ware med> fore deforentIoealte the econ colaction abo at UC tao. 89254), auld ot be consiered an saiype: Aone lento Beton © Rove, N. Ser, Fl 23: 67. 1928, ye: Mento, Baja Cakforia Sur: San Joos del Cabo, 28 Sep. 1890, TS: Brandegee 196 Acteiype, NY, Bere dosgnated:islecteypes, 8 CBLRY, Ue cs, US) Annes iaerior Brin & Rlwe, N. Amer. FL 23: 66. 192 Type Mexico. Durango, munis of sou ‘en Duras, 15 Aug, 1897, Rore 2317 Wecarype US no. 301227, hee desgnsted solocttype, NY Shrubs to (3) m tall, stems usualy slender, erect or drooping. Pinnae I~-jugates petioles 1 $.2(-5) em long; rachis 0.33.4 em long, rechillae 1,9-6.5(-11.3) om longs leaflets (3-)4-12(-17) peirs per pinna, distal leaflets ohlong-obovate to ‘oblong-very widely obovate, rarely oblong, the proximal ones sometimes becoming elliptic. the ds lal leaflets cblique at base somotimes becoming acute proximally, rounded to acute at apex, usually imycronate, 8-31 x 3-21 mm, wsuslly becoming smaller proxially, glabrous to villous stipules lan ceolate to ovate-lancealate, to 9 mim lng, glabrous to villous. Capitula axillary, some imple peeudopanieles to 13.5 em lang: peduncles 14. (5.8) cm long at anthesis. Calyx 1-5-2 mm long, elabrovs, the teeth triangular to oblong-lanceolate: corolla 3-5 mm long, glabrous: filaments red-pur ple. or white in the bacal bulf end rod-purple oF pink in the distal half. Pods to 9 = 0.9 em st ‘maturity, glabrous Seeds rhoiubeid-ovoid to widely thomboid-oveid, 4-5 % mo, brown or gray, the areola lighter usually, pleurogram ievegular. So matic chromosome number 2 = 26. Distribution (Pig: 44). "This subspecies occurs slong the Pacific slope of Mexico, from northern ‘Sonora south and southecet to the Isthmus of Te Ihuantepec, It has alo been collected in southera Raja California and on the Revilagigede Islands (lsle Socorro and Isla Clarién). A number of eo) Tections have been made fron the districts of Yau topes and Tlacobala, Oaxaca, and fer ff loealities in the Balsas Bosin. It is relatively common ia disturbed or undisturbed habitats of Hernéndez Zapoteca 849 tropical thorn woodlands and tropical deciduous of tropieal semievergrcen forests, 1! found om step calcaroous slopes, on vleanic rock, or along, the ‘margins of soasonal streams at 0- 1,430 Common memes tosapalo (Son073) Husje de caballo (Oaxaca), Addiional specimens examined. MFXICO: ts et IFORNSUN: 6-10 au. SE of Tdka Santos Cals San Tnons hyo nll edo Rancho La Darers, Breed. love We Akstrad 41273 (CAS, MENU). cara Mp, at Arriaga long Mea hey. 195 near Monte Bonito, Poa 3405 (C48, MO, US) clwwames Mpa de Batolae, Bo! La Buta, near Arroyo Ia Buta, Bye 7720 (MEXUY SW Qua, Peter 297 (UM, Gl, K, MERU, NY UC, USp La Janta, 18 kau NE de Basin, Tenorio © Tass 4545 (MERU). cotiuse Sooree Ia, athony gn (UC Revilageds Telande, Claréa Telesd, above Sulphur Bay, Howell 2371 (CAS, GH: Reville Isles, Garde Ln, Mason 1506(C8S, GM} evi ‘ge Isles, Mason 601 (CAS above Bo Sada Fi Sol Colina, MeViygh a Roe: 1000 MICH} cia famine Evert, Ia Soot, Miranda 8732 (MEXU Vennanas, Mrnnda 6775 (MEXU Mansanilo, Paiier 06 (CH, Ky MICH, NY. UC, US 19 kam ol SW de (Clears Monro, Sota & Covi 2571 MEXUS slong Colima ade of Rio Tuxpan bekow hy. 110 beige, Stevens & Fairhura 1834 (CAS, OH, MICH, MO, TES) "stu: Pied le Cesta near Aeapaleo, Carlion $051 (F, US 14.7 mi N of Chipaniages Dan etal, 20519 GENCE, MO} Dise. of Copura, Pangaea, Hinton ot al 6966 (A. BM. 1. GH LL. MO, NY. US), Disc Gakana, Teopan, Hinton TORO {BMF C, GH. K, LL, MEXU, MO, NY, US} Disc, Calene, Moyes, Hinton 114005 (GH, TL. NY, USx Mpo. de be Uinony car Triapa-Ploya. Aza, 3 tal SSE de ln den. 9 fo Un, Tn of 22 (MERU, MO) Lx Povnea, Laagtesse 35 (6, GH, K, MEXU, P, US) Igual Capon, Primate 10057 (CAS, & GH, LL, MICH, UC, US, WIS} Achotl, Reo 1916 NY, US pio. de Zamgango del Ro, Case Werde, corea se Xeshpala, Reedovests 160417 WAS, TEX), 10 kin al NW de Petan, pore camino al Ca tualte, Soto oF el 3967 (WIS). Bia de hasten, Sins 150(CA5, MEXU, MO, NY}: 5 wi 5 of Ica, Weiser eal 17MZ21 (OM, MEXU, NY, TEX, US) sisi 8 kn dela Boes de Forstn, cerca de Puerto Vallarta, Delgado ot 1 139(C Mpa. Le Huerta, Estrin Chamela, M diernd Lett 250 (MEX, My ills Borosen Barts Se Navidad fan La Manuals 9 Bahis Teneatta, MeVaugh 20988 (MICH error af N de San Patric, Peres Pree 1256 (MEX carrera al Cazsemate, Pog 2 (ENC Corcovado Canyon, I is NE af At ea Goalie, Withur & Wilhur 2385 (MEXU, MICE, Eticor Dist. of Tmascetpee, Tenavae, Hinton $183 (CH, K, Uj Dis. of Tenascalepre, Aeatilan, Hinton (6995 th G. CH, K, LL, MICH, RY, UC, US) Dist of ‘Terascaltepee, Patanal Hinton 7094(4, F. MICH, MO, UG. US) aieiosesn= Agua, kan al We lg Cour de Cachin, Guerrvo er al 1012 {RAL Mpio. Jrgspeo, re Aun Banca ¥ Lézaro Céedena I Hernandes (Chacéa 185 (MEXU, MO} Dat. Coaleomasy Vile Vie tor Hinen 12558 (BM, CAS, F. GH, MENU, NY UC. US) Dist datseaoro, San one Pari, Hinton 29055) (Mi, CAS. FG GH, Ky MENU, MICH, NY, TES NEXTLUC, US, Hetranea of to Canrita atthe beige 9 prey SE of Apstangin, Mevaagl. 17974 HICH) enire FL ‘Quoiaile y Arero Benen, Tuzla, Modiano cab Sond (MEXUn Mo, Tigucheo, Agta de Ine Coy Sree La Crovesta ye Goya Soto 199 (CAS, MERU Sip. de Anteage, en cerry ake 4 nerio, Soro 4 Sion 1636 MO} care Citro Cominene Paya’ An ‘Pam de Arteage, Soro & Sur 1099 (MEX, TE Wide 7 kal do Arteaga, Souse & Soto 4033 (CAS, MEXU), Mio. de Mirtamo, Purr de fs Cian 6 eval Ede Murtaio, ear HosanwZatéeuao, erie frat 1548(MO) sores: 10 kn de Tel, Ges Poripa 1058 (MBXU. Caro Aealipn, Péaques 2046 (MERC) naxuca: Dit. de Tertepees de Roo Verde a Iuuiepee, Conzess 4994 (Uf ntepec, Fier 3497 (EMERY. TS) Mp. Totolapan, 9 ke Es Fegan por carta Tekuanepec, Ht Herndndes & Toes 307 HERO. Nos Mpin. San Moréo de Mar, 13.8 bn NW Sin Mote del Mar Heras: Torres 209 (MEXU, "Yauepec, 17 in NW La Reforma po cae WE Tiemindes Torres 210) MEXU, MO Mio. Nejpa, 3 kn Nefapa de Madero por camo fhe estmcs tn care Ooxaea Tasuantees, Her ‘hindes & Torres 212 (MEXU. MOR Dist. Toei, ttre fan Jone de Gracia y Totlapan. 2, Herndndes Toros 215 (MERU, MO}, Ditr Moola enlrente de ta Virgen Juni 22 hin Ede Talpan, ff erodes (€ Tovres 217 [MEXU, MOY Dist Jvehio, 3.3 kin E Le Ventona por caer, Tapenstpos, H. Herndndes & Torres 117 (MERU. MO} Distr. Tessie lp del Cerro Ginga, erent E, nl ented ian Ries Acgumigits, H Heratades & Ramirez 928 (MEXUE Distr Yautepee 26 kn def desc a San Barolo autopr orto de Son Batol Yautepec) deb eae. Oasace-Tehuastepec, H. Herndndes & Romie: 990 (MEXU, 29 km de San Jose de Gracia, soe can. Ossare- Tehuantepec, entre San Jou de Gracy San Joan Gucgosacha, Fl Herudndes & Renirez 952 (MEXUs Disc. Talla, 9 km NW de Totlepan por farra Orxoen H Hernindes & Roinies 943 MENU Zipolte poe el canine al arroyo, Shapiro 236 (CAS, MEXt, [G5i Dice Caan, Tomelln, Sous. etal 6246 G0 Dis. Juul, 7 lam a NW de Bo Grande, Sousa oral 7079(B, CAS, MEXU, MOx Ds. Juqule, km al WNW de Po. Eeonibio, Sousa eto! A400 (CAS, MERU, MICH. Dit duchiin, Moe de San Mid Chimaipa, Visa Teron, 3 tan a NE de El Parvo, Souse 8907 (GAS, MEXU, NY} 2 ka al SW Ga Sen Jost do Cora, orea de Le Cant ear Os ace Tehuantepec, Teles Angles 123 (CAS, MEXUs | km al SW de fl Goaclss a 21 an al NE de EN Gaon, Tlles & Nogatioes 253 (MEX) Cerro Trew Cruces, al S de I Lin cual etd U1. kin AT SW cel encangue de carr Tekusmepee-Onxoes-Boc- f= Ales Terres 4202 (MERU, MO). stNatox: Bahia Toplatampo, inthe Siero Novscit, Gentry 1 460 (CEEMXT MICK cern de Fuerte 18-34 1 Nok lw Mochi, Geary 24527 (LL, MX, TS) Matin, Ortego 7305 (CAS, 6 K, MEXU, US} ED Qa, 34 ni 5 cf Bora, Skorepa a2 (MO} Mpio. de. Reso. ‘imino de Heradra H1 Alano-Carcy Yauco, 6 nal INE el Ejido debs de Ponce. Tenorio a 2816 (S10) seote Catan de Nacapties ct 6 bra NE Bahia San askin Polen of 82124 (MERU Enso Grande (Gan Pedro Bay 0 3m. Efeom S ead of Bas, Felgor feat, 11579 (MEXU): San Boreacde, Rio Mayes Gerry 1690 (AL FB, LL, MEXU. 1D, NY, UG, US), Cerro Annais of the Missouri Botanical Garden del Viejo SW of Caborea, Gentry 14459 (US Sera Bojliuacame SE of C4 Oberg, Goncry 1488 (LL, Usp Son Barnardo and weinity, Arzoyo Gochico, Gentry et ak 19280 (US Hacon:e de Sax Mugua, near Dx Apes, Padoer 16 (0M, GH, Ky MENU, NY, US) This polymorphic, highly variable subspe ‘long with Z, formose subsp, formove, represents the phylogenetic core within the species, Leaflet size is highly variable within and between popula: tions, ard inal variation pattern is evident, especially in the northern portion of its range. possibly a refletion of deereasing precipitation from South to north, There & some variation in filament calor within Z. formosa subsp. roset, although this character is consistent at the populational level ‘The distribution of the diferent color morph i= not known with precision. The following genera ation can, however, be drawn: the cost wide- spread pattern of filament color is the white/red- prrpe patra, which curs fem southern Oaxaca to Sonora and Chiusbus. The populations fem the Ishi of Tehuantepoe, at the easter ati butonal init (eg, HL Hermindes & Ranives 918, Sousa & Filles 8687), ond afr poplatons from Michoacin (e., 1. Herndndes & Chacén 185, Soto 1636) have entely red-purple fl rents. Populations from the distits of Tacolula and Yautepec, Oaxaca, have white/pink filaments (evga H. Heradndes & Ramires 930, 043 “Zaporeca formosa wep. rset inclrely elated tw malay. formowa and to subsp. mollicata sl subep.sehoti It can be distinguished from subsp, formosa by the color of its laments aod by is usually smaller leaets. However, plants of these ‘ova subspecies sometimes resemble each other closely. Tn some localities of Guerrero, the to subspesies hoenme syrapatsio and apparently in: tergrade morphologically. These celloctions are not assiqnod 1o a particule subspecies; they have been listod separately under subsp. formose (and see discussion under this subspecies) As will be diseussed below under the respective subspecies, paierns of morphological variation, es peciully in leaflet size, are essentially cont hetwoen Z. formosa subsp. rose’ on the one had and subsp. molliewla and subsp. schodtit on the ther. ARhough each of these i relatively easily ‘deatifiable, their integrity breaks dosen breause of the existence of moephological intormodites. Za ipoteee formeca subsp. roset ean be ditinguiched from subsp. mollicula by having more-develaped Teaf parts (64, longer petioles and racillae, vst ally more numerous and larger leaflets, and usually larger stipules (Table 14). Similarly, subsp. rosed an be distinguished from subsp. schotti primarily NEXTPREV Volume 76, Number 3 1989 Dy having more numerous and larger leaflets (Table 14), Nevertheless, the differeneas hetweon these two subspecies are les marked than those sepa rating subsp. rosed and subsp. mollicala, Geo Graphically, subsp. mollicula and subep. schottit sre peripheral to but contiguous with subep. ravi ‘This fact, slong with their cloxe morphological af- fois, suggests that subsp. molficuda and subsp. Schott were derived from subsp. row’. Further discussian of the presumed relationships of these three taxa is provided under the respective auby species 12e. Zapoteca formosa subsp. mollieula (M. Martens & Galeotti) Il. Hera, stat. et comb. nov. dearia mollicula M, Martens & Galeoti, Bul. Acad. Roy. Sci Bruxelles 10: 313. 1843, Aanestia moliicula (M. Martens & Calectti) Briton & Rove, N. Amer. FI. 28: 64. 1928. Callinadea moliicule (M. Martens & Galeati) Standley. Publ. Field Columbian Mus,, Bot Ser. 4: 309. 1929, vrs: Mexieo. Purbla: Phine de Tehuacdn, 5,500 ft. H. Galeotte 3216 (lectotype here designated, BR). Caltandea unijaga ose, Cont, US. Nat, Herd. 5 198. 1809, dmslia wage (Rone) Beinn & Rose IN. Amer F-28:64, 1028, 1YeH: Blanco. Oaxten Ccicatlan, 1,800 f:, 26 Sep. 1894, LC. Sith 203 (lectcryge hore desgraied, GH: toeetotypes, US 2 Fragments Caltiansea ungugn var. pucblensis JF. Mabe Cots Gray Herbs 59. 4. 1919. s90t Monto. Poel vty of Sen Las alidanapa, war Cavaea, Ba. FancsdeTlaeuieso, Jane 1908, C4 Purp 3185 Alecttype, B, haze designed botypes, BM, CH, NY, MO, UC, US 2p, Breet shrubs to 2(-2.5) mv tals stems slender, sometimes with exfoliating bork; branchlets pilose in the younger parte, glabrate, ometimes glabrous ‘broughout. Pinnze I(-2)-gates petioles 3-L} ram Jong; rachis 4-8 cna long; rachile 6~ 18 men Tongs Teaflets (2-)3-5(-6) pps per pinna, oblong.che foxate to wilely obovate, rarely elliptic, the distal ones oblique at base, 4-L4 2-9 miu, the pro: Jmol ones smaller, becoming rounded or acute, rounded or rarely retuse at apex, usually muere: nate, all membranous, finely pilose oF villous t totally glabrous: stipules lanceolate to ovate-lan seolate, sometimes narrowly triangular, t0 3 mm long, finely pilose to glabrous. usualy elite, Ca pitula axillary: peduncles (1-2-)2.3-1.5 em long at anthesis, Calyx a, 2 mi long, the teeth tric angular, glabrous: corolla ca. 5 mm Tongs Filaments ‘shite in the basal hall, ced-parple im the distal hall. Pauls to 7.5 > 0.8 em at maturity, glabrous, Seeds ovoid, ea.5 > 4mm, browns pleurogram icegular. Hernandez Zapotoca 851 Distribution (Fig. 44). Endemic to the Te Ihuaedn and Cuieatlin valleysin the states of Puebla NEXT ad Oaxaca, whero i primarily occurs on dry. cealearvous slopes, in arid serub and tropical dry forests at 550-1,450 a. Hull (Poet). Common name Adduional specimens examined, MEXICO Pub Indo W de Aone, Chane of ol. 2459 (MEXU Con callin, Deigndo 45 (CAS, MEXU, Tebtocin, Rose 0 fat 8S (NY. US. Zacoiin, al SE de Zapories de fim Salinas (LB20'N, SP26W) Zara 109 (XALs txses: Teewavien, Cotcutt 41 MENU. US} Coie Atahusea, Le'Hoera, 29 kan al NE de Tepelmeme de Morel, Craz 2565 (ENCE, MENU, Dut. Cuca. Tomelis, £1 Horadnies & Torres 218 (MEXU, MO) Distr. Cuicalén, Toren por care. Santa Main, 1. Hernéndez & Torres 221 (MEXU, MO} 7 kon N Cie fatlan por caer. a Testilin del Cao, 12 Honaindes {& Torres 226 MENU, MO}, Distr Toon, B bea NW Teconvaca grea. Toot, H. Hoses Pores 2IMMEX, MO) Dir. Teta, kn Sle Teeoravacs por care Cuicatln, H. Hernéndes & Ramos 87 (MEXU, Diss. Toot, 1 Lin Tomolin por ol camino lb cart Cueatlan- Toot, Ml Hernandes & Ramircs 870 (MEXUs Dir. Teor, on el sesomboque del ‘amino vio ¢ Tomelin, sbrela ear. Cicalsn- Oana, Wh Hernbndez & Ramiree 87? (MENU) leros al SE 4 Cusaran, Miranda 4577 (MENU Caieatlin, Neds YoaN(Us omelin Canyon, Rose Hough 4671 (US), Dar, Cuicatlan, 15 hro'S de Donogle, Reeders 94957 (ENCD, Caf de Tomelin, Souse ofa! 6008 (OM, MEXU, $10) Zapotecs formosa subsp. mollicula ropresonts fan extreme of morphological variation withic the species in petiole length, number of pairs of pine, land namber and sizeof leaflets. Its distinguished from the other subspocies by its 1, rarely 2jugate Pinuse with fener and smaller leaflets and by its omevhat shorter internoles. Leaflet size, however, is extremely variable in this subspecies, both at an intra- and imterpopulational level (eg, Cras Cie eros 2563, H. Herndndes & Torres 213% Rose & Hough 4672, Sousa ot al, 6904, 6905). Sim iacly, vetiuce i highly vatiable, and does not dlicplay any coherent pattern Zapetces formosa subs. mollcula elearly is closely relatad to the highly variakle aukep. rose Perhaps the most taxonomically meaningful char acters separating these taxa. ave the usually more reduced leaves of subsp. mollicula, However, varie ation loaf characters (0... petiole length, nucnber of pinnae, number of lafiets, sie of leaflets) is esseatially continuous, expecially whon plents of sulwp. rosed from the districts of Tlaeclula and Yautepec, Oansea (ex.. Dejgasto et al. 724, Ht Hernindes & Torres 207, 210, 212, 215, 217, Hesndndez & Ramirez 930, 932. 943, Magatla: res et al. 199, Sousa et al. 6549) are comparedh some plants of subsp. mollirula (eg. Cruz Cisneros 2563, Hornéndes & Torres 218, Her- inde: & Ramive 874, 876, 877, Rote & Hough 4071, Sousa et al. 6904, 6005). PREV 12F. Zapoteca formosa subsp. schottil (Tor- rey ex. Watson) H. Hem., comb, nov. Cal- Liandra schow Tortey ex S. Watson, Pro. Amer, Acad. Arts 20: 364. 1885, “Callian- dra portoricensis Benth. vae." Torrey, Emo: ry Rep. 2: 61, 18595 Isely, Madrofi 21; 280. 1972, This species vas doubtfully referred to in Toreey*s Botany of the United States ancl Mexican Boundary Survey loc it.) as “Cal Viandra (portoricensis Benth, vat.)." Subst quently, S. Watson hosed his Calliandra “choition the description provided by Tore siting, a Pringle collection (272). snneslia schon (Torrey ex 8. Watson) Briton & Rose, IN. Amor. Fl. 28:67. 1928, Callianden scho' 1 subsp, schottt (Torrey ex S. Watson) Fel= ger 8 Towe, J. Ariz, Acad. Soi 6: 83. 197 TyPe: Mexico. Sonora: Pimeria Alta, Sierra Verde, Arroyo de los Samotas, 20 Aus. 1855, Schott sn. (loetotype designated by Tely (Ma. ‘rojo 21: 280. 1972), GH: tolectotypes, NY 2). US (ragment). shrubs 10 1,5{-2) mam tall stems slender, densely branche: branchlets glabrous to finely pi se. Pinnae 1-2-jugates petioles 1.23.5 em long. rathic 0.41.7 emlongs eachillee 1.94.5 emlong Leaflets (3-)4-64-7) pairs per pinna, ablong-o ovate, 12-18) x 2-6(-7) mun, oblique at bese, rounded to acute at apex, usuelly mueronate, mea branows, glabrous to finaly pubescent: stipules nar. rowly triangular to lanceolate, sometimes chghtly curved, to5: mm long, glabrous to finely pubescent Capitula osilarys pesmeles 1.8-4.5(-6) em long at anthesis. Calyx ea. 1.5-2 mim longs glabrous, the eth tiangular-loneeolate: corclla ea. 3 mm long: filaments white in basal ball, pink in distal half. Pods to 7 > 0.6 em at maturity, glabrous, rarely very sparsaly and minutely pilose. Seed thomboid-oveid, 5-8 x 3.5-5 mm, brown, the areola lighter: pleurogeam ieeqular Distribution (Fig. 44). Enemie torueky. arid slopes in a few loealites ofthe northern portion of the Sonoran Desert in Pima County, Arizona (6.84 Biaboquivari Mts, Santa Catsliaa Mts.) and north fem Sonora at 1,100- 1,400 mi, This subspecies vecupies the nerthemamest part of the distribution of the species ‘Annals of the Missouri Botanical Garden Auddivional specimens examined USA, sn20%s Puna Co. W of Tacs, Robles Pas Bortrom 409(US) Coyote Mountains, Gooding 2457 (CAS Monto an NEXT Yor. Dabogicar Moomins Gooding 241-45 (AV Beboguivart Canyon, Baborivani Mocstains. Gould Hoel! 32535 CAS, F, GH, NO, Nv), Colina Moun tain, Sabin Caron, Jones na (CAS. Fo Ul MOLNY UG, US), Toro Canyon Baloguvar Mountains, ewe) & Becton 10192 CAS, MICH) Soh Caryn Keane) E Prctcy 14960 (CAS, Gil NV, USES in. bees Foseral Pron Car Heaters Soca Catalina My Porker ef al. S81 Gil, NY, UC). Mixx sos Sera iu, Gontry Weber 27985 (A, US noth western Blin Pringle 272 (S177 me SE of Mag Salen Star Brgy 169 (MEXLy Cen de Agi enrgn Whi 3609 (MICS, CE Zapotece formosa subsp. svholtit ix closely ve luted to und was probably derived fromm subsp. roses. ‘The foreoer has generally smaller stature, less nt reerous, smaller leaflets, and atuller pods (Table 15}, These two taxa show geographically eontig- uous distributions (Fig. 44), and there i clinal variation in leaflet sey with subsp. sohotit oc- ccupying the extreme pesition. Although the pat- terns of morphological variation are apparently continuous between the twa taxa, subsp. schotté is relatively easily recognizable as separate taxo- rome unit with a somewhat axbiteery boundary In the Bahoquivari Mountains and in northern So ‘ora and Chihuahua, a number of difficulty placed imermediate specimens have boen collected, Zapoieca formosa subsp. schott eould be eon fuse with subsp. mollica, since they have eaters of comparable size. However, subsp. sehottié can be readily distinguished by the longer internodes and more elongate petioles and rachillae (Table 14). Further, although vestituee is highly variable in subsp. motliculn, most of its individuals have denser vestture than de plants of subyp. sehatte: - 13. Zapoteca seutellifera (Bonth,) H. Hern, ‘comb nov. Calliandra seutellifora Bonth. J Bot. (Hooker) 2: 189. 1840; Benth,, London J. Bol. 3:98. 1844: Benth.. Trans, Linn. Soc London 30: 542, 1875; Benth. in C. Martins, Fl, Bras. 15(2p 412. 1876. Fouilleea sew tellifera (eath,) Kuster, Revis. Gen, Pl 189. 1891. TyPE: Broil. Terntory of Rondnia: Rio Madeira, Riboirdo, in siecia arenasis, May 1828, Riedel 1334 (eetorype, K, here des ignated: inolectotypes, A, K, LE (3), NY— fragm., P, US) Seandent shrubs; branchlets toreto or suban ulate, the surface costate under magnification, slabrows to finely puberulent, usually villous on the yyoungor parte. Leaves glandular, the pinnae 2-3-PREV Volume 76, Numbor 3 1989 Hernanaez Zapoteca 853 jugate; petioles terete with o conspicuous aden) channel, 3.5-B ex long, glabrous to sparsely wile Joos, villows in the area of the ebannel; rachis 1~ 5 em longs tachillae 5-7 em longs glands clove to the base of the petioles, at the paints of insertion of the pinnze, and sometimes between the distal pairs of leaflets, scutelform, the larger ones on the petites to 2 mm diam, glabrous 10 poberub abaxiallys leaflets 6-11 pairs per pina, oblong: obovate, falcate or subfaleato, oblique atthe base, rounded to subseute at the apex, mucronate, the median ones 15-20 5-7 mn the proximal ones spnaller, mersbranous, labrous, sometines ciliate, primary and secondary leaflet veins conspicuous abaxilly: stipules leafy, linear-triangular, to 4 mm Jong, glabrous to villou:. Capitula axillary: podun- les Solitary or fasciculae at distant nodes, glabrous to sparely villous, ea, 6.5 cm long at anthesis Flowers pentanierous; ealyx cup-shapet, 11.5 min Jong, finely ciliate, the tecth triangular, acutes eo rolla carspanulate, 45 mn long. glabrous. the Joes lanceolate; flamemts e3. 2.5 em long, white inthe basal hal, piok atthe distal lf, the starsinal tube ca. 1 mm long: ovary ca. 1 man long labrous Pods acute, rostrate, ca. 9% 0.7 em, thickly mombranous, finely puberulent, especially at the margin. Distribution (Fig. 44). Bodemic to the Am- ‘zon Basin where it has heen collected at scattered localities, In Brazil knoven fram the states of A Zonas and Paré, and from the tersitory of Rondinia, algo in Bolivia’ from the departments of &l Beni ad Santa Cruz, It usually occurs in open sandy Aéitional sperms examined, nari: snnzonas ‘Ammzonac ad ini fa, Sls, Spruce 1386 (BM, GoT- GH. NY, Py rans Ms de Tt, entrada Stntdrénn“Coiak, BM 168. Ken LOL1 com ums pene wacko de 3 kas Jeneo da mata, merger desta do Rio Jomasiza sl d Para, T0PA0'S, 39°19 W. Amare! ab 3251 (MO, NY}. Boule LL WEN grows Vaca Dies Gucyoramerin, 10-495,” 05422W, “Krapraickas & ‘Sohinint 35129 (MD), sata coz: prow. Sor, Buenas ta, 450m, Sieinhach 7047 (A, UNF, GG, K, NY, Mo, ‘This morphologically uniform species is closely related to Z- formosa but differs by having sou telliform extrailoral glands, seandent habit, and usually emaller, inear-trangular stipules, More lover. it ean be distinguished from Z. formose subep formosa by ite white/pink, rather than whitey fi ments. The eeutellform glands, whieh aro con stant throughout the known populations of Z. se tellifera, are on excellent diagpostic Feature, as they occur only in this species of Zapatece, They are fully developed at the earliest stages of leaf development, which probsbly reflocts their defen sive fanetis 14, Zapoteca tehuana 1. Hern., sp.nov. TYRE: caltvated at MO, Jan. 1, 1986, # Hernan. des 950 (holotype, MEX: isotype, MO). (Original cource: Mexico. Oaxaca: Distr, To: Jrauntepec, El Manguito, SW El Limén, 2 LLL km SW de la entrada 2 Bucnos 3 tm, R. Torres 4167 (MEXL), Aa types to be distributed, Figure 45, Frsices, coutbus geoelius. Finns (1 JX Suga, tolls 10-21 jugs in utraque inna, ablongmianconkt Yel ovatolancraat, mombranaces vel aabslerace, frdinatie venarum erideser trochidedruma stale folinse ca 2 mm hinge. Neanerin hremnonomatu mates 20 = 26, rubs ca, 11.8 m high; stems slender, 68. 5 10 mm diam. at haze, the bark amooth except for al extoliations; branchlets torete, fissured under agification, glabrous. Pinnae (1-)2(-3)jugate Petoles ezlandular, terete, with a conspicuous fdaxial channel, 0.5~3 em long, glabrous, see times sparcoly villous; rachis 0.71.8 em Tong in 2-jugate pinnae, to 3.8 em long in S-jugate pinnae; rachillae 19 7 cay longs leaflets 10-21 pairs por Pinna, oblong-laneeolate to avate-lancectate, the distal pairs sometimes obovate, at base oblique, at ‘9p0x acute to suhacste, sometimes mucronulate, tho meilian ones 9-15 % 2-4 mm, the proximal and distal pairs cmallor the enterior proximal leaf: Tet reduced in sie, rarely lacking, ell membranous to thinly chartaceous, usually deving light green, usally glabrous throughout, occasionally pilose snd ciliata, with conspienous brachidodramous vera tions stipules leafy, triangular, ca. 2 mm long slabrous, rarely pilose, Capitola axillary: peduncles fasciculate at distant nodes, ifr, (2 10cm “Tong at anthesis, Flowers pentarserous, or rarely totramerous: ealy ox, 22.5 mm long, elabrous fr pilose, usualy ciliate, eceh tooth with « central vein, tiangular, ca. 1 mm longs earola campan: late, ea, 55.5 i long, glabrous, the lobes ovate lanceolate, acute; filaments (20-J24-25 mn long, white in the basal half and pink in the distal hall ‘or white throughout, the staminal tube ea. 2 mm Tong: ovary ea. 1.5 mm long, very shostly (= 0.3 ram) stipitate. Pods Linear, at apex subacute, ros tellate, to 12 en long when mature, thickly Dranous, glabrous, reticulate-veined. Seeds widely ‘ovoid to widely rhomboid, fattened, 5-6 x 4mm, ray, the areols light gray: pleurogram ierogular, dervateates! by « datk fine, Somatic chromorome number 2n = 26 NEXT854 Froune $5. Zapotecn sehuaon pole 0. Cease I of venation i cleuradTeaBel—E, Fl ‘Annals of the Missouri Botanical Garden NEXT A. Traces wih ilerewences at anther —B. Branehlet with debised WF Asher, —C. Dioacied ower. Felpad.—1 Sees, (Drown fam ling pent 7. Hemdndes 500 MENU, MO po fm Tres #107 MEXU)} Distribution (Fig. 84). This epeciesis ender vo a resirited area southwest of the Isthmus of Tehuantepee in Oaxaca, Mexico, where it grows fn secondary vegetation derived from trapieal de- cishuous forest mixod with probably rekictval cle rents of temperate deciduous forests. Adidivional speconens esamined. MEXICO, OxXaca: Dist, Tetantopec, eafada en of Arroyo de les Micas 2 al SW de HI inn y Lé kal NW de Tehuantepec, Garvie & Terres 1679 QMEXU, MOy Dist, Tete pe, El Limin, ariba de Bl Chicoapoce, SW’ de Te Aveitepec,entrande yor amino a Bueoos Airs, H. Her éndes & Torres 303 MEXU, MO}, Dist. Tebanepee,PREV Volume 76, Number 3 1989 ELMangit, al SW de BL Lins el eval sé 8 111 AISW dls estrada aBwnen Aron Forres 4767(MEXU, Moy, Dit Toboansepen, 15 km SWW de Buenos Aire: hela Tenanae, entrardo per Hierba Sent, Torres Maritues 7375 (MEXUy,Subida al Coe Cuicgola por isdera SL. Toes ot als 208 (MEXU}. Zapotecs tehuavua isa narrowly endemic species probably not closely related to species of subs, Zapoteca, and further knowledge on this specirs could provide evidence for its placement in a sep srate subgenus, I difere from the ramaining specie feubg, Zepotece primarily by having eonpicuous brachidodromous leaflet venation and. relat thick lets, Tt might be confused with x the species with oblong or avate-lanceokat leaflets of comparable size (et, Z. portoricensis and Z. Tambertiana) but difors by having leaves uswally having fewer pairs of pinnae, smaller stipules, white or white/pink filaments, and shorter stature. The felationship between Z. tehuana and Z. aculeata are discussed under the later 1. Zapoteca subg. Aculenta H. Hotn., subg nov, TYPE: Zopotera aculeata (S Benth.) H. Her Pinas I> rara2ugee fae sine ect fare gles file tat charts a + abr vel ue inca, mein ices, cedintons vena con ‘pieuaystiplae spineccenten, Capita axiler. Policia ‘aworbus lenbeulorbs presi. Lagumine corse ga te, Leaves glabrous or nearly so, the pinnae 1, racely 2-jugate, the petiole without nectates, leat lots 3-7 pairs por pinna, moderately largo. char taccous to coriaceous, sith a very conspicuons ‘venation; stiplesepinescens. Capita axillary. Po- Iyads with Tens-shaped thickenings. Pods coria- ecu, glabrous, ‘This eubgenus comprises «single spociee, 2, faculente, characterized by medium size, chart. eous to coriaceous leulets, and spineseent stipules, 15, Zapoteca aculeata (Spruce ox Benth.) T. Hern, comb. nov. Calliandra aculoata Spruce ex Benth, Trans, Linn, Soe, London 30: 541 1875. 1998: Beuador, ad pedes m. Tangara Inus, Sep. 1857, Spruce 505d (leetotype, K, here designated: iolectotypes, BM, F, G,(2) S. (2). The ieoloe totypes are dated as fellows: BM, 1857-9: F, 1857-9; G, 1857-9, 1861; CH, 1857-9; K, Aug. 1857; LE, 1857-9; NY, 1857-9; P) 1857-9, 1861; 8, 1857-9. Figure 46, Hernandez Zapoteca 855, Trees toca. 10m all; ranchiets terete, cestate, labrous. Pinnae I-, rarely 2-jugate; petioles eglan ular, terete, with a conspicuous adaxial channel, $25 mm long, villous, raroly glaheato: rachis 5 om loog when pinnae 2-jugate:rachillae 112.5 «em long: leaflets (3-)1~6(-7) pairs per pinna, the proximal anterior one usually reduced oF lacking, ll ovate-lanceolste, at base oblique, at apex acute to acimninate, the distal ones 1.56.8 ¥ 0.4 2.8 fem, the proximal ones smaller, all chartaccous to coriaceous, glabrous, the buso sometimes villous: leaflet venation conspicuous; stipules spinescent, Fei, adpressed to the branchlets, sometimes be coming divarieste, Iinearlancealste, sometimes scarica, heaked, 2-10 mm long, glabrous, Capita ry! peduneles usually fasciculate at distant nodes, rarely solitary, (2.5-)5-9 em long at an- thesis, Calyx shorly, sometimes irregularly den: tate, ea. 2.5 men Tong, glabrous, ube teeth some times ore than 3, sballowly triangular to deltate; coralls camipanulate, 5-6 rm long, glabrous, the lobes ovate-ancealate, aes Rlamients ea. 2.5 em Jong, red-purple, the staminel tube e8. 2 mm long; ovary ca, 1-1.5 mm long. glabrous. Pods about 10-125 * 0.9-1.1 em, subacute 29 rounded at apex, sometimes rostellate, coriaceous, glabrous, the surface reticulate-veined. Distribution Pig. 47). Zapoteea aeueata has been eallzetad from natural populations at the foot of the Mount Tunguraha io the Beuadorian Andes (Province of Arubato) and fran cultivated individ- valsat public gardensin Ambalo, Basin, and Quite Gallections come from elevations af 1,900: 2.850 m, Additional specimens examined. —Ecvavon, uncut ade tS 0! Bain, Pendarad Summers 10M (F, GH, US), Cuctivateb. Bovabon. 9 cunt! Quo, deosta Safs 13599 (F). FUNcuHAKL Ambuto, deaita Soll 9488, 9523 (Fy Rose & Rose 22363 [GHLNY, US. Zapoteca aculeata represents by itself 4 sub- gems of Zaboteca. It is a poorly known, unis Takable species charactorived sy spinescent ti ples, medium sie, and corigeoous, ovate-lanecolate Jeaflets, which are usually larger than the estea: tally membranous leaflets of subg. Zapoteca. Iris not obviowsly closely related to any other spe ‘of the genus; homever, as shown in the chapter on morphology, an analysis of sts Iealet architecture revealed outstonding similarities to that of Z. te fuanas. Zopoteee aculeata is known from only 8 few collections made in a restricted area of the Bcundorian Andes and from cultivated individuals NEXT856 Annals of the Missouri Botanical Garden Data of from Acosta 13599 (MO), ‘at Ambato, Ratios, and Quito: it has not been cole Folia ine ne lecvel for more than 40 yeurs, Jug fs 1PREV Volume 78, Number 3 1989 Fron 47. ors. Loaves without noctaries, labrous oF nearly 20, the pinnae 1 juga leailete 11% pairs per pin, chartaceous to coriaceous; leaflet venation con- spieuous: stipules leafy. Capitola in long, simp ceompaind yeoudopanieles, Polya with lens:shaped thickenings. Pods eoriacoou, glabrous. Seeds with out plenrogeam. This subgenus comprises two relatively special jae species, Z. micracephate anil Z. amazonien, ‘which are characterized by a set of characte that aze highly advanced in Zapotoca: leaves re duced to one pair of pinnae, with 1-1! pairs of large, chartaceous to coriaceous leallets capitals mized in long, spe or icles, mnpound preutopan 857 Dsnbuton of Zapoteca eculeata (tangles, Z. microcephaie (open square) end Z. aneznnica 16. Zapoteea microcephala (Briton & Killip) Hi Vern, cou. nov. Calliandra micraceph- fala Britton & Killip, Ans. New York Acad. Sei, 35: 154, 1936. Tyre: Colombi: 1760- 1808, Mutis 07 (holotype, USno, 1561096: isotypes, MA (1) not eon photos at MEX, MO). Shrubs; branches terete, the suelace srooth, plebraws. Pionae I-ugate, petioles eglandilar, slender, with « pair of channels running longitw. dinally in the adaxial surface, (30-)40-70 mm long, glabrous abaxially, usually scabrous between, the channels rechillae 13-19 anim longs leaflets 14 pairs per pinna, the proximal anterior lati lucking. all falcately lanceolate, at base strongly NEXTPREV 858 Gamngarison of Zapoteen miroeeplale wi ‘Annals of the Missouri Botanical Garden 7 anasonica. raion NEXT Ninian Pere Magdalena River Valley, Glob Dewibaton Habit rot arabe Pains of leaflets par giana 1s Pete length (0-10-10 mmm Leaf pubescence ciate Peoudopaniels| Lengtt 20-30 em Stipa lrgoh 1-2 um wile Fine calor oblique, sharply long-acumsinate a ape the distal leaflets 7.5- 10.5 % 2.5-4.5 em, the proximal one smaller, all ehartaceous, glabrous, usually ciliate: leaflet venation eonspicnous, with 2 primary and atleast I secondary veins prominent; stipules leafy, Uriangula to narrovly tlangular, ea. 1-2 munlong, glabrous, Capitula in terminal, elongate, simple preudopaicles a, 20-80 cm longs pesiuneles fas ticalate,carely solitary, fliform, 2. 4.5 cm long, ft anthesis, Calyx ea, 2 mm long. the teeth tn angular. < 1 in long: corolla carmpanulate, ca. 3-4 rom long, the labes ovate-lanceolate; stamens white, the starninal tube ea. 2 mm long, Pox to 15 x 1.1 em when mature, at apex acute or retuse, ccoriacens, glabrous, the valves reticuls evened, Distribution (Fig. 52). Endomie toa reacted ‘area of the Median Valley of the Magdalena River in Colombia. Additional specimens examined. Covownta sastobie viiity of Puerto Brro, between Coracé ae Magdaloa rivers, Heaght 1711 (P, K, US) Haught 1794 (US wthout loca, Mate 3400 (UA, US) This narrowly endemie species ie eleuly allied te Z, amazonica, from which itis geographically * olated (Pig. 41). Zapotecee mierocephale cat be readily distinguished frorn that species by 1% p ‘of smaller leaflets por pinna, simple (nonrarified), shortce poeudopenielos, «mailer stipules, and whito filaments. Table 15. provides an account of the ‘main comparative characters distinguishing these lovely related species. Zapotecs microcephala x known from only four collections dl has nt heen collected since Haught (1711, 173) found it in 1935 in the lorstcaly rich ares of the Magialena River Valley, Colombia, 17. Zapotecs amaronien (Benth) H. Hers comb, nov. Calliandra amazonica Benth London J. Bot. 3: 94. 1844 Benth, London scant shrub 1 20-42 rt abrows compo 40 en cee prple J. Bot. 5: 101. 1846; Benth, Trans. Line Se, Londan 30: 538. 1875: J. F. Mache. Fieldiana, Bot. 13: 67. 1943. Ings bawhinie folia “bawhiniaefolia® Pooppiz, Nov. Cen. Sp. Ph. 8 80, 1. 290. 1845. (The epitet is [presumably alo based an Poeppig 2070 used bby Bentham i the description of Caliaeadra famazonien.) TYPE of Z, amazonea: Pet. Loreto: in sylvis provinciac Maynas eircura ‘Yurimaguas. 1824, Poeppig 2070 (letotype, G, hove designated: isclectorypes. F.C, LE pro parte, NY, P). The specimens at Gand P age dated 1834, whereas those at F and NY are dated 1891. Figure 43, Scandont shrubs 3-5(-8) m talk stems slender, cca. 15 mim diam, the bark smooth; branchlets spreading, terete, glabrous, Pinnae I-jupates pet fakes eglandular, 200-42 sam lng, usually glabrons rachilla + corrugete, with a conspicuous adaxial channel, 10-21 mm long (excluding the pulvini ‘usually glabrous, but occasionally sparsely willows in the extremes: Tellets [ paie per pana, sleately lanceolate-avate, 7-22. x 2.8-9.2 em, strongly oblique basally, acurninate to caudate, chartaceous to coriaceous, usually drying lighter green abaxi- ally, = lustrous adaxial, glabross of rarely gla- bate; leaflet venation conspicuous, with 2 primary and at least 1 secondary vein prominent; stipules leafy ovate-lane lero acute at apex, 22% 3-8 mmm, glabrous, Capitula arranged in terminal elongate, compoud pxeudopanicles to 40 long or more, tho pedunelns faceivulate raraly solitary, 0.8-7 cm long at anthesis, Calyx dentate, ca, 2.2 mm long. the teeth shallowly tangular or dltate, ca. 0.7 min long, glabrous, rarely ciliates corolla campanulate, ea. 4-5 mun lang, glabrous. the lobes lanceolate; laments shite in the bass! half and purple:red i the distal half, the tube ea 2.5 mm long: evary about 2 mu long, glabrous,PREV Volume 78, Number 3 1989 Feuer 48. Zapotec spyooth bark. -C. Leaf is Pods to 24 | em sthen mature, acute at ape, Jabrous, the valvos retieulate-veined ly rhomboid to narrowly allipsoi lat 4% 5 mam, dark brown, without ple Hernéndez Zapoteca Pastaza in the Depa 859 <0. Primacy stem showing F. Pod. G. Sccd. From, Fig. 47). This species has been hay sites of undisturbed coin Peru, from Coronel Porillo (Rio neniv of Loree, especially NEXTPREV 860 slong Rio Huallaga. Collections come from 360= 900m, Aditi! apocimens examined Pent WSts: Sa le ad Satipo, Woythornks 5943 (F, GUL LL. MO, NY. {US}, uukaur sve Rio Muallga ot Ting Mari, Croat 21102 (MOY Tengo Maris, km 138, toes te dl Ag la, Ferrevra 082 (USy, Pacitea, Puerto Ine, Just N of town (18'S, TH58W), Poster 9332 (MO) Paste Uo de Ponta (40'S, 75°28'W), Foster 9209 (MOS tong Rio Huallaga, veer vachusla Chontalaguay Mexia 005 IMG Fy Ce GH, MO. NY. S, LC. US Lemes rato, Rupa Rupa, ‘Tigo Marla (18S, 75°59°H), Pronomars eral. 11249 (F, Ue Tingo Maria, Stork & Horton 9477 (8, 6, UC), Lowes Coronel Porto, 3-5 orae arta. del caverio Abwa0, sora Rip. Abi Dias 61 (MO, NY) mines 9110 Panel serra ‘ero in Alto Make de Drs fos Shinty, emery rt (al 27203 (MO), san marnin: Maisal Cece, Toesrhe ‘Nuevo, Cerzo Sin 13 iim W.Tocache Nawvo slong eal t Puerto Paana (8°00, 7692°W), Plowman Sehune 1115 (Fy Marcel Civerc, Depto. Teche, a de Tocache Nusve, Funda le Campine. Scho 3371 FG, NY, USk Mariscal Caecea, Dept. Tocarki Nuvo, Cons sarge foto Rip Haya Ske £497 (FG. NY, Uy Maral Cees, Depo. Tocache ‘Nuevo, Muipaco de Baha, Probara, | im ajo deeery booadira Rio Miso, Sehunke #237 (F. GH, MO, US Mariscal Ceres, Deto, Tocaebe Nuevo, Qoekroda de Santis FE Pacrto Para, Sohunbe 643 (MO, KV Marae Cicer, Puerto Panna (omrgen derecho del Rio Malaga, SoPunke 691 9(MO Meriaeal Cores, Dept TTocache Noewo, Quebrala de Husqusha imargen i> ualaga), Schunke 2053 (MO, NYp Tarapet, Sprace 1494 1B, BR. F.C. GH. KP. NY‘: Alto Rio Hua, Tarapei, Wiliams 0588 0, US). Zapotece amazonica and Z.microcephala rep resent a distinct evolutionary ne within the genus: these closely related species probably diferentiated from a cormmon ancestor after becoming separated from the rest of the gens, Zapotec amazontca, which is remarkably uniform morphologically, ean bie readily distinguished from Z. microcephala by ite scandont habit, ono pair of largor lofi pins, larger tiles, elongate peoudopaniles, and Wwhive/purple.ced filaments. The diferences be: tweon the two species were discussed further under 2 microcephala (ver also Table 15} Zapoteca amazonica probably is the most spe~ alized species in the genus, presenting a combi nation of derived charactors: scandent habit; 6x- treme reduction of leaflet number eoncomitaa: with ej relatively complex leaflet ve- nution patterns; capitala organized in compen, ter- ‘minal, compounl pseudopanicles. The specialized nature of this species was discussed in the section ‘on morphology. Lnunyrone Chuo AuSuies, AN, 1982. ‘The palmeinate and p |26uid= TDR 22424697 2104740729353 ‘Anais of tne ‘Missouri Botanical Garden Pranee eto) inlet Divernon inthe Trop ics, Collin Unie, Pres, New York. Aust: ML 1982. Qiainrary palyslogeal shes ‘he Ariston. Basin, Fp. 67-79 in G, Pane eater, Bislegeal Divoscaion in the Tropes. Colmbia Unive Prose, New York AursaNDEn,M-B. 1969. Dlfevental staining of sorted ‘al mnaborted lle. iain Terhowle S40 11712 Angoxo, MT K. 1981 Dering syste aed polit natn biogy ey Leguminase: Bp. 723-769 tn B M. Pail & P. He Raven (ature, Advances tn Legume Spstrtice, Pert 2. Reval Botan Gardens, Kew Rietmond, Surrey. Enalend Bwin, Ti G&L Bakin, 1985, Floral nectar augue ‘onsitents in elation 0 painat type. Pp. LI TL nC. Jones & RI. Lil eitoe) Handhook of Experimental Pellnston Biology. Van Nostrand Feintold, New York. SB J Hsin 1957. ‘The goliation of Parkin by ists ands stendant evcluiany pele lem Ewhutnn TL: 480 860 Bawa. S197. Breeding systems of ee species ‘fa lowland topical omunry. Evolution 28: B5— 2 Bowtie, C. 1819. Contributors towards 4 fora of ‘Sout Amerisa—ersmeraton of plant collect by Me. Setomargk in Brisk Guiana, Hook J Bo, 2 12-14, BAS. Kote fn Mimoness, with « syoupais of the species Lenn J. Bot. 3 83-112 1875. Revision of the sohorder Mimoeae ‘Teate, Linn. Soe, London 30; 338-06), Bewrtss,B. 1977. 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