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1-1-2006 Nomenclatural Innovations in Neotropical Salicaeeae Mac H. Alford University of Southern Mississippi, [email protected]

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Recommended Citation Alford, M. H. (2006). Nomenclatural Innovations in Neotropical Salicaeeae. Novon, 16(3), 293-298. Available at: https://aquila.usm.edu/fac_pubs/8569

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Nomenclatural Innovations in Neotropical

Mac H. Alford

L. H. Bailey Hortorium, Cornell University, 228 Science, Ithaca, New York 14853, U.S.A. Current address: Department of Biological Sciences, University of Southern Mississippi, 118 College Drive #5018, Hattiesburg, Mississippi 39406, U.S.A. [email protected]

ABSTRACT. M. H. Alford is described e transferido para Abatia Ruiz & Pav6n, com base em as a new of Salicaceae based on Neosprucea analises de seqiiencias de DNA do plastideo e devido kuhlmannii Sleumer from Brazil. The deciduous h ausencia de descontinuidades morfologicas entre os perianth, papillate teeth, presence of receptac generos. Em razao do nome Abatia spicata (Turezani ular disk glands, and analyses of plastid DNA data now) Sleumer ji existir, Abatia angeliana M. H. suggest that the new genus is more closely related Alford e publicado para substituir Aphaerema spicata to Merrill and L. 0. Miers. Um lect6tipo para Aphaerema spicata tambem en Williams than to Banara Aublet or Neosprucea Sleu designado. 0 genero monotipico Priamosia Urban, da mer. The monotypic genus Aphaerema Miers of Brazil ilha de Hispaniola, originalmente descrito fora de and Argentina is transferred to Abatia Ruiz & Pavon, Xylosma G. Forster com base em seus poucos estames based on analyses of plastid DNA sequences and on (quatro, em vez de oito a numerosos), e transferido the lack of morphological discontinuities between the para Xylosma, e um ne6tipo e' designado para Xylosma genera. Because the name Abatia spicata (Turczani domingensis (Urban) M. H. Alford. now) Sleumer already exists, Abatia angeliana M. H. Key words: Abatia, Aphaerema, Argentina, Ba Alford is published as a replacement name for nara, Brazil, , Hispaniola, Macrothu Aphaerema spicata Miers. A lectotype for Aphaerema mia, Neosprucea, Priamosia, Salicaceae, Xylosma. spicata is also designated. The monotypic Hispanio lan genus Priamosia Urban, originally described as Based on recent phylogenetic analyses of Salica separate from Xylosma G. Forster based on its few ceae using both morphological and molecular data stamens (four, instead of eight to numerous), is (Alford, 2005; Chase et al., 2002), several taxonomic transferred to Xylosma, and a neotype is designated changes were deemed urgent and sufficiently well for Xylosma domingensis (Urban) M. H. Alford. supported to precede a forthcoming treatment of the RESUMO. Macrothumia M. H. Alford e descrito como family in the Kubitzki-edited series The Families and um novo genero de Salicaceae, com base em Genera of Flowering (S. Zmarzty, in prep.). Neosprucea kuhlmannii Sleumer do Brasil. 0 perianto Salicaceae now include a large part of the former deciduo, as papilas nos dentes das folhas, a presenca Flacourtiaceae (Alford, 2005; Chase et al., 2002), and de discos glandulares receptaculares e a analise de the taxa considered here were all formerly placed in dados de DNA do plastideo sugerem que o novo Flacourtiaceae. Phylogenetic analyses indicate that genero e mais proximamente relacionado a Ahernia the monotypic Aphaerema Miers is nested within Merrill e Pleuranthodendron L. 0. Williams do que Abatia Ruiz & Pav6n, that the monotypic Priamosia a Banara Aublet ou Neosprucea Sleumer. 0 genero Urban is nested within Xylosma G. Forster, and that monotipico Aphaerema Miers do Brasil e da Argentina the usually treated as Banara kuhlmannii

NOVON 16: 293-298. PUBLISHED ON 7 NOVEMBER 2006.

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(Sleumer) Sleumer characteristics is indicative of Ahernia[= and PleurNeosprucea kuhlmannii Sleu mer] belongs anthodendron. in Because a this speciesseparate differs in several genus allied more closely to Pleuranthodendron key characters from those other genera (see de L. 0. Williams and Ahernia Merrill. scription and key below), a new genus is erected here. Another possibility is to lump all three entities into

MACROTHUMIA, NEOSPRUCEA, AND BANARA Ahernia, the name with priority, but because their interrelationships are still unresolved (Alford, 2005) In 1950 Sleumer described a new species from and because Pleuranthodendron is a relatively com Brazil as part of his recently established genus mon genus of the Neotropics, the more conservative Neosprucea Sleumer. Sleumer (1936, 1938) had and less disruptive approach is the creation of a new segregated Neosprucea from Banara Aublet based on genus. the former having spiciform racemes with 4- or 5 merous and linear-elongate anthers. In contrast, the remaining species of Banara usually Macrothuinia M. H. Alford, gen. nov. TYPE: have paniculate inflorescences (rarely racemose, Macrothumia kuhlmannii (Sleumer) M. H. Al fascicular, or with flowers solitary) with 3-merous ford. Figure 1.

flowers and globose to ellipsoidal anthers. Both genera Genus novum, Aherniae Merrill affine, a qua fructibus have highly intruded parietal placentation and in majoribus et numero petalorum differt, et Pleuranthodendro clude several species with that are 3-veined L. 0. Williams affine, a quo inflorescentia haud ramosa et from the base. Sleumer's (1950) new species, N. numero seminum et tepalorum differt. Arbor, folia simplicia, alterna, ovata vel oblongo-elliptica, ad apicem petioli kuhlmannii, had strongly 3-veined leaves and highly glandulis duabus rotundis crassis operculatis instructa, intruded parietal placentation, but he saw no material margine inconspicue serrato, venatione foliorum actino with flowers. Regardless, he was convinced that the droma. Inflorescentia terminalis racemosa fasciculatiformis new species belonged in Neosprucea due to its large vel umbelliformis, sepala et petala 3 vel 4, post anthesin (> 3 cm diam.). Such a large fruit had never decidua, stamina numerosa, antherae ellipsoideae, glandulae disci praesentes, ovarium superum. Capsula globosa, 3-5 cm been observed in Banara, and the few available diametro, multiseminata, pseudo-8-9-locularis, placentibus of Neosprucea were larger than those of most Banara. 8-9-lamelliformibus, in cavitatem profunde prominentibus, Following the collection of additional specimens, pericarpio fragili subcoriaceo. Kuhlmann (1953) reinforced Sleumer's claims, point to 20 m tall, 30 cm DBH. Leaves simple, ing out that N. kuhlmannii was 4-merous and had alternate, ovate to oblong-elliptic, 6-13 X 3-8 cm, flowers disposed in a racemose inflorescence. Kuhl apex acute to shortly acuminate, base rounded to mann did not mention anther shape. These character subcordate, chartaceous, deciduous, margin subentire istics matched those of the described species of to inconspicuously serrate with papillate apices of the Neosprucea and were unlike those of Banara. teeth, venation actinodromous, petiole 2-4.5 cm long, In 1978, however, Sleumer transferred the species apex of petiole bearing a pair of thick, projecting, cup to Banara without comment, presumably because he shaped glands. Inflorescence a terminal, congested, had then seen flowers and realized that the globose to fascicle- or umbel-like raceme of 3 to 9 flowers; ellipsoidal shape of the anthers was characteristic of and 3 or 4, creamy yellow, ca. 1 cm long, Banara, not Neosprucea. Sleumer (1980) maintained deciduous in fruit; stamens numerous (ca. 100 to 130), this treatment of the species as Banara kuhlmannii anthers ellipsoidal, longitudinally dehiscent; disk (Sleumer) Sleumer in his monograph of the Neotrop glands present, consisting of a single whorl of 6 to ical Flacourtiaceae. 8(to 9) pad-like or crateriform disks, ca. 1 mm diam., The first specimen of Banara kuhlmannii that I with stamens arising inside, outside, and between the examined (A. M. de Carvalho et al. 6689, MO) was glands; ovary superior; style 1; stigma consisting of annotated by Ron Liesner in 2000 as "If Flacourtia several minute craters at the apex of the abruptly ceae, new genus! To me it looks like a Flacourt! Need expanded style. Infructescence of few fruits, often fls [flowers]." Although the species had already been consisting of only 1. Capsule globose, 3-5 cm diam., described, his intuition was correct. The species has multiseeded, with 8 or 9 highly intruded parietal several characters atypical of either Banara or placentae, fruit wall subcoriaceous but brittle, base of Neosprucea. The glandular apices of the leaf teeth style persistent. Seeds exarillate. are papillate (violoid) instead of spherulate (salicoid) or torus-shaped (see Leaf Architecture Working Macrothumia is a new genus allied to Ahernia and Group, 1999; Hickey & Wolfe, 1975; Alford, 2005), Pleuranthodendron. Macrothumia has parietal placen and it has receptacular disk glands and a perianth that tation, congested, fascicle- or umbel-like racemes, is deciduous after anthesis. This combination of three or four petals per , and large fruits

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Figure 1. Macrothumia kuhimannii. -A, C. Flowering branches. B. Leaf, with close-ups of teeth and glands at the apex of the petiole. D. Floral bud. E. Flowers at anthesis. -F. Longitudinal section and cross section of a flower showing receptacular disk glands. G. Stamens. H. Pistil, close-up of stigma, and cross section of ovary, showing highly intruded parietal placentation. I. Mature fruit. Scale bar equals 3 cm for A-C and I, 5 mm for E and F, and 3 mm for G and H. Length of D equals 1.3 cm. A, B, D, F, G, and H drawn from Mort et al. 10641 (NY, herbarium sheet, spirit collection, and photos), C and E from Thomas et al. 12530 (NY, herbarium sheet and photos), and I from Pirani et al. 3384 (NY).

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(> 3 cm diam.) 3b. with Raceme axillary, notmany congested; fruit seeds, whereas Ahernia has small, obovoid to ellipsoid (< 2.5 cm 10 to 15 petals per flower and smaller fruits (< 2.5 cm long); petals 10 to 15; Philippines . . . Ahernia long), and Pleuranthodendron has large disorganized panicles and one seed per fruit. ABATIA AND APHAEREMA Etymology. Macrothumia (ptcypoO uio [f.]) is Abatia, Aphaerema, Pseudoscolopia Gilg, and Ho a Greek word for long-suffering or enduring patience, malium Jacquin are the only genera in Salicaceae that a virtue prized by taxonomists. have opposite leaves. Aphaerema and Abatia are closely related Neotropical genera, Pseudoscolopia is a South Macrothumia African genuskuhlmannii of a different tribe, and Homalium (Sleumer)is M. H. Alford, comb. nov. a pantropical Basionym: genus of a different tribe with Neosprucea only a few kuhlmannii Sleumer, speciesLilloa from Madagascar 23:that have opposite 248. or whorled 1950. Banara kuhl mannii (Sleumer) leaves. Aphaerema is a monotypic Sleumer, genus from Brazil Blumea 24: 118. 1978. TYPE: and Argentina Brazil. of small (< Espirito0.5 m) with 8(to 12) Santo: Colatino, Rio Doce, 30stamens andNov. no staminodes (Angely,1943, 1962). Abatia J. is G. Kuhlmann 6559 (holotype, aRB genus of ninenot montane seen; species from Mexico isotypes, and LIL not seen, RB photo Centralat andUSMS). South America of small- to medium-sized shrubs (to 4 m tall) commonly with numerous stamens Distribution. Brazil (Bahia, Espirito Santo, Minas (ca. 20 to 35) and an outer ring of filamentous Gerais). staminodes. Baillon (1872) noted that two new species Representative specimens examined. BRAZIL. Bahia: of Abatia, since described as A. stellata Lillo (1919) Belim & Pinheiro 2878 (NY), Carvalho et al. 6689 (MO), and A. mexicana Standley (1931), have as few stamens Mort et al. 10640, 10641 (NY), Morn & Thompson 11016 (4 to 8) as Aphaerema. Baillon (1872) considered the (NY), Thomas et al. 7063 (NY), Thomas et al. 12530 (MO, NY). Espirito Santo: Kuhlmann 6449 (FSU), Pirani et al. genera closely related and did not believe that the 3384 (NY). Minas Gerais: Mexia 5294 (BM, NY). absence of staminodes was alone sufficient for generic status. Thus, he treated Aphaerema as part of Abatia. A thorough description of the species may be found Furthermore, several other species of Abatia, A. in Sleumer (1980), as Banara kuhlmannii, although americana (Gardner) Eichler, A. glabra Sleumer, and he failed to mention the key character of receptacular A. microphylla Taubert, have intermediate numbers of disk glands and reported, with the material at his stamens (12 to 20) and form an overall morphological disposal, that the flowers were 3-merous (or rarely 4 continuum between the larger, common species of merous by splitting of the petals) instead of 3- or 4 Abatia with numerous stamens and staminodes and the merous. Information about the use of Macrothumia diminutive Aphaerema with few stamens and no kuhlmannii as refreshment for both humans and staminodes. Despite the clear continuity between wildlife may be found in Kuhlmann (1953). Aphaerema and Abatia, their segregation has been maintained, with some authors producing dichotomous KEY TO THE GENERA OF SALICACEAE WITH LEAVES 3-VEINED FROM keys with number of stamens as a principal character, THE BASE, PAPILLATE (VIOLOID) LEAF TEETH (NOT SPHEROIDAL OR TORUS-SHAPED), AND PAIRED GLANDS AT THE BASE OF THE LAMINA presumably unaware of the less common species or OR APEX OF THE PETIOLE their characters (e.g., Hutchinson, 1967). Further evidence of their close relationship has been inferred la. Placentation axile, or parietal with later develop ment of a pseudo-axis ("pseudoaxile"); perianth from analyses of plastid DNA (Alford, 2005). persistent in fruit; a pair of elliptic leaf glands Although considering the two genera congeneric, embedded in the base of the lamina (see photo in Baillon (1872, 1875, 1876) never made the proper Alford, 2003)...... transfer of Aphaerema to Abatia, because he did not lb. Placentation parietal; perianth deciduous in fruit; a pair of circular leaf glands projecting associate from the the epithet of Aphaerema spicata Miers with lamina-petiole junction or apex of the petioleAbatia (see (ICBN Art. 33.1, Greuter et al., 2000). photo in Alford, 2003). Aphaerema spicata is thus transferred formally to 2a. Inflorescence a panicle; seed(s) l(orAbatia 2) perhere, but because that epithet is blocked in fruit ... Pleuranthodendron Abatia by Abatia spicata (Turczaninow) Sleumer, 2b. Inflorescence a raceme, this sometimes con gested and fascicle- or umbel-like; seeds a substituteper name is introduced (cf. ICBN Art. 11.2 fruit numerous. 11.4, Greuter et al., 2000). 3a. Raceme terminal, congested and fascicle or umbel-like; fruit large, spherical (> 3 cm diam.); petals 3 or 4; Brazil Abatia . . angeliana M. H. Alford, nom. nov. Replaced ...... Macrothumia name: Aphaerema spicata Miers, Proc. Roy. Hort.

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Soc. London 3: 295. 1863. probablyTYPE: the simple Brazil. reduction Parani:of the laminar part of Sdo Jeroinimo da Serra, the stipule,1861, leaving J.only theWeir terminal tooth.427 A similar (lectotype, designated phenomenonhere, hasK been 000187414; observed in Azara (Charlton, duplicates, BM, K). 1994). The one reliable character differentiating Xylosma and Priamosia is stamen number. A lectotype is chosen because Miers did not Urban and Ekman in Urban (1930) later described designate a holotype. He referred to a single collection Xylosma microphyllum Urban & Ekman based on in the protologue, J. Weir 427, of which one specimen a sterile specimen. This species turned out to be exists at BM and two at K. Thus, the three specimens a synonym of Priamosia domingensis Urban, indicating are syntypes. All specimens are of excellent quality, that the morphological similarity between Xylosma and and the particular specimen at K is chosen due to the Priamosia could confuse even the botanist who amount of material available on the sheet. described Priamosia. Later authors, including Barker Etymology. The epithet honors and Dardeau Jodo (1930), Alberto Sleumer (1980), Angely and Liogier (1917-), Brazilian botanist and (1981), editor have maintained of the segregation small of the and genera, short-lived series Flacourtiaceae: but Sleumer Se'rie (1980: cientifica127) recognized that devo Priamosia is tada ao estudo geral das plantas "very closedestafamilia. to Xylosma." Female or sterile specimens of Priamosia domingensis are virtually indistinguishable KEY TO THE GENERA OF SALICACEAE from WITH Xylosma, OPPOSITE and analyses OR of plastid WHORLED DNA data place LEAVES it clearly within that genus (Alford, 2005). Because la. Receptacular disk glands present; Priamosia stamenswas distinguished often based on inone unreliable fascicles opposite petals; ovary character half-inferior (stipule presence) ....and a single reliable ...... Homalium character (stamen number, 4), which is highly variable, lb. Receptacular disk glands absent; stamens never in fascicles opposite petals; ovary superior. albeit not overlapping, in the closely related genus 2a. Petals absent; inflorescence a many-flowered Xylosma (8 to numerous [ca. 50] stamens), there is no raceme; style not parted at apex; New clear World reason to maintain the two genera. Thus, I propose ...... Abatia the transfer of Priamosia to Xylosma, and the needed 2b. Petals present; inflorescence a few-flowered cyme; style 2- or 3-parted at apex; combination South is provided here. Africa . Pseudoscolopia Xylosma doniingensis (Urban) M. H. Alford, comb. PRIAMOSIA nov. Priamosia domingensis Urban, Repert. Spec. The monotypic genus Priamosia Urban from Nov. Regni Veg. 15: 412. 1919. TYPE: Hispan Hispaniola was described in 1919. Urban suggested iola [Dominican Republic]. Civ. Santo Domingo: Cordillera Central, prov. de la Vega, Constanza, that it is allied with Xylosma and perhaps Azara Ruiz & Pav6n, genera that also have axillary inflores hillsides, pastures, thickets, ca. 1200 m, very cences, receptacular disk glands (lacking in some common, 10 Nov. 1949, E. L. Ekman H14082 Azara), and no petals, characteristics typical of the (neotype, designated here, NY; duplicates, A not tribe Flacourtieae (cf. Warburg, 1893; Gilg, 1925; seen, F not seen, G not seen, GH not seen, K not Lemke, 1988). Urban (1919) pointed out that Azara seen, LL not seen, S, US not seen). differs from Priamosia in having leaf-like stipules and Xylosma microphyllum Urban & Ekman, Ark. Bot. 23A(5): that Xylosma differs from Priamosia in having more 88. 1930. TYPE: Haiti. Massif des Cahos prope Las stamens. Xylosma has eight to numerous (ca. 50) Caobas ad Chapelle Ste.-Claire solo detritu calcario stamens, while Priamosia has only four (Sleumer, obtecto cr. 850 m alt., E. L. Ekman H5546 (holotype, S 04-192; isotype, S 05-961). 1980). Priamosia and Xylosma are also usually dioecious and have armed branches, while Azara is A neotype is designated because the holotype (H. usually bisexual and never armed (Sleumer, 1980). Eggers 2285, Bf) was destroyed in World War II. Sleumer (1980) and Liogier (1981) further recorded Photos of the holotype exist at F and NY, but no that Priamosia has stipules while Xylosma does not, isotypes have been found. Very few quality specimens but this is inaccurate. Although leaf-like or scale-like of Xylosma domingensis exist, but Ekman H14082 stipules do not occur in Xylosma, except perhaps in (NY) was chosen as neotype because the collection the short shoots of Xylosma bahamense (Britton) comes from the same area as the holotype ("Valle de Standley, stipules essentially reduced to single glands Constanza") and has numerous duplicates. do occur in numerous species (Alford, pers. obs.). Because stipules of Salicaceae are sometimes glan Acknowledgments. I thank Melissa Luckow, Sue dular toothed, a gland in a stipule's position is Zmarzty, Robert Kiger, Lucia Lohmann, and an

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anonymous reviewer Greuter, W., J. McNeill, F. R. Barrie, H. M. forB rdet, V. discussions and suggestions leading to this Demoulin, T.paper, S. Filgueiras, D. H. Nicolson, P. C.William Silva, J. Dress for proofreading the Latin description, E. Skog, P. Trehane, N. J. Turland & D. L. HawksworthAlice Calvente for preparing the (editors). 2000. International Code of Botanical Nomen abstract in Portuguese, Bobbi Angell for the excellent clature (Saint Louis Code). Regnum Veg. 138. illustration, Hickey,and L. J. & J. A. Wolfe.Arne 1975. The bases of angiosperm Anderberg, Luis Fernando da Conceicao, Lucia phylogeny: Vegetative Davila morphology. Ann. Missouri Bot.Freire de Carvalho, Mia Ehn, Tom Wendt, Gard.and 62: 538-589. the curators of BH, BM, FSU, K, LL, MO, NY, RB, Hutchinson, and J. 1967. The SGenera offor Flowering Plants. the study of their herbarium Dicotyl dones. Vol. II. Clarendon Press, Oxford. specimens. I especially thank the Harvey Fellows Kuhlmann, J. G. 1953. Flacourtiacea [sic]: Neosprucea Program of the Sleumer n.g. Arq.Mustard Serv. Florest. 7: 3-7. Seed Foundation, the Andrew Mellon Foundation, Leaf Architecture Working Group (A. Ash,the B. Elllis, L. J. Graduate School of Cornell University, and Hickey, K. Johnson, the P. Wilf & S. Wing).Harold 1999. Manual of F. Moore, Jr., Fund of the L. H. Bailey Hortorium, Leaf Architecture: Morphological Description and Cate which provided funding for gorization of Dicotyledonous and Net-Veined Monocoty this study. Thisledonous Angiosperms. Smithsonianwork Institution, Washing was part of a dissertation submitted in partial ton, D.C. fulfillment of the requirements for a Ph.D. in plant Lemke, D. E. 1988. biologyA synopsis of Flacourtiaceae. Aliso 12: at Cornell University. 29-43. Lillo, M. 1919. Rese a fitogeogr fica de la provincia de Literature Cited Tucum n. Estud. Bot. Presentados Primera Reuni n Nac. Soc. Nat. Tucum n 1916: 210-233. Alford, M. H. 2003. Claves para los g neros de Flacourtia ceae de Per y Liogier, del A. H. 1981. AntilleanNuevo Studies I. Flora of Hispaniola: Mundo. Arnaldoa 10: 19-38. -. 2005. Systematic Part 1, Celastrales, Rhamnales, M lvales, studiesThymeleales, in Flacourtiaceae. Ph.D. Dissertation, Cornell Violales. Phytologia Mem. 3. University, Ithaca, New York. Angely, J. 1962. Sleumer, H. 0. 1936. berFlacourtiaceae die neue Flacourtiaceengattung do Brasil meridional. Estudo e revis Spruceanthus. o Notizbl.do Bot. Gart.genero Berlin-Dahlem 13: Aphaerema. Delimita o de 362-363. sua rea geogr fica na Am rica do Sul. Flacourtiaceae 6: 1-L -. 1938. Die Gliederung der Flacourtiaceae-Prock B illon, H. E. 1872. Notes sur les Bixac es. Adansonia 10: iinae Gilg. Notizbl. Bot. Gart. Berlin-Dahlem 14: 45 248-260. 52. -. 1875. The Natural History of Plants, Vol. IV. -. 1950. Algunas flacourti ceas sudamericanas. Lilloa Reeve, London. 23: 247-251. -. 1876. Dictionnaire de Botanique, Vol. I. Hachette, -. 1978. New combinations and a new name in Paris. neotropical Flacourtiaceae. Blumea 24: 118. Barker, H. D. & W. S. Dardeau. 1930. Flore d'Ha ti. Service -. 1980. Flacourtiaceae. Pp. 1-499 in C. T. Rogerson Technique du D partement de l'Agriculture et de L'En (editor), Flora Neotropica, Monograph No. 22. New York seignement Professionnel, Port-au-Prince. Botanical Garden, New York. Charlton, W. A. 1994. Elaboration of stipular structures in Standley, P. C. 1931. Studies of American Plants V. Publ. R. & P. (Flacourtiaceae). Acta Bot. Neerl. Field Mus. Nat. Hist., Bot. Ser. 8: 293-398. 43: 373-382. Urban, I. 1919. LVIII. Sertum antillanum. VII. Repert. Spec. Chase, M. W., S. Zmarzty, M. D. Lled , K. J. Wurdack, S. M. Nov. Regni Veg. 15: 397-415. Swensen & M. F. Fay. 2002. When in doubt, put it in -. 1930. Flacourtiaceae. In Plantae Haitienses et Flacourtiaceae: A molecular phylogenetic analysis based Domingenses novae vel rariores a cl. E. L. Ekman 1924 on plastid rbch sequences. Kew Bull. 57: 141-181. 1928 lectae. VIII. Ark. Bot. 23a(5): 82-89. Gilg, E. 1925. Flacourtiaceae. Pp. 377-457 in A. Engler Warburg, 0. 1893. Flacourtiaceae. Pp. 1-56 in A. Engler & (editor), Die nat rlichen Pflanzenfamilien, 2nd ed, K. Prantl (editors), Die nat rlichen Pflanzenfamilien, Vol. Vol. 21. Duncker & Humblot, Berlin. 3(6a). Wilhelm Engelmann, Leipzig.

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