Three New Species and A Nomenclatural Synopsis of Urera (Urticaceae) From Mesoamerica DCJMB4038

Three New Species and a Nomenclatural Synopsis of "Urera" (Urticaceae) from Mesoamerica Author(s): Alexandre K.
Monro and Alexander Rodrguez Reviewed work(s): Source: Annals of the Missouri Botanical Garden, Vol. 96, No. 2 (Jul., 2009), pp. 268-285 Published by: Missouri Botanical Garden Press Stable URL: http://www.jstor.org/stable/40389934 . Accessed: 19/01/2013 13:34
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THREENEWSPECIESANDA NOMENCLATURAL SYNOPSIS OF URERA FROM (URTICACEAE) MESOAMERICA1

Abstract
AlexandreK. Monro2and AlexanderRodrguez3
Urera Gaudich. is unique among Mesoamerican Urticaceae in having bright, fleshyfruits.Within Mesoamerica, there is confusionover the application of manynames, especially U. corallina (Liebm.) Wedd., U. elata (Sw.) Griseb., and significant U. eggersiiHieron. Three new species, U. fenestrataA. K. Monro& Al. Rodr. (Costa Rica and Panama), U. guanacastensis A. K. Monro & Al. Rodr. (Costa Rica), and U. lianoides A. K. Monro & Al. Rodr. (Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Peru, and Bolivia), are described and illustrated on the basis of stami nate flowers, staminate and habit. The affinitiesof the new species are discussed. The firstrecord of ant inflorescences,stem, leaf morphology, associations forthe genus is documentedin relationto U.fenestrata.In addition,a key is presentedto the 10 species of Urera U. recognized forMesoamerica; nomenclaturalreview is given in which U. mitis(Veil.) Miq. is lectotypified; baccifera(L.) Gaudich. ex Wedd., U. caracasana (Jacq.) Griseb., U. mitis,and Urtica nitida Veli, are epitypified; and Urera denticulata and a list is providedof morethan Miq., U. eggersii,U. subpeltataMiq., and U. subpeltatavar. morifoliaMiq. are neotypified; 900 exsiccatae from13 herbaria.
Resumen
Urera Gaudich. es un gnero nico entre las Urticceas de Mesoamerica por presentarfrutos lustrososy suculentos. En Mesoamerica existe una confusinsignificativa la aplicacin de muchos de los nombres,especialmente en las especies U. en corallina (Liebm.) Wedd., U. elata (Sw.) Griseb. y U. eggersiiHieron. Tres nuevas especies, U. fenestrataA. K. Monro& Al. Rodr. (Costa Rica y Panama), U. guanacastensis A. K. Monro& Al. Rodr. (Costa Rica) y U. lianoides A. K. Monro& Al. Rodr. (Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Peru, Bolivia), son descritas e ilustradassobre la base de sus florese inflorescenciasestaminadas,tallo, morfologa hojas y hbito. Las afinidades de las nuevas especies son discutidas. de Se registrapor primervez la asociacin de una hormigacon el gnero,especficamenteen U. fenestrata.Adicionalmentese presenta una clave para las 10 especies reconocidas por Urera en Mesoamerica; se entregauna revisinen la cual U. mitis (Veli.) Miq. es lectotipificada;U. baccifera(L.) Gaudich. ex Wedd., U. caracasana (Jacq.) Griseb., U. mitisy Urticanitida Veli, son epitipificadas; Urera denticulata Miq., U. eggersii, U. subpeltata Miq. y U. subpeltata var. morifolia Miq. son neotipificadas;y se suministrauna lista de ms de 900 especmenes existentesen 13 herbarios. Key words: Flora Mesoamericana, IUCN Red list, Mesoamerica, Urera, Urticaceae.
The genus Urera Gaudich. comprisesshrubs,trees, and vines that occur most frequently riparianand in disturbed vegetation in Mesoamerica. Within the Urticaceae, Urera is characterized by fleshy fruits of (formedby the inflation the tepals), penicillate or capitate stigmas,glabrous pistillodes, and hairs with bulbous bases thatare stinging some species. Urera in has a nearly pantropical distribution(Neotropics, Africa, Australasia, and the Pacific Islands) but is absent from Asia (pers. obs.). Currently,a single species, U. kaalae Wawra, has a Critically Endan-
gered (CR) status,accordingto IUCN Red List criteria (IUCN, 2001). WithinMesoamerican Urticaceae, Urera is unique in having brightfleshyfruits. is also characterized It stems thatfrequently release a waterylatex when by cut (a trait shared with Myriocarpa Benth.) and, in some species, stinging, bulbous hairs. It is forthese stinginghairs that the genus is most widely known, hence the widespread local name of "chichicaste," which is derived fromthe Nahuat word "tsijtsikast" meaning "to vibrate" (Bonilla A., pers. comm.). It is
1We are to grateful NormanRobson (BM) forhelp withthe Latin diagnoses; Charlie Jarvis(BM) and Sandra Knapp (BM) for commentson the manuscript;Roy Gereau (MO) and Melanie Wilmot-Dear(K) forreviewingthe manuscript;RosemaryWise Victor Steinmannforsending images of type materialat IEB; and the curatorsat BM, C, F, GH, (OXF) forthe illustrations; INB, K, LL, MEXU, MO, NY, P, PMA, TEX, and US forthe loan of,and access to, collections. Some of the paratypematerial was collected withsupportfromDarwin Initiativegrant15/027. 2 Departmentof Botany,The Natural HistoryMuseum, London, SW7 5BD, United Kingdom,a.monro@nhm.ac.uk. J Instituto Nacional de Biodiversidad,ApartadoPostal 22-3100, Santo Domingode Heredia, Costa Rica, arodrig@inbio.ac.cr. doi: 10.3417/2006121
Ann. Missouri Bot. Gard. 96: 268-285. Published on 7 July 2009.
Volume 96, Number2 2009
Monro& Rodrguez New Species and Synopsis of Urera
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also fromthese stinginghairs that Urera derives its limitedeconomic and medical importance:U. baccifera (L.) Gaudich. ex Wedd. is used as "livingfences" in Guatemala and Costa Rica (Standley& Steyermark, 1952; Burger,1977) and is also the focus of research for its anti-inflammatory properties(Badilla et al., 1999). In addition, Urera includes species that are used to treatarthritis (Gonzales, 1994; House et al., 1995), fever (House et al., 1995), hemorrhage (Gunchez, 1999), erysipelas (Gunchez, 1999), and syphilis(House et al., 1995), and species that are of as moderateimportance food forLepidoptera(Janzen & Hallwachs, 2005). The genus Urera was firstdescribed by Gaudichaud-Beaupr in 1830, for which he proposed the subtribe Urerinnae (as Urereae) of the family Urticaceae, which was later raised to tribal rank (Urereae) by Weddell (1856). This was subsequently renamed Urticeae by Friis (1989). Urera was expanded to include the monospecificgenus Scepocarpus Wedd. by Friis (1989), and chloroplastDNA sequence data (trnL-F)suggests that Urera could be sister to Poikilospermum Zipp. ex Miq. within the Urticaceae (Monro,2006). since attention no Urerahas attracted monographic Weddell (1856, 1869), and, to date, no subgeneric classificationhas been published (althoughWeddell did divide the species intounnamedgroupsaccording and distribution). to inflorescencestructure Systemlocalized atic workin Urerahas largelyresultedfrom floristic treatmentsin and adjoining Mesoamerica 1952; Burger, 1977; Pool, (Standley & Steyermark, 2001; Steinmann, 2005). 138 published species epithets There are currently Plant Names Index, forthe genus (The International 2008), of which 133 appear legitimate. Of these to epithets,16 have since been transferred Laportea Gaudich., Gyrotaenia Griseb., Dendrocnide Miq., BoehmeriaJacq., and GirardiniaGaudich. by subsequent authors. Of the remaining 117 names, Friis (1989) estimates that there are ca. 35 good species and Pool (2001) estimates35 to 75. WithinMesoamerica,thereare 25 specificepithets for 10 recognized species of Urera, and there is significantconfusion over the application of many names, especially U. corallina (Liebm.) Wedd., U. elata (Sw.) Griseb., and U. eggersii Hieron. This, combined witha lack of regional keys, the similarity of form and habit, and the extent of overlapping variationin charactersbetweenspecies, has made the of determination collections difficult.The result is that a significantproportionof herbariummaterial Mesoamerica is misidentified. from
Materials and Methods This workwas undertaken partof the revisionof as Mesoamerican Urticaceae for the Flora Mesoamericana project. The definitionof Mesoamerica is as given in the Flora (Davidse et al., 1994): a region bounded to its north the Mexican statesofYucatn, by Campeche, Tabasco, Quintana Roo, and Chiapas, and to its south by the Panama-Colombia border. In addition, material from Mesoamerica and areas adjacent to Mesoamerica (Oaxaca and Veracruz [Mexico], Greater Antilles, Colombia, Venezuela, Ecuador, and Peru) fromBM, C, F, GH, INB, K, LL, MEXU, MO, NY, P, PMA, TEX, and US was examined, resulting in 995 collections that were examined and determined,850 from Mesoamerica. Determinations are listed in Appendix 1. The nomenclatural revisionwas based on the examination of the original published descriptions for all 24 accepted names, as well as typematerial. The macro-morphologicalcharacters used most of frequently previous authorsforthe delimitation by species are leaf shape, leaf margin morphology, inflorescence morphologyand structure,trichome fruit size and color, distributionand morphology, cystolith shape, and stigma shape. In this study, and emphasis was also placed on stem morphology withpersonal observationsin the field habit,together that some species release a thin,waterylatex when cut. Althoughno referenceto this was found in the descriptions on collection labels, a number of collections had a dark stain on the rim of the cut stem. This was taken as an indication that these collectionsmayhave released a watery latex, and it is forthis reason thatthis was noted in the observations made and in the descriptionsbelow. All materialwas at examined using a stereomicroscope X64 to X400 were made and magnification, up to 138 observations foreach specimen sampled. These observationswere then used as a guide to delimit taxa within a species concept and in the preparation morphological of of a key to the identification the Mesoamerican species. Once species were delimited,theywere then matchedto type material. TaxonomicTreatment Urera Gaudich., Voy. Uranie 496. 1826 [1830]. TYPE (designated by Britton& Wilson, 1924: 243): Urera baccifera (L.) Gaudich. ex Wedd., Ann. Sci. Nat., Bot., sr. 3, 18: 199. 1852. Wedd.,Prodr.(DC.) 16(1): 98. 1869. TYPE: Scepocarpus
manni Scepocarpus Wedd.
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Annals of the Missouri Botanical Garden
Keyto the Mesoamerican Species Urera of of with Distributions each speciesare givento country levelwithin for Mesoamerica, theexception Mexico and speciesthatare onlyknown in case they givento statelevel. Global are a from singlecountry, which Flora distributions the key are givento regional in West Indies),following level (e.g., SouthAmerica, in are Mesoamericana keys, (Davidseetal., 1994).Someofthecharacters thefollowing e.g.,cystoliths, protocols are on visible dried in of and material. Several only types hairs (i.e.,bulbous, straight, curved) found thesurface oftheleavesandyoung and HereI define bulbous hairs those as with swollen a stems Mesoamerican of Urera. inflated base giving base a bulbous bottle-shaped the or appearance.
la. Leaveslobed;stem l.U. laciniata white latexwhencut releasing Ib. Leavesnotlobed;stem latexgray latexor not;when released, (never white). releasing 2a. Stem leaves 2-3.2 mm; and with the ca. leavescoarsely dentate, teeth tepals spacedevery 0.5-1 cm;achene spines; 1. U. baccifera the or of to covering basal 1/4 less ofachene prior theinflation tepals 2b. Stemand leaves lacking or whentoothed teeth the crenate, entire, spaced < 0.5 cm spines;leaves dentate, of of to achene0.75-2 mm; 3/4 apart; tepalscovering or more theacheneprior theinflation thetepals. 3a. Adaxialsurface leavessparsely of or pubescent glabrous. 4a. Leafmargin or hairsnever crenate, crenate-serrate, sinuate, entire; urticating. shallowly of hollow. or section stemnever 5a. Shrub smalltree;leaf-bearing without domatia theaxilsof in 6a. Ratioofleaflength width than abaxialleafsurface to less 4:1; 4. U. glabriuscula fruits ororange orange-red to thesecondary veins;mature fleshy pink with in 6b. Ratioofleaflength width to than greater 4:1; abaxialleafsurface domatia theaxilsof fruits red 5. U. guanacastensis thesecondary veins;mature fleshy or section stem of ca. 5b. Shrub, shrub, vine;leaf-bearing lax hollow, 5-10 mmdiam. the 0.125-1 mm; flowers ca. 7a. Young shoots to staminate pubescent, hairs pubescent densely 8. U. lianoides 1.25 X 1.25 mmimmediately to anthesis prior or the 0.125-0.25 mm; 7b. Youngshootssparsely pubescent glabrous, hairswhenpresent 3. U.fenestrata staminate flowers 2 X 2.5 mmimmediately to anthesis ca. prior or or 4b. Leafmargin hairsurticating noturticating. serrate, crenate, dentate; prominently or 8a. Youngstemglabrous sparsely the pubescent, hairs ^ 0.25 mm;stemcoarselysulcate, inflorescence bulbous staminate fenestrate; leaves,and petioles hairs; stems, frequently lacking 3. U.fenestrata to 110 mm;mature fruits fleshy red-pink 8b. Youngstem to the stem but sulcate never coarsely sparsely densely pubescent, hairs> 0.5 mm; with without or to bulbous inflorescence stems, hairs;staminate fenestrate; leaves,and petioles 80 mm;mature or fruits fleshy orange orange-pink. 9a. Tertiary of to venation abaxialleafsurface cream pale green, lamina; noticeably palerthan or stemsparsely 6. U. killipiana forked notforked; stipules pubescent 9b. Tertiary of or the where venation abaxialleafsurface darker rarely palerthan lamina, paler thanthe laminapale brown orange-brown color;stipules forked; to in not stemdensely pubescent. never or 10a. Leavesovateor cordiform, obovate lanceolate; hairsurticating 2. U. caracasana 10b. Leaves obovate, or hairsurticating or lanceolate, elliptic, ovate,nevercordiform; not 9. U. simplex 3b. Adaxialsurface leavespubescent densely of to pubescent. Ila. Leaves bllate; staminate at peduncle unbranched base for 40-80 mm; pistillatepeduncle unbranched base for 27-98 mm at 10. U. verrucosa lib. Leavesnotbllate; staminate pistillate at and unbranched base for< 25 mm. peduncles 12a. Leavesovateor cordiform, obovate lanceolate; never or hairsurticating .... 2. U. caracasana 12b. Leavesobovate, or never hairs or cordiform; urticatingnot . . . 9. U.simplex lanceolate, elliptic, ovate, Ureradenticulata 4: Miq., Fl. Bras. (Martius) 192. 1853. TYPE: Brazil. MinasGerais: Chacha Viosa,rd.E from 10 Fazendada Creciuma, May1930, Y. Mexia valley, 4679 (neotype, MO here,BM!; isotype, not designated seen). Urera var. Arch. Mus.Hist. Wedd., baccifera hrrida (Kunth) Nat. 9: 151. 1856. Basionym: Urtica hrrida Kunth, Nov.Gen. Sp. [HBK] (quarto ed.) 2: 4L 1817. Urera hrrida 4: (Kunth) Miq.,Fl. Bras.(Martius) 192. 1853. BM!). TYPE: Colombia."Santander, MagdalenapropeAn& Urera 4: gosturade Carare," Humboldt Bonpland 1639 armigera Presi)Miq., Fl. Bras. (Martius) 192. (C. 1853. Basionym:Unica armigeraC. Presi, Bot. designated de Rooij [1975: 302], P!). (lectotype, by Bemerk. Presi): 110. 1844 [1845]. TYPE: Brazil, Urtica nitidaVeli.,Fl. Flumin. Icon. 10: t. 20. 1827 [1831]. (C. "near Rio de Janeiro," Lhotsky s.n. (lectotype, TYPE: Fl. Flumin. Icon. 10: tab.20. 1827. EPITYPE: J. Brazil.MatoGrosso Sul: Mpio.Paraguai, do Serrades designated de Rooij [1975: 302], PR notseen). by 1. Urera baccfera (L.) Gaudich. ex Wedd., Ann. Sci. Nat., Bot., sr. 3, 18: 199. 1852. Basionym: Unica bacciferaL., Sp. PL, d. 2, 2: 1398. 1763. TYPE: Plumier, PL Amer.: tab. 260. 1760 (lectotype, designated by de Rooij [1975: 302]). EPITYPE: Jamaica. Stony Hill, 13 Mar. 1898, Fawcett 7177 (epitype, designated here,
271
FazendaCurrupira, Araras, 1510'S, 565'W, 24 Jan. 1995, Dubs 1770 (epitype,designatedhere, K!; E duplicates, notseen,ESA notseen). Urticagrandidentata Liebm., Kongel. Danske Vidensk. ser. Selsk. Skr.(Trondheim) 5, 2: 296. 1851. TYPE: Costa Rica. Cartago:"Irasu," Jan. 1848, A. rsted 14283 (lectotype, by designated de Rooij [1975: 302], C!). An epitypeis selected forUrera bacciferabecause the type illustration, although accurate, is not sufficient support the unambiguous fixingof the to name to this species. Fawcett 7177 was chosen and vegetativematerial because it includes flowering to thatconforms Linnaeus' (1763) description"Urtica the West Indies, foliis alternis,cordatis" and is from as was the Plumier materialthat formedthe basis of the type illustration. An epitypeis selected forUrticanitida because the to althoughaccurate,is notsufficient typeillustration, the unambiguousfixingof the name to this support species. Dubs 1770 was chosen because it includes floweringand vegetative material that conformsto Vellozo's description and is from Brazil, as was the material that formed the basis of the type illustration. A neotype is selected for Urera denticulata because, althoughde Rooij (1975: 302) cites Martius s.n. at M as lectotype for U. denticulata, no such collections could be traced either at M (M. Esser, pers. comm.). The pers. comm.) or BR (P. Stoffelen, collections database fromthe Field Museum's Deof partment Botany(Field Museum of NaturalHistory, of 2006), whichincludes the negativesof photographs material taken by Macbride, does, European type however,include a negative(no. 8845) cited as being a from Martiuscollectionthatrepresents typematerial of U. denticulata at M. Such a negative cannot be considered isotypematerial,but it does indicate the existence of a type specimen existingpriorto World War II. It may be that de Rooij based his on lectotypification this photographor that he did see the type collection at M prior to 1975, and that this collection has since been lost. The specimen designated as neotype was selected because it material and includes good leaf, stem, and fruiting is fromthe same countryand state as the material cited in Miquel's description.
US), nigua, niguilla (El Salvador: P. C. Standley & 22394, GH), ortiga(Costa Rica: R. Anderson S. Mori 147, F; Panama: P. C. Standley30536, US), ortigade los caballos (Mexico: A. Schott 796, BM, F), rscate bien (Honduras: A. Molina R. 868, F, GH). Habitat and distribution. Evergreen or seasonal fromsea level to 1400 m, forest, riparianvegetation, Mexico to Panama, Colombia, Peru, Bolivia, Brazil, Paraguay. Comments. This species most closely resembles Ureralaciniata Goudot. ex Wedd. These species can be distinguished from each other based on the presence of latex and spines and leaf margin morphologyas follows: (1) for U. baccifera, leafbearing stem releasing gray but never white latex when cut, leaf margin coarsely dentate; (2) for U. laciniata, leaf-bearing stem releasing white latex when cut, leaf margindeeply lobed or laciniate. examined. BELIZE. Cayo: ChiquiSelected specimens 671 (BM, bul,Las Cuevas,1643'N,8859'W,A. K. Monro BRH, MO). BOLIVIA. Cochabamba: A. M. Bang, N. L Britton H. H. Rusby1209 (A, BM, GH, MO). COSTA & on RICA. Alajuela: Ro Sarapiquat bridge rd. to Colonia 9 del Viejo hwy., Virgen Socorro, mi.SE ofSan Jos-Puerto 1016'N, 8411'W, T. B. Croat 68307 (BM, MO). EL de SALVADOR. La Libertad: Cordillera Balsam,San A. the Julin towards Pacific, 1341'00"N, 8938'32"W, rd., K. Monro al. 3676 (BM,ITIC,LAGU,MO).GUATEMALA. et del Izabal: Montaas Mico,4-5 kmW ofSantoTomsde et W. Castilla, D. Stevens al. 25612 (BM,MO). HONDURAS. 26275 (MO). Atlntida:1542'N, 8651'W, R. L Uesner 20 Ro NICARAGUA. San Juan:NearCaoChontaleo km 3589 (BM,MO). D. NE ofEl Castillo, Neill& P. C. Vincelli C. & A. PANAMA. Darin: Ro Cocalito, Whitefoord Eddy 249 (BM, MO). PARAGUAY.Canindey:Karapa,Saltoa P. B. & G. 5 kmdel puesto, Marn, Jimnez M. Chocarro 763 Distr. Huancabamba, (BM,FMB).PERU. Pasco: Oxapampa, limite Sector ParqueNac. Yanachaga-Chemillen, Grapanazu, & E. R. Rojas,K. Meza,J. Lingan, Camavilca M. Villaran 1832 (BM, MO). 2. Urera caracasana (Jacq.) Griseb., FI. Brit.W. I. 154. 1859. Basionym: Urtica caracasana Jacq., PL Hort.Schoenbr.3: 71. 1798. TYPE: Jacquin, PL Hort. Schoenbr. 3: t. 386. 1798. EPITYPE: Venezuela. Araugua: Tovar, 1854-1855, A. Fendler 1275 (epitype,designated here, K!).
Ureraalceifolia(Poir.) Gaudich.,Voy. Uranie,Bot. 497. Urtica 1826 [1830].Basionym: Poir., alceifolia Encycl. Local names. Bringa mosa (Panama: C. WhiteGuyana. Suppl.4: 227. 1816.TYPE: French (Lamarck) s.n. foord & A. Eddy 249, BM), chichicaste (Guatemala: /. s.d.,Martin (lectotype, by designated de Cayenne: A. Steyermark 38770, F; Honduras: P. C. Standley Rooij [1975: 304], P!). Nov.Gen.Sp. [HBK] (quarto Urtica Kunth, ed.) 2: tiliifolia 20526, F; El Salvador: P. C. Standley 22344, F; Bolvar: TYPE: Colombia. 34. 1817, as "tiliaefolia." chichicaste Nicaragua: P. C. Standley 11202, F), & Humboldt Bonpland1633 (lectoRo Magdalena, cuyanigua (El Salvador: P. C. Standley 21880, GH, by designated de Rooij[1975: 306], P!). type, Urera Wedd.,Arch.Mus. (Kunth) US), chichicastenigra(El Salvador: S. Caldern 1539, jacquinii var. ulmifolia Urticaulmifolia Hist. Nat. 9: 145. 1856. Basionym: cow itch (Belize: P. H. Gentle2781, F, GH, NY, NY),
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Kunth,Nov. Gen. Sp. [HBK] (quarto ed.) 2: 141. 1817. TYPE: Colombia. Bolivar,Humboldt& Bonpland 1427 (lectotype,designated by de Rooij [1975: 306], P!). Urera mitis(Veil.) Miq., Fl. Bras. (Martius) 4(1): 191. 1853. Basionym: Urtica mitis Veil, Fl. Flumin. 10: tab. 19. 1827 [1831]. Ureracaracasana var. mitis(Veil.) Wedd., Prodr.(DC.) 16: 90. 1869. TYPE: Fl. Flumin. Icon. 10: tab. 19 (lectotype, designated here, tab. 19!). EPITYPE: Brazil. Amazonas: Marapata, Municipality of 4568 (epitype, Carauary, 25 May 1933, B. Krukoff designated here, BM!). Urera corallina (Liebm.) Wedd., Prodr. (DC.) 16: 90. 1869. Basionym: Urtica corallina Liebm., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk.Math. Afd.,ser. 5, 2: 295. 1851. TYPE: Costa Rica. Alajuela: Monte Aguacate, rsted 14282 (lectotype,designated by de designation here, C sheet Rooij [1975: 304], further
and regionas the the same country and is from holotype. var. A neotype selectedfor Urerasubpeltata is de becausealthough Rooij(1975: 306) cites morifolia collection no 633 fitting Spruce (M)as lectotype, Spruce at either M (M. Esser, couldbe traced thisdescription or Miquel pers. pers.comm.) BR (P. Stoffelen, comm.). cites two description, (1853: 190), in his original in "Martius from Brazilian the collections Amazon, and ad do silvisamazonicis, Barra Rio Negro" "Spruce either of couldbe traced neither which ad Santarm," or at M (M. Esser,pers.comm.) BR (P. Stoffelen, pers. at located A was,however, comm.). Sprucecollection Prov. "In BM with annotation, vicinibus the Santarm, the on as Par."Thisis selected neotype thebasisthat "26/2003/2"!). Urera capitata Wedd., Ann. Sci. Nat., Bot., ser. 3, 18: 201. andstate is the collection from samecollector, country, 1852. TYPE: Bolivia. Yungas: Dec. 1846, Weddell fruit and and includes goodleaf,pistillate, immature 4317 (holotype,P!). material. Urera subpeltata Miq., Fl. Bras. (Martius) 4(1): 189, pl. 66. because for A neotype selected Urera is 1853. Urera jacquinii var. subpeltata (Miq.) Wedd., subpeltata Arch. Mus. Hist. Nat. 9: 145. 1856. Urera caracasana s.n. de although Rooij(1975: 306) citesMartius at M var. subpeltata(Miq.) Wedd., Prodr.(DC.) 16: 90. 1869. no for as lectotype U. denticulata, such collections TYPE: Brazil. Bahia: 1839, Blanchet 927 (neotype, or at either M (M. Esser,pers.comm.) couldbe traced designated here, BM!; isotype,G not seen). The specimen BR (P. Stoffelen, comm.). Urerasubpeltatavar. morifoliaMiq., Fl. Bras. (Martius)4(1): desigpers. 190. 1853. TYPE: Brazil. Par: Santarm, was selectedbecause it includes natedas neotype July1850, R. Spruce s.n. (neotype,designated here, BM!). material and and goodleaf,stem, pistillate staminate Urerajacquinii var. miquelii Wedd., Arch. Mus. Hist. Nat. 9: as the and andis from samecountry state thematerial 145. 1856. Urera caracasana var. miquelii Wedd., citedin Miquel's1853 description. Prodr. (DC.) 16: 90. 1869. TYPE. Peru. Gay s.n. De Rooij (1973: 306) citesMakins.n. (nottraced) P not seen). (holotype, Urera acuminata Miq., Fl. Bras. (Martius) 4(1): 190. 1853, for as type(holotype?) Urera jacquiniivar.miquelii. nom. illeg. non Urera acuminata (Poir.) Gaudich. cites only Claude Gay's Weddell (1856), however, and from Peru in his description this is collection of caracasana Jacquin's original description Urera maintained holotype. as is based solely on staminatematerial.Original Local names. Chichicaste (El Salvador: E. material thenamehas notbeen locatedat BM or for havebeenat W has Sandoval 1854, BM; Nicaragua:P. C. Standley that LINN,and anymaterial may blancopicante(El Salvador: beendestroyed couldnotbe traced 10712, F), chichicaste probably (material at W). Basedon thetype itis 504, MO), chichicaste illustration, notpossible E. Sandoval& R. Chinchilla todistinguish E. between caracasanaand U. corallina cujaniguade altura(El Salvador: Sandoval& R. U. Wedd.on sterile characters alone. Material Chinchilla chichicaste 1182, MO), (Liebm.) rojopicapica(El examined thathad been determined U. corallina Salvador: O. Martnezs.n. (ISF225), MO), pan as (includingthe holotype)by Weddel and as U. caliente (Honduras:C. Nelsonet al. 3955, BM), caracasanadid notuncover significant any morpho- migirillo (Costa Rica: A. Sanchez 10, F), miguito differences. epitype selected An is becausethe (CostaRica: /. A. Echeverra 268, F), ortiga C. logical (Costa to is notsufficient Rica: /. A. Echeverra 268, F), pan caliente(El C. illustration, accurate, type although the of 113, F), zulsimtezla support unambiguous application thenameto Salvador:M. L van Severen thisspecies. A. G. (Mexico: Mndez 8945, BM). A lectotype designated UrticamitisVeil, is for Habitat and distribution. Cloud forest,shade becauseVellozo(1827) does notrefer theplateas to coffeeforest, QuercusL.-Liquidambar pine forest, as typematerial his description predatesthe type h-Pinus L. from levelto 2300 m. Mexico sea forest, Tabulae 19 can be considered concept. type original El Belize,Guatemala, Honduras, Salvador, (Chiapas), material.An epitypeis selected for Urera mitis Costa Rica, Panama, Colombia,Venebecause the typeillustration, is Nicaragua, accurate, although and Argentina. Peru,Bolivia, Brazil, notsufficientsupport unambiguous to ofthe zuela,Ecuador, the fixing name to this species. The epitypewas selected of Comments. Herbarium material thisspeciesis because it includesbothleaves and inflorescences commonly determined Urera corallina. Thisspecies as
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most closely resembles U. verrucosa(Liebm.) V. W. Steinm. The two species can be distinguishedfrom each other based on leaf textureand inflorescence peduncle size as follows:(1) forU. caracasana, leaves chartaceous,staminatepeduncle branched to base or unbranchedat base for2-13 mm,pistillatepeduncle branchedto base or unbranchedat base for2-20 mm; (2) for U. verrucosa, leaves bllate, staminate peduncle unbranched at base for 40-80 mm and densely pubescent,pistillatepeduncle unbranchedat base for27-98 mm.
pubescent, the hairs 0.25-0.5 mm; leaf blade 103320 X 64-220 mm, ovate, broad-ovate, or broadelliptic, chartaceous; adaxial surfacesparsely pubescent or glabrous, the hairs 0.375-0.675 mm,weakly appressed, weakly curved or crooked; the cystoliths punctiformto oblong, radially arranged; abaxial surface sparsely pubescent to pubescent, the hairs 0.25-0.5 mm, appressed, weakly curved; the cystoliths oblong, parallel to veins; primary veins 3, to or veins visible to primary quinternary hexternary the naked eye, the lateral primary veins visible for2/3 of the leaf length;domatia not presentin the axils of Selectedspecimensexamined. ARGENTINA. Jujuy: San the secondaryveins; base cordate or obtuse; margins 9633 (K). BELIZE. S. Pedro,SierraSta. Barbara, Venturi Caracol Maya ruins, 14 km W of Las Cuevas, entireor weaklycrenateto serrate;apex subcuspidate Cayo: 1132 (BM, MO). BOLIVIA. to cuspidate. Peduncular bracts 2-3 mm; bracteoles 1645'N,8807'W, T. Hawkins ca. Cochabamba: Carrasco, 11 kmbelowSehuencas, R. 0.375-0.5 mm; staminateinflorescences1 to 16 per J. 10275(BM,K). BRAZIL.Roraima: Ilhade Marac, I. Wood stem,peduncle branchedto base or unbranchedfrom de S SEMA EcologicalReserva, of Cachoeira Fumaa,W. base for2-21 mm,pubescent,the hairs to 0.125 mm, 546 Milliken (K). COSTA RICA. Puntarenas: W.Haber& the whole inflorescence15-110 mm, bearing 160 to W. Zuchowski9294 (MO). ECUADOR, s. loe., Eggers 15611 rd., (K). Pichincha: newAlluriqun-Quito Km 4, P. J. M. 350 flowersin a symmetrical cyme with3 or 4 orders Maas & L Cobb4770 (K). EL SALVADOR.La Libertad: of dichotomous branching; the flowers borne in P. "Laderas La Laguna," Lemas de Cuscatlan, Mpio.Antiguo clusters of 10 to 35, pedicellate to subsessile, the s.n. "WB-01217"(BM, LAGU, MO). GUATEMALA. Juwhen presentto 0.5 X 0.125 mm. Pistillate tiapa: D. Dunnet al. 23234 (MO). HONDURAS.Ocotepe- pedicels 1 inflorescences to 4 per stem,the peduncle branched de que: Alrededores Beln Gualcho,C. H. Nelson,E. Romero,A. Rubio & M. Pereira 3955 (BM, MO). MEXICO. to base or unbranched at base for 2-26 mm, pubChiapas: Rancho Puy Ukum,sobre la carr. a 2 km de escent, the hairs ca. 0.125 mm, erect, straight, the A. G. Bochil,Mpio.de Bochil, Mndez 8945 (BM, MEXU). whole inflorescence10-80 mm (47-80 mm in fruit), NICARAGUA. Jinotega:"El Recreo,"4 kmal N de Sta. P. & Gertrudis, Moreno J. C. Sandino7897 (BM, HNMN). broaderthanlong (or as long as broad),bearing224 to PANAMA.Chiriqui/Bocasdel Toro: AlongContinental 750 flowersin a symmetrical cyme with 5 orders of S. dichotomous Divide on trailin Zona Palo Seco, 0847'N, 8213/W, branching,the flowersborne in clusters Knapp & J. Mallet 9169 (BM, MO, PMA, SCZ). PERU. of 3 or 4, pedicellate to subsessile, the pedicels when Loreto: Ro Itaya,T. B. Croat18829 (K). VENEZUELA. X 0.125 mm, glabrous. Staminate Falcn: Sierrade San Luis, ca. del Puentede Joboentre present to 0.25 flowers2-2.5 X 1.25-1.5 mm immediatelyprior to 99249 (K). Curimagua San Luis,J.A. Steyermark y anthesis; tepals 4, ca. 3 mm; stamens 2.5-3 mm; ca. 0.5 mm diam., glabrous. Pistil3. Urera fenestrata A. K. Monro & Al. Rodr., sp. pistillode late flowers ca. 1 X 0.5 mm, the lateral tepals nov. TYPE: Costa Rica. Guanacaste: Monte0.5-0.675 mm,asymmetrically ovate; the dorsal tepal verde,Cordillerade Tilarn,Pacific slope, above 0.50-0.675 mm, ovate, with a subapical dorsal Quebrada Cuecha, 1540-1600 m, 6 May 1976, thickening,the ventral tepal ca. 0.375 mm, ovate; V. J. Dryer 179 (holotype, F!; isotype, CR!). stigma penicillate, erect. Fruit prior to inflationof Figure 1A-E. tepals ca. 1.25 X 1 mm;basal 3/4 ofachene obscured caracasanae(Jacq.)Griseb. nova Urerae similis, by tepals, the laterals ca. 1 mm, the dorsal tepal Species ramulis staminatis sed ab ea floribus petiolisque 0.675-0.75 mm, the ventral ca. 0.5 mm; achene tetrapartitis, nervis atque foliorum espinosisramulissaepe fenestratis 0.75-1 X 0.75 mm, asymmetrically elliptic, keeldiffer. visibilibus lateralibus 2/3longitudinis per shaped, surface smooth.Fruit when ripe with tepals ca. 1.5 X 1 mm, red-pink, Shrub, lax shrub, vine, or small slender tree, inflatedand berry-like, or orange when fresh. dioecious (?). Main stems arching2-10 m, stems not red-violet, pink, spines; young releasingwhitelatex when cut; without Local names. Ortiga {A. Smith 100, F, NY). shoots glabrous or sparsely pubescent, the hairs is Habitat and distribution. Urera 0.125-0.25 mm, erect, straight to weakly curved; fenestrata found in and cloud forest, sectionsof stem9-30 X 4r- in premontane, montane, of internodes leaf-bearing frequently at or shadedareas close tostreams smallrivers, disturbed to red-brown, 10 mm, pale brown coarsely sulcate, when ^ 5 mmdiam. 800-3000 m. It occurson thePacificand theCaribbean fenestrate and frequently hollow, de pubescent; slopes oftheCordillera Tilarnin Costa Rica and the Stipules 5-18 mm,lanceolate, not forked, and Panama. de petioles 52-350 X 1.5-1.75 mm,glabrousor sparsely Cordillera Talamanca in Costa Rica
274
Figure 1. A-E. UrerafenestrataA. K. Monro & Al. Rodr. - A. Habit, withstaminate inflorescences.- B. Leaf abaxial surface. - C. Stem. - D. Pistillate inflorescence.- E. Pistillate flower.(A, B: Dryer 179 [F]; C-E: Kirkbride Duke 770 & [NY]). F- I. U. guanacastensis A. K. Monro& Al. Rodr. - F. Habit, withstaminateinflorescences.- G. Leaf adaxial surface. - H. Pistillate inflorescences.- I. Immaturefruit.(F-I: Delgado 24 [MO]).
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for IUCN Red Listcategory.Conservation Urera must considered LeastConcern be as fenestrata (LC) to (IUCN, 2001), according IUCN Red List criteria to the owing thefactthat specieshas beencollected 28 timesin several localitiesin Costa Rica and Panama.
9 mi.from overRo Grande Orosi, Gentry de A. 2043 bridge caminoentre (MO); a la veradel Rio Turrialba, Pacayasy SantaCruzde Turrialba, Gmez-L 8078 (CR); Tausito, of S J. TresRos,cerros la Carpintera, Khan, Tebbs A. R. de R. M. & 1318 (BM); ca. S of Tapanti, de Vickery alongRo Grande 806 R. R 3803 (F, NY); Orosi, W.Lent (CR); Tausito, W.Lent Sector Dos Amigos, Mora543 G. ParqueNacionalTapant, Ro Macho,SectorEl Embalse,G. (INB); ReservaForestal Forestal Macho, Ro sendero atrz Fromthe Latin"fenestratus," mean- Mora643 (INB); Reserva Etymology. V. 393 when da casa a direito, Nilsson& R. Manfredi (CR); El to the stems which, referring ing"windowed," Interamericana L J. Poveda306 (CR); El Sur, Empalme, greaterthan or equal to 5 mm diam., become Guarco, de de Cerro ParqueNac. Tapant-Macizo la Muerte, windowed. la Muerte, 600 m N de Chespirito, Solano 77 (INB); D. characteristically caminoa Moravia, antesdel Ro Pacuare,L O. Turrialba, Discussion. This species is characterized the Williams J.J. Crdoba by 4668 (CR); Guanacaste: Palmira, & and stem windowed, leaf-bearing sections the fog zone,A. Smith100 (F, NY). Heredia: Parque Nac. gnarled, leaves. BraulioCarrillo, indented or to entire subentire weakly Zurquistation, just above Los Guarumos ovate, 2450 B. Carrillo, been has Urera frequently determined trail, Boyle/. (BM,CR); ParqueNac. Braulio fenestrata most sector A. Chacn1881 (CR); La Palma,Ro Bajo be as U. elataorU. caracasanaandmay distinguished de Zurqui, M. BraulioCarrillo Nati.Park,N. Garwood, Honduras, from themas follows: forU. caracasana,stems Gibby, J. Hampshire C. J. Humphries (BM, CR); (1) & 386 R but Quebrada Mochte, diam.hollow, than when I. el borde Cerro de L greater orequal to5 mm Zurqui, D. Gmez, M. 21040 (CR); never fenestrate, lacking spines; petiole lacking A. Chacn, G. Herrera, M. ChavarrCa 323 de A. & R. Rodrguez (CR); or serrate-dentate, Porrosat Barva, Jimnez serrate, spines;the leaf margins of headwaters Ro Cantnde Heredia,Cordillera Central, flowers to crenate-serrate dentate;staminate (4)5- SantoDomingo, slopeofVolcnBarva,M. Grayum 7237 N thanor (INB); S slopeofVolcnBarva,W.H Hatheway 1354 (CR); (2) parted; forU. elata, stemswhengreater 258 Estacin but neverfenestrate, ParqueNac. Braulio Carrillo, Barva,G. Rivera equal to 5 mmdiam. hollow, 259 Ro Porrosat, m N de parada 50 G. Rivera smallspinespresent; petiolewithsmallspines;the (INB), de Paso (INB); Rivera 432 (INB); S slopes of de buses Llano,G. or leaf marginsserrate,serrate-dentate, dentate; VolcnBarbabetw.Ro Cirueles & & Sacramento, Utley J. stems K. U. flowers staminate (3) 4-parted; for fenestrata, sendero 2319 (NY); ParqueNac. BraulioCarrillo, Utley thanor equal to 5 mmdiam. hollow, haciaQuebrada de Zamora625 (CR). Limn: Cantn whengreater Zrqu, Cordillera J. Talamanca, ca. campRo Lori, Bittner with fenestrate prominent 5 mm windows, lacking Talamanca, smallspines;theleafmargins 1882 (INB), 1894 (INB). Puntarenas: Cantnde Buenos lacking petioles spines; Tres fincaBenito Estacin Acua,E. flowers Aires, Cantn Colinas, ZonaProtectora Alfaro staminate to crenate serrate; or entire weakly Las Tablas, de CotoBrus, 745 (INB); E. 1260 (INB); Cantn Las Tablas,sendero Echandi, Alfaro 4-parted. Las have observedants of the genera de CotoBrus,Zona Protectora Tablas,I Camp.ACLA, Both authors near the E. 1569 Lund,and PheidoleWestwood (For- caminoa Cerroechando, Alfaro of (CR); Crematogaster W. Divideca. 2-5 kmE & SE Monteverde, C. of Continental stems thehollow micidae, occupying Myrmicinae) 8639 (CR), 8752 (CR). Puntarenas& Burger J. L Gentry of this species in the field. Collections these ant Alajuela border: 2-5 km E & SE of Monteverde, C. W. Nacional de Burger& R L Liesner 8639 (F); ReservaSanta Elena, are available at the Instituto species 19 D. Monteverde, S. Penneys (CR). San Jos: Vsquezde collections. Biodiversidad entomology a senderofrente Coronado, Parque Nac. BraulioCarrillo, 486 480 L de COSTARICA.Alajuela:La Palma San Estacin Zurqu, Acosta (CR,INB),L Acosta (CR, Paratypes. de INB); valleyof Ro Hondura 136 A. (belowLa Palma),NE of San Ramn, M.Brenes (5544)(CR,F, NY);La Palma & W. 220 A. SanRamn, M. Brenes (4208)(CR,F); rd.toFinca Jernimo, C. Burger R G. Stolze4912 (CR, F); Ro NE belowLa Palma, T. N Zarcero,B. Croat Clarovalley La Hondura off 15, LosEnsayos hwy. ca. 7.5 mi. of drainage), (Ro & W. 44 V. de 43488 MO); (MEXU, CordilleraTalamanca,J.Dryer of San Jernimo, C. Burger R G. Stolze7646 (F); 233 179 V. de (CR);Cordillera Tilarn, J.Dryer (CR), (CR), bosquecillosresidualesentre las Nubes y Cascajal de 5535 SantaElena, Coronado, Gmez-Laurito (CR); nearquebradaBajo J. Reserva de 234 (CR); Cantn San Ramn, 2500 (BM);Acosta 3 s 19 Mquina, kmNE ofCascajal,R W.Lent D. 100 de Cordillera Tilarn, mNWofStation, Penney ca. de Cerros Escaz,El Cedrai, Zona Protectora N (INB); Vara Blancade Sarapiqu, slope of Central Palmichal, Zona A. betw. Cordillera, Pos & Barbavolcanoes, F. Skutch de Ia casa, S. Lobo 661 (CR); Cantnde Aserr, la Escaz- Carpintera, La de Cerros Escaz,Cerros de near Central SanJuan Laja,ca. Protectora 3602 (NY);Cordillera & /. A. L Daser,AltoHierbaBuena, F. Morales L 15 km N of Zarcero, 0. Williams, Molina R, T. P. cimadel Cerro 5866 faldaNW, F. Morales de Bohs3787 (INB);Cerro /. Forestal Cedrai, 28972 (F, NY); Reserva & Williams D. N. Gibson del C, Grecia,cuencasuperior Ro Rosales,G. Herrera G. (CR, INB); Tarraz,San Carlos, Bajos de La Virgen, A. Ros Paraso, ros 542 (BM, INB, MO); above Quaker confluencia Negro Blanco,Albergue Umao& H. Gmez y Finca San 2352 (MO). Quesada1161 (CR); Turrubares, Luis,San Rafael, & at settlement Monteverde, Utley H Utley J. 9313 (INB); Altode la A. Ro de Cartago: Cantn Paraso,ReservaForestal Macho, de MelvinChavarra, Rodrguez 635 & ca. J. 1799 (INB); TapantHydroelectric Palma, de 6 kmN ofSanJernimo,Utley K. Utley Ro Pejivalle,E. Alfaro de ReservealongRo Grande Orosi,T. Croat36151 (CR); (CR). PANAMA.Bocas del Toro: Chiriqutrail betw. & & J. zum quebrada von Higuern Gutierrez, H. KirkbrideJ.A. Duke Richtung Orosi, farhweg Rio Macho,in westlicher of at I.C.E. Reservation, 770 (NY); Caribbean Fbrega foot "Falso slopesofCerro 5162 (CR); Tapanti P. Stausee, Dbbeler
276
A. 4855 Fbrega," K. Mojito& S. Cafferty (BM, INB, MO, 4856 (BM,INB,MO,PMA).Chiriqui:vie.ofGualaca PMA), ca. 10.7 mi.from La Planesde Homito, Fortuna to dam rd. T. 5111 (BM);betw. Bambita Cerro & site,T. Antonio Punta, B. Croat 10593 (MO);betw. Palo Alto& topofridge (divide) et nearCerro PateMachoaboveRo Palo Alto, D'Arcy al. W. 12658 (BM); Boquete, M. 349 Bajo Chorro, E. Davidson (F); near Fortuna & Dam,R. J.Hampshire C. Whitefoord214 (BM), 203 (BM);lower of A. reaches trail Cerro to Pando, K. Monro, S. Knapp, Mallet, Mallet, Mallet V.Mallet A. I. & 3520 (BM, J. 257 & Velo,R. E. Woodson R. W.Schery (MO); MO);quebrada & vie.ofBajo Mona& quebrada R. Chiquero, E. Woodson R. W. Schery of 593 (GH, MO); Bajo Mona,mouth quebrada P. & Ro R. Chiquero, along Caldera, E. Woodson, H. Alien R. 1005 (MO,NY). J.Seibert
3 June1990, R. Delgado 24 (holotype, INB!;

isotype,MO!). Figure 1F-I. sed ab ea Wedd. similis, simplici Species nova Urerae foliis ramulis angustioribus supraparcepubescentiglabris, subtus domatiis bus vel galbris praeditis, pedicellisflorum differ. rubro breviorbus staminatorum atquefructu
Shrub to small tree,dioecious(?), main stems whitelatex,without 2-A m, not releasing arching, of shootsglabrous;internodes leafspines; young sections stem of 3-10 X ca. 2 mm, pale greybearing not brown, hollow, lackinga darkstainon the cut of portion the stem. Stipules 5-12 mm, narrowly 6-90 forked, lanceolate, pubescent; petioles sparsely toward tosparsely 4. Urera glabriiiscula V. W. Steinm., Acta Bot. X ca. 0.5 mm, glabrous pubescent Mex. 71: 22. 2005. TYPE: Mexico.Veracruz: leaf base, the hairs 0.25-0.375 mm,strongly apleaf blade 55-230 X 8-55 mm, Mpio. San AndrsTuxtla,Volcn San Martn, pressed,straight; 1300 m,2 Apr.1985,CedilloT. 3175 (holotype, narrowly lanceolate, lanceolate to oblanceolate, to adaxialsurface chartaceous subcoriaceous; MEXU notseen). sparsely IEB!; isotype, toward toglabrous, hairsmost the frequent pubescent Localnames. K'analzulzimtez A. 0.375-0.5 mm, (Mexico: Mndez theleafbase,hairs weakly appressed, G. 9054, BM). the inflated, densely straight; cystoliths punctiform, abaxial surfaceglabrous;the cystoliths scattered; Habitat and distribution. Montane deciduforest, to inflated, occasionally appearing ous and evergreen forest, Liquidambar-Taxodium punctiformoblong, veins3, and scattered parallelto theveins;primary Rich,forest from Mexico 100-2800 m. El Salvador, veins to and quarternary primary tertiary occasionally Guatemala. Oaxaca,Tabasco,Veracruz), (Chiapas, veins visibleto the naked eye, the lateralprimary Comments. Urera glabriuscula is most similar to finer and visiblefor1/3of leaf length; thanmidrib U. lianoidesin its glabrous sparsely or in pubescent domatia veins, present theaxils of thesecondary leaves thatare entire discretely or divided. The two composed hairs;base obtuse;margins of asymmetrifrom basedon cally discretely to crenateto serrate; speciescan be distinguished eachother apex pungent and achene subacuminate. habit,stem,stipule,staminate flower, 1-2 mm; Peduncular bracts bracteoles as shrub 0.25-0.5 mm,staminate ca. inflorescences 15 per (1) morphology follows: forU. glabriuscula, or small tree, leaf-bearing sectionof stem never stem,peduncleunbranched from base for6-9 mm, with 20 less hollow, internodes than mm; apex pubescent, hairsto 0.25 mm, wholeinfloresthe the stipule 0.75 mm diam.; achene cence 5-40 mm, forked; minutely pistillode in ca. bearing 110 flowers a weakly surfaceverrucate; (2) for U. lianoides, vine or asymmetricalsymmetrical with or 5 orders 4 to cyme section stem of hollow of dichotomous shrub, scrambling leaf-bearing the subsessile, branching; flowers at ca. 5-10 mmdiam.withinternodes than borne clusters 5 to 7; pistillate greater 2 in of inflorescences 20 mm;stipule 0.375 mm to21 perstem to7 in fruit), peduncle apex notforked; pistillode the branched (4 smooth. diam.;achenesurface tobase orunbranched base for at 2-12 mm (6-17 mm in fruit), the sparsely pubescent, hairsca. 0.1 mm, Selected examined. GUATEMALA. specimens Quezaltethe 5-40 inflorescence mm 34292 (F). MEXICO. Chiapas: erect, (23straight; whole nango: J. A. Steyermark G. 30 mm in fruit), ColoniaMaravilla, Mndez A. thanbroad (or as long as Mpio.Maragritas, Tenejapa, longer 9054 (BM, MEXU). Oaxaca: Hector Hernndez A. M. y in 30 (59 broad), bearing to250 flowers to67 in fruit) Chacn 487 (MO). Tabasco: Mpio. Teapa, en Cerro 4 a symmetrical with or5 orders dichotomous of cyme Universidad Madrigala 500 m al E de Puyacatengo, flowers borne clusters 3, in of pedicellate, Autnoma E. S., Chapingo, M. Martnez J. ColonicoSoto, branching, A. M. Hanan-Alipi, A. Hernndez, Martinez N. rarely2 to 4, the pedicels subsessileto 0.25 X M. A. & 34699 (BM,MEXU).Veracruz: A. Gentry, Lott 0.175 mm,glabrous.Staminate E. Peregrino 1-1.25 X flowers & UNAM class32227 (BM, MO). tropical botany 1.50-1.75 mmimmediately priorto anthesis; tepals 4, ca. 2 mm; stamensand pistillodenot seen. 0.4-0.6 X 0.25-0.6 mm,lateral 5. Urera guanacastensisA. K. Monro Al. Rodr., Pistillateflowers & nov.TYPE: CostaRica. Guanacaste: ca. 0.4 X 0.25 mm, ovate;dorsaltepalca. 0.4 sp. Parque tepals NacionalGuanacaste, de Cantn Liberia,Esta- X 0.25 mm, ovate, the ventraltepal ca. 0.4 X cin Cacao, 1055'45"N,8528'15"W,1100 m, 0.25 mm, erect.Fruit ovate;stigma prior penicillate,
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to inflation tepals withachene entirely of obscured by tepals, the laterals ca. 1.75 mm,the dorsal ca. 1 mm, the ventralca. 0.675 mm, achene 1-1.375 X 0.751.25 mm,elliptic,surfacesmooth, fruit whenripe with 1-1.25 X ca. 2 mm, tepals inflatedand berry-like, red. Habitat and distribution. Ureraguanacastensis is found in montane and cloud forest,disturbed and undisturbed forest, at 820-1350 m, known only fromthe Cordillera de Guanacaste in the Cantn de Liberia, Guanacaste ConservationArea in Costa Rica. IUCN Red List category. Conservationfor Urera guanacastensisis consideredas Near Threatened(NT) according to IUCN Red List criteria (IUCN, 2001). This is based on an evaluation of the potential of distribution U. guanacastensis, the threat to its habitat withinthat area, and the numberof existing records for the species. Extrapolatingfromthe 12 collection localities of U. guanacastensis plotted on Google Earth, this species' potential distribution covers an area of ca. 680 km2. All 12 records are from forestlocalities (Google Earth, 2008; collection label data) that formpart of Costa Rica's protected ca. areas' network.Currently, 50% of the potential is distribution deforested (Google Earth,2008) and no localities. We therefore deforested recordsexist from assume that ca. 50% of the original population has been lost and that the future of this species is dependent on the maintenance of Costa Rica's ProtectedAreas Network.
domatia; pedicels of staminate flowersgreater than 1 mm; mature,fleshyfruit orange. Paratypes. COSTA RICA. Guanacaste: Cantn de Cordillera Guanacaste, de Liberia, ParqueNac. Guanacaste, Estacin Cacao, sendero Maritza,L ngulo88 (INB); EstacinCacao, bosquesprimarios orillasde bosque,W, y E. Bello 2237 (INB); EstacinCacao, C. Chvez95 (INB); de de ParqueNac. Rincn La Vieja,Cordillera Guanacaste, Estacin Pailas,R. Espinoza771 (INB); Estacin Las Cacao, sendero casa de Fran, Gamboa (CR, INB); Estacin a B. 45 VolcnCacao, sendero entre Estacin potrero Los Mengo, y Naranjos,W, B. Hammel& E. Chavarra17536 (BM); sendero Potrero, sur,//INBio 183 el EstacinMengo, lado (BM, INB); EstacinCacao, cerroCacao, senderoa Casa Fran, A. Mora37 (INB); Estacin W, Cacao,cerro Cacao,M. Moraga 60 (INB); Estacin Cacao, CerroCacao, W, F. Las Quesada275 (INB); Sector Pailas,Ro Colorado, Aguas de Arriba,G. Rivera651 (INB); Cordillera Guanacaste, sendero lagunaSantaMara,W, G. Rivera a 1152 (INB). 6. Urera killipiana Standl. & Steyerm.,Fieldiana, Bot. 24: 427. 1952. TYPE: Guatemala. Quetzaltenango:Volcn Junil,1700 m, 8 Aug. 1934, A. F. Skutch982 (holotype,F!; isotype,GH!). Local names. Chichicaste (Guatemala: P. C. Standley 76980, F), chichicaste comn (Guatemala:
P. P. C. Standley 64714, F), nigita (Guatemala: C. 75549,F). Standley
Habitat and distribution. Disturbedvegetation, from montane riversides, 900-3000 m,Mexico forest, (Hidalgo, Queretaro,Oaxaca, Tabasco, Chiapas), and Guatemala, El Salvador. Comments.Urerakillipianais most frequently
Etymology. Urera guanacastensis is named after determined U. or as eggersii(= U. simplex) U. the Guanacaste ConservationArea, where all known from caracasana.These speciescan be distinguished recordsof this species have been collected. Discussion. Ureraguanacastensis is characterized lanceolate, lanceolate, or oblanceolate, by narrowly to sparsely pubescent leaves thatfrequently glabrous possess domatiaon the abaxial surfacecomposed of a clusterof hairsin the axils ofthe secondaryveins,and fruits (notunique fleshy by the red color of the mature the within genus). It is mostlikelyto be confusedwith whichit maybe distinguished U. simplexWedd., from by leaf width, pubescence, and the presence of domatia, as well as by the length of the staminate pedicels and color of the fruitas follows:(1) for U. guanacastensis, young stems glabrous; leaves 838 mmwide, adaxial surfaceverysparselypubescent to glabrous,abaxial surfacewithdomatia in the axils of the secondaryveins; pedicels of staminateflowers less than 0.5 mm; mature,fleshyfruit red; (2) for U. stems pubescent, frequently densely simplex,young so; leaves 37-210 mm wide, adaxial surface pubescent, frequently densely so, abaxial surface lacking
and basedon venation, each other stipule morphology, as stemindumentum follows: forU. killipiana, (1) or stems stipule apexforked sparsely pubescent, young veinsof the abaxial leaf the not forked, secondary in domatia their venation without surface axils,tertiary the ofabaxialleafsurface noticeably than lamina; paler stems U. densely pubescent, young (2) for caracasana, veinsof the notforked; secondary the stipuleapex with abaxialleafsurface frequently domatia (flap-like venation of of or tufts hairs)in theiraxils, tertiary tone as darker ofthesamecolor or abaxialleafsurface stemsdensely the lamina;(3) forU. simplex, young the veins forked; secondary apexnot stipule pubescent, in without domatia their of the abaxial leaf surface or of darker venation abaxialleafsurface axils,tertiary ofthesamecolortoneas thelamina.
examined. EL SALVADOR. San Selectedspecimens San 727 (GH, NY). GUATEMALA. Salvador: 5. Caldern 15338 (MO). Finca Armenia, Marcos: San Marcos, Dwyer de MEXICO. Chiapas: Ocozocoautla Espinosa,T. B. Croat
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40599 (MO).Hidalgo: Puerto Obscuro nearKm328 onhwy. betw. SantaAna& Chapulhuacn, H. JacalaDistr., E. Moore 5049 (BM). Oaxaca: B. Hammel M. Merello & 15457 (MO). B. Queretaro: 3^1 kmal S de La Parada,Mpio.de Jalpan, Servin 557 (BM). Tabasco: C. L Gilly& E. Hernandez X. 320 (GH, MEXU). 7. Urera laciniata Goudot. ex Wedd., Ann. Sci. Nat., Bot., sr. 4, 18: 203. 1854. TYPE: [Colombia] Nouvelle Grenade. "Quindui" [Quindio?], La Bolsa, 1844, /. Goudots.n. (lectotype, designated by de Rooij [1975: 308], P #00281783!). Unica girardinioides Seem., Bot. Voy. Herald 194. 1854. TYPE: Panama.1846-1849,B. Seemann 494 (lectoby type, designated de Rooij[1975: 308], BM!). Both epithets,Urera laciniata and Urtica girardinioides,were published in 1854. Accordingto Stafleu and Cowan (1988: 139), the probable month of publicationforWeddelFs publicationis March,while that Seemann'spublicationis July for (Stafleu& Cowan, 1985: 476), thereby to givingpriority Ureralaciniata. Local name. Pringamoza (Panama: J. A. Duke 9266, MO). Habitat and distribution. Ureralaciniata has been collected fromriversidescrub and fromsea level to 1200 m. Its range extendsfrom Honduras,Nicaragua, Costa Rica, Panama, Colombia, and Peru to Bolivia. Comments. The deeply lobed laciniate leaves and large (2-2.5 mm) asymmetrical fruit are unique amongst Mesoamerican Urera, and this species is unlikelyto be confusedwithany other. Selected examined. BOLIVIA. La Paz: Franz specimens senda Azariamas- Fermin San tamayo, ParqueNac. Madidi, sector E A. C. G. Mutun, Ticona, Arajo V. Torrez, Perez, M., & 149 Jove A. Urbano (BM,MO). COSTA RICA. San Jos: A. F. Skutch 4266 (GH, MO,NY). HONDURAS.Gracias a Dios: J. Saunders1204 (NY). NICARAGUA.Jinotega: CaoLitutus, Bocay,W.D. Stevens, H. Beach, Schal Rio J. J. & 0. M. Montiel 16650 (BM, MO). PANAMA, loc: s. J. 1846-1849, B. C. Seemann494 (BM). PERU. Junin: El Cataract Tirol, K. Monro, T. Pennington A. Daza A. R. & 3993 (BM,MOL). 8. Urera lianoides A. K. Monro & Al. Rodr., sp. nov. TYPE: Costa Rica. Alajuela: San Miguel Oeste, Naranjo, subiendo por ladera sur del Cerro EsprituSanto hasta bosque residual en el flanco noreste del mismo, 1005'20"N, 8424'20"W, 1000-1200 m, 24 Nov. 1988, G. Herrera 2326 (holotype, INB!; isotypes, BM!, MO!). Figure 2A-F. V. similis, Species novaUrerae glabriusculae W. Steinm. sed ab ea habitu in caulis5-10 mm lianiformi, foliifera parte
non diametro cava,stipulis apice integris furcatis, pistillodio minore atqueacheniolaevi differ. Vine, scramblingshrub,dioecious (?); main stems cane-like, to 25 m, youngshootspubescentto densely pubescent, the hairs 0.25-1 mm, erect or weakly appressed, weakly curved or crooked, occasionally straight;intemodes of leaf-bearingsections of stem 21-84 X 3.5-10 mm,red-brown, occasionally yellowgreen, hollow where 5-10 mm diam., lacking a dark stain on the cut portion of the stem. Stipules 412 mm, narrowlyovate or lanceolate, not forked, sparsely pubescent to pubescent; petioles 11-90 (-170) X 0.75-1.5 mm, sparsely pubescent to pubescent, the hairs 0.125-1 mm,appressed, occasionally erect, curved, occasionally crooked or straight; leaf blade 72-280 X 29-110 mm, narrowlyovate, obovate, elliptic, or oblanceolate, always longerthan wide, chartaceous and occasionally bllate; adaxial surface sparsely pubescent, the hairs 0.5-0.75 mm, appressed, weakly curved or crooked; the cystoliths occaoblong or occasionally fusiform, punctiform, scatteredand occasionally sionallyinflated, randomly parallel to veins, rarely arranged radially around a hair base; abaxial surfacepubescent,the hairs0.1250.5(1) mm, appressed or erect, weakly curved, occaoccasionally straight;the cystolithsfusiform, sionally oblong, parallel to the veins, occasionally veins 3, occasionally 5, randomlyscattered;primary to veins visible to the naked eye, primary quinternary the lateral primary veins visible for1/2-2/3ofthe leaf length; domatia not present in the axils of the secondaryveins; base subcordateor obtuse,occasionally cunate; marginsdentate, occasionally crenatedentate or nearly entire; apex cuspidate. Peduncular bracts 1.25-2 mm; bracteoles 0.25-0.5 mm. Staminate inflorescences 1 to 12 per stem, peduncle branched to the base or unbranched fromthe base for 4.5-9 mm, densely pubescent, the hairs to 0.25 mm,the whole inflorescence 19-56 mm,bearing 192 to 384 flowers an asymmetrical in cymewith4 or 5 orders of dichotomousbranching;flowersborne in clustersof6 to 12, sessile; pistillateinflorescences ca. 6 per stem, unbranchedbase of peduncle 7-13 mm, 5-11 mmin fruit, denselypubescent,the hairs0.1250.25 mm,erect,straight; whole inflorescence12the 40 mm (12-45 mm in fruit), broader than long, bearing 112 to 448 flowersin an asymmetrical cyme, with 4 to 6 orders of dichotomous branching; the flowers borne in clusters of 3, pedicellate, the pedicels 0.5-0.675 X 0.125-0.25 mm, glabrous. Staminate flowersca. 1.25 X 1.25 mm immediately prior to anthesis; tepals 4, 1-1.5 mm; stamens ca. 1.75 mm; pistillode ca. 0.375 mm diam., glabrous. Pistillate flowers0.5-1 X 0.375-0.5 mm, white to yellow-green;lateral tepals 0.375-0.75 mm, asym-
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Figure 2. Urera lianoides A. K. Monro& Al. Rorir.- A. Habit withstaminateinflorescences.- B. Cluster of staminale flowers. C. Staminateflower anthesis.- D. Leaf ahaxial surfacewithcystoliths. E. Habit, withpistillateinflorescences. at - F. Immaturefruit.(A-D: Johnson1600 |GH]; K, F: Shank & Molina R. 4405 [NY]).
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metricallyovate, dorsal tepal 0.25-0.75 mm, ovate, ventraltepal 0.25witha subapical dorsal thickening; 0.75 mm, ovate, stigma penicillate. Fruit prior to inflationof tepals ca. 1.5 X 0.75 mm; basal 3/4 of achene obscured by tepals, lateral tepals ca. 1 mm, dorsal tepals ca. 0.75 mm,ventraltepals ca. 0.5 mm; achene 1.25-1.5 X 1-1.25 mm, asymmetrically elliptic,keel-shaped,surfacesmooth.Fruitwhenripe, withtepals inflatedand berry-like 1.5-2.25 X 1.251.75 mm,orange when fresh. Habitat and distribution. The new species is found in premontane,montane,and cloud forest,in disturbed and undisturbedforest,fromsea level to 1300(1900-2500) m. Mexico (Chiapas), Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Peru, and Bolivia. IUCN Red List category. Conservationfor Urera lianoides must be considered as Least Concern (LC) according to IUCN Red List criteria (IUCN, 2001), owing to the fact that the species has been collected 50 timesin several localities across Centraland South America.
Cordillerade Tilarn, Reserva Biologica Monteverde, de Refugio Eladio, Valle del Ro Peas Blancas,E. Bello 5793 (F, A. 5181 (INB); La Palmade San Ramn, M. Brenes A. La NY); MataCartago, Palmade San Ramn, M. Brenes 6308 (F, NY); quebradaAzul (San Carlos),A. M. Brenes El 23063 (NY); Guatuso, ParqueNac. VolcnTenorio, Piln, a colectadoen bosque primario orillas de senderoLos de 874 del Misterios Tenorio, L Chvez (INB); Cantn Los /. Ro Cao Negro, de Natural Vida Silvestre Chiles,Refugio & Fro,Finca Betel,K. Flores M. Flores130 (INB); Reserva Ro Biolgica Monteverde, Peas Blancas, Finca Wilson W. Atlntica, Haber& E. Bello 7403 (INB, Salazar,vertiente W. Poco Sol 13 km S Fortuna, MO); ReservaMonteverde, 9377 (BM,INB); pala,Bijagua,El Haber& W.Zuchowski 1293 (BM,INB); G. Piln,cabeceradel Ro Celeste, Herrera R. N side ofVolcnArenal, W.Lent2922 (F); pala,Zona aboveRo 3 Protectora Miravalles, kmW ofBijagua,ridges & Zapote,D. S. Penneys W.Haber710 (INB); ParqueNac. G. FincaJulio La de Rincn La Vieja,Colonia Libertad, Soto, Rivera1485 (INB); Colonia Blanca, Finca Ro Negro,G. Cerro Rivera1563 (INB); San Carlos,ParqueNac. Arenal, V. A. Chato,sendero que lleva a la Laguna, Rodrguez, H.
Ramrez, G. de La 0 & G. Soto 6254 (INB); Colegio
Santa Clara, Cantnde San Carlos,A. S. Agropecuario,

Weston,D. F. Weston& J. Weston3104 (MO). Cartago:
Cantn de Turrialba,Valle del Reventazn, Carolina, P. Moraviade Chirrip, Campos176 (INB); 1/2km S of Chitara, near rd. #CR233, R. W. Lent 237 (NY). de Cordillera Tilarn,1de Guanacaste: Cantn Tilarn, Etymology. From the English "liana" or "vine," 2 kmW of Lago de Cote,13 kmN ofTilarn, continental derived fromthe French, "lier," which, in turn, is divide on SE, slope of Volcn Tenorio,W. Haber & W. Finca 11624 (INB). Heredia: L. R. Holdridge's to Zuchowski the derivedfrom Latin "ligare" or "to tie," referring El La Selva,Ro Puerto Viejo at quebrada Sura& quebrada the liana-like habit of this species. G. El Salto,ca. 1 mi.abovejet. Ro Sarapiqui, B. Rossbach R. 35 bankabout kmNE ofAlajuela, J. Discussion. Ureralianoides is distinguished its 3690 (GH); roadside by & 4536 (NY). Limn: Level areas betw.Siquerres scandent habit, hollow stems, sparse non-urticating Taylor overtheRio bridge steephillsS oftherailroad fruit. Material of U. Rio Pacuare, hairs, and orange,mature,fleshy Pacuare, W. C. Burger & R. L Liesner6952 (NY); borde de fromothercollections lianoides fromChiapas differs et lago DabagrihastaRo Liei,L D. Gmez al. 23129 (BM); of this species in having densely pubescent young Costado et hastaRo Liei,L D. Gmez Oestede lagoDabagri el sobre Ro ToroAmarillo, al stem and abaxial leaf surfaces,with very shorthairs al. 23172 (BM);vecino puente M. 5.5km al SO de Gupiles, A. Jimenez 1067 (F); (0.5 mmor less). This species correspondsto "sp. A" de pantanosos-yolillosos Suerrey Dos Bocas, drenajesde in the treatment Urera in the Flora de Nicaragua of P. los RosParismina Reventazn, J.Shank& A. MolinaR. y (Pool, 2001: 2495). Urera lianoides most closely 4265 (GH); pantanosos-yolillosos de de Goldengrove, drenaje resembles U. glabriuscula with which it shares a P. Ro Reventazn, /. Shank & A. Molina R. 4405 (F). Forestal Golfo de Puntarenas: Canton Osa, Refugio Dulce, in distribution Chiapas, Mexico. The two species can BahaChai,entrada de Chocuaco, be distinguished from each otheras follows:(1) forU. Pennsula Osa, Los Mogos, de of 3591 (INB); foothills Cordillera Talamanca, R Aguilar lianoides, a vine or scramblingshrub, ca. 5-10 mm G. S just N ofSantaElena on Fila Cotn, ofAguaCaliente, diam., leaf-bearingstem section hollow, with inter- Davidse, G. HerreraCh. & M. H. Grayum28277 (BM); Osa, nodes greaterthan 20 mm; stipule apex not forked; Ballena,entrando Puertecito, hastael Cerro J. Escalera, por & Gonzlez C. Aragn2327 (INB); Cantnde Coto Brus, 0.375 mmdiam.; achene surfacesmooth;(2) pistillode de Cordillera Talamanca, La ParqueInternacional Amistad, for U. glabriuscula, a shrub or small tree, ca. 5Estacin Pittier, Agua Caliente,E. Navarro211 (INB); stemsection never hollow, cantnde 10 mmdiam., leaf-bearing Osa, Pennsulade Osa, Rancho Quemado,F. with internodes less than 20 mm; stipule apex Quesada 167 (INB); deep forest 4 near airport, mi. W of P. minutelyforked; pistillode 0.75 mm diam.; achene Rincnde Osa, Osa Pennsula, H. Raven21623 (MO); de Forestal Golfo de Canton Osa, Refugio Dulce, Pennsula surfaceverrucate. del finca Grupo Fila Rincn, Osa, Rancho Quemado, Guerra, M. Piln de Conservacin, Segura 4 (INB); Reserva Biolgica Ballivian, Paratypes. BOLIVIA. Beni: Provincia de San Cordillera Tilarn, Luis de Monteverde, Monteverde, Laja, 130 km de San Borja,M. Moraes670 (K). La Paz: 118 S. ca. Provincia Yungas, 2 kmal S de Coreico, G. Beck bosquesen la cuencadel Ro San Luis,K. Taylor (INB); Nor 1173 & /. 21926 (K); Santa F Tacana, al lado de la parcela Rincnde Osa, area N ofairport, Utley K. Utley de de (F). San Jos: Cantonde Prez Zeledn,Cordillera permanente Santa F, G. Bourdy1746 (K). Tarija: 329 SantaElena,E. Alfaro (INB); W Las R. Provincia Arce,108 kmde Tarijahacia Bermejo, Ehrich Talamanca, Nubes, 494 (K). COSTA RICA. Alajuela: Cantnde San Ramn, part of montaas Jamaica,ca. 3 km NE of Bijagual de
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Cararareserve, H. Grayum al. 5878 (BM); Portobelo M. et S. Turrubares, hwy.to 4 km up Rio Guanche, Knapp1016 Zona Protectora Cangreja, La Mastatal Puriscal, F. (BM);Ro Guache, J.Sytsma de K. 1617 (BM). Darin: Parque /. Morales (INB); Acosta, 727 Fila Ro Nac. Darin,Serrana Sapo, lmite Parquehastala de del Colorado, San Jernimo, H. & S J. Colorado, F. Morales7383 (INB); Acosta,Aserr, Agua cima, Herrera J.Polanco795 (BM).Panam: Piriat, et 558 (BM). PERU. Buena,cabecerasQuebrada Laja, J. F. Morales al. 9937 of Pan-American Hwy., C. Hamilton A. 2863 (GH, MO,NY). Madre de Dios: ParqueNac. del Manu,Rio Manu,Cocha (INB); vie.ofEl General, F. Skutch GUATEMALA. ca. Peten: Dolores, 100 m del cementano, CashuStation, Terborgh R. B. Foster & 6488 (K). J. lado N, R. Tun 0. 1323 (BM, F). Suchite pequez: Finca 1479 (GH). HONDURAS.Atlantida: Moca,A. F. Skutch Vic. of La Ceiba, bank of Danto river, slopes of Mount 9. Urera simplex Wedd., Prodr. (DC.) 16(1): 90. 8454 T. & J. 1869. TYPE. Colombia. Cundinamarca:"ad salto Cangrejal, G. Yuncker,M. Koepper K. A. Wagner (GH, MO, NY). Corts: Aldea de Corinto alrededores y de Tequendama," Mar. 1856, Triaras.n. (holo55 C. frontera Guatemala, km al 0 de Puerto con Corts, type,P 00281785!). 2926A E. M. Nelson, Vargas, Erazo,M. Garca& M. Sierra de (BM, MO). Olancho: Vaguadadel Ro de la poblacin Urera Bot. 20: Hieron., Jahrb. eggersii Syst. 3. 1895. TYPE: Culmi, C. Nelson & E. Romero4738 (MO). MEXICO. Pichincha: kmW ofSantoDomingo los 20 Ecuador. de rd. to Pichucalco, along Chiapas: 2-4 kmbelowIxhuatn 1000 ft., Nov.1961,P. C. D. Cazalet& 25 Colorados, D. 19906 (MO); long Muni,of Solosuchiapa, E. Breedlove T. D. Pennington 5150 (neotype, here,K!; designated 60 gravel rd. betw. Palenque & Bonampak, mi. SE of B isotypes, notseen,NY notseen). rd. 40192 (MEXU,MO); alonggravel T. B. Croat Palenque, V. rzedowskii W. Steinm., Bot.Mex.71: 37. 2005. Acta 88-90 mi.SE ofPalenque,T. Urera betw. Palenque& Bonampak, TYPE: Mexico.Veracruz: Mpio. San AndrsTuxtla, 40230 (MO);alonghwy. B. Croat 195,betw. Chiapade Corzo 8 kmal N de San Andrs Tuxtla, LagunaEncantada, & Pichucalco, 6 mi.NWofPuebloNuevoSolistahuacn, ca. 1828'N,9510'W,350 m,4 Apr.1981,J. I. Calzada Caribbeanslope, Selva Negra overlooking along mirador IEB 8105 (holotype, notseen;isotype, ENCB not seen). vie. at lookout point trailto ColoniaPinabeto, Km99, T. B. Donn. Sm., Bot. Gaz. 23(1): 14. 1897. camino Ureratuerckheimii 65180 (MO);Crucero & Croat D. P. Hannon Corozal, TYPE: Guatemala. AltaVerapaz:Pansamal, 1160 m, S. E. Boca Lacantum, Mpio.Ococingo, Martnez Palenque1243 US May1887,H. vonTuerckheim (holotype, not Corozalsobreel 15439 (BM); a 14 kmal NW de Crucero seen;isotype, NY!). camino Palenque-Boca Lacantum,Mpio. Ococingo,E. Martnez 16642 (BM). Veracruz: camino BastonalS. A neotypewas selected for Urera eggersiibecause T. R. Tebanca, Mpio.Catemaco, Cedillo 3439 (BM);Estacin & M. B. Los Biolgica Tuxtlas, Hammel, Merello S. Sinaca thetypecollectioncited by Hieronymus, Eggers 14466 15492 (MO). NICARAGUA.Chontales: On ridgetop of in (B), was destroyed enemyaction duringWorld War P. A. W. A. Cordillera Amerisque, Gentry, D. Stevens, Grijalva 43950 (MO). Granada: VolcnMombacho, II and only photographsof the holotype could be & P. P. Moreno O. located (F, MO). The neotypewas selected on the basis A. HaciendaUPE-Pancasn (antesdel crter), Grijalva, & R. M. Rueda 2924 (BM); NE del Volcn thatit includes good leaf and fertile materialand was Vanegas a de en Mombacho, el emplame los caminos Sta. Isabel y a from same country the holotype. as the 2602 (BM, MO); Volcn Mombacho, P. Cutirre, Moreno subiendo la Finca Las Delicias,1.5 kmde la hacienda, Local names. Bilsimtezla (Mexico: A. Mndez T. por 4103 (MO); NW slopesofVolcnMombacho, 6738, P. P. Moreno BM), chenek'mut(Mexico: A. Mndez T. 4863, 27693 (MO);NWde M. 10 kmS ofGranada, Nee& J.Miller BM), chichicaste (Guatemala: P. C. Standley 68232, de cafetales FincaCutirre camino VolcnMombacho, que y lleva S del volcn, C. Sandino1273 (BM, MO). Zelaya F; Mexico: M. Heath & A. Long MA 44, BM), J. Sur]: alongtherd.betw. chichicaste huevo de cangrejo (El Salvador: E. [ReginAutnomadel Atlntico & NuevaGuinea& Verdun, 5. Miller J. C. Sandino1094 Sandoval & H. Rivera 1252, MO), sakil zulsimtez J. del la Caoel Pato,1.5 kmsobre ribera Point, (MO);Monkey laa (Mexico: A. Mndez T. 6238, BM), tzotznizzul s. 12397 (MO). PANAMA, loc.:5. Hayes Cao,P. P. Moreno 2008 simtez(A. Mndez T. 9066, BM), zulsimtezla(Mexico: H. Allen 683 (K); Canal area,vie. ofMaddenDam,P. A. Mndez T. 7022, BM). 106 (F); rd.S-ll, Island,S. Aviles (F, NY); BarroColorado NWofEscobal,T. B. Croat12466 (BM, NY); rd.alongRo Habitat and distribution. Disturbed and undisNW partofCanal Zone (area W of Pia-RoMediadivide, sea and humidscrub from cloud forest, 1600 turbedforest, Locks& Gatun Limon Lake),/.M. Johnston Bay,Gatun of lake,NWpart CanalZone level to 2500 m. Mexico (Chiapas, Tabasco, Vera(GH); rd.alongW sideofGatun (area W of LimnBay,GatunLocks& GatunLake),/. M. cruz), Belize, Guatemala, Honduras, El Salvador, et 1706 (GH); 12 mi. S of Coln,E. L Tyson al. Johnston Costa Rica, Panama, Colombia, Ecuador, 4478 (GH). Bocas del Toro: Ridge N of Campamento Nicaragua, 4506 (BM, INB, MEXU, Peru, Bolivia, Brazil. & A. Luchio, K. Monro E. Alfaro MO, PMA); Ro Cricamola,betw. Finca St. Louis & Comments. Materialofthis species has frequently P. R. Konkinto, E. Woodson, H. Allen & R. J. Seibert to and referred (Flora of Guatemala, 1923 (GH, NY); 10-15 mi. inland S frommouthof been determined T. as W.H Lewis, D. Dwyer, S. Elias & Flora de Nicaragua, and Flora Costaricensis) Urera J. River, Changuinola 867 K. R. Robertson (GH, K, UC, US). Chiriqui: Bajo elata, U. eggersii,or U. tuerckheimii. Examinationof 349 (MO). Cod: M. Chorro, BoqueteDistr., E. Davidson typematerialofthese species indicatesthatU. elata is of Foothills Cerro Piln,nearEl Valle,J. Duke& M. Correa to a species endemic to Jamaica, while U. eggersiiand abovevillage farm potato 14670(1) (MO); El Valle from are 7923 (BM). Coln: From U. tuerckheimii conspecificwith U. simplex.Pool & Cerro J. Pilon, Dwyer M. Correa
282
El Costa Rica, Panama, U. may mala, Honduras, Salvador, (2001) indicates tuerckheimii U. simplex) (= to U. aurantiaca Wedd. fromSouth Peru,and Bolivia. correspond America(Argentina, Bolivia, Paraguay);however, Comments. Ureraverrucosa mostclosely resembles thatthe two of comparison the holotypes suggests from can The species bedistinguished U.caracasana. two with characterized by each otherbased on leaf texture speciesaredistinct, U. aurantica and inflorescence ovate cordiform or leavesandrelatively pistillate short leaves size as follows: forU. verrucosa, (1) inflorescences known from SouthAmerica peduncle and only 40at unbranched basefor staminate bllate; peduncle Bolivia, Brazil, and Paraguay).Some 80 (Argentina, mm,denselypubescent; pistillate peduncleuncollections U. simplex CostaRica andPanama, of from U. at 27-98 mm; for caracasana, branched base for (2) 1939 (GH) and Folsomy Page 5984 leaves e.g., Kennedy to staminate branched base chartaceous; peduncle (PMA), are unusualin the possessionof narrowly orunbranched basefor 2-13 mm; at pistillate peduncle oblanceolate leaves, whilesome material branched baseorunbranchedbasefor pubescent at 2-20 mm. to from Chiapas (Purpus7039, NY) is characterized examined. BOLIVIA. La Paz: M. Selectedspecimens by denselypubescentleaves. Urerasimplexmost resembles elata. The twospecies can be Lewis882155 (K). COSTA RICA. Cartago: N of Cartago, U. closely 959 M. Tebbs A. R. Vickery (BM). & R. Ro Reventada, fromeach other based on petiole EL SALVADOR. Khan, distinguished Cordillera Ahuachapn: Lago de Ninfas, and size peduncle as follows: Grande Apaneca,NW ofJuayua, Davidse, Sidwell, K. morphology inflorescence G. de M. 37383 (BM, ITIC, & (1) for U. simplex, petiole lacking small spines; A. Monro, A. Renderos C. Cortez On CA Chimaitenango: hwy. staminate flowers (2) LAGU,MO).GUATEMALA. occasionally 4-parted; 5-parted, to to & turnoffPatzm Solla,14.8 mi.NNWofturnoff for U. elata, petiole withsmall spines; staminate betw. & to 23.5 mi.SSE ofturnoffSolla,T. B. Croat D. P. Patzm, flowers 4-parted. M. 64732 (BM,MO). MEXICO.Veracruz: Orizaba, Hannen
Selected examined. BELIZE. Stann Creek: specimens W. 400 Middlesex, A. Schipp (BM,F, GH,NY). BOLIVIA.La EstacinBiolgica Paz: N Yungas,valle de Huarinillas, S. Tunquini, G. Beck24612 (K, LPB). BRAZIL.Amazonas: E. Ule5465 (K). COLOMBIA.Putumayo:Umbra, Klug G. 1741 (K). COSTA RICA. Puntarenas: Cantonde Golfito Dos Brazos RoTigre, de orilla Quebrada de Pizote, Jimnez, 95 G. Cordero (BM,INB,MO).ECUADOR.Pichincha:P. C. D. Cazalet& T. D. Pennington 5073 (K). EL SALVADOR. A. et Ahuachapn:FincaL'Esperanza, Jujutla, K. Monro al. 2997 (BM, ITIC, LAGU, MO). HONDURAS. Olancho: S. Catacamas,Ro Catacamas, slope of Sierrade Agalto, Blackmore G. L A. Heath1916 (BM). MEXICO.Chiapas: & Finca Mexiquito, A. Purpus7039 (BM, F, MO, NY). C. NICARAGUA. Matagalpa: Macizode PeasBlancas,Finca San Sebastian, TllezV.,R. Riviere, D. Stevens, M. O. W. O. Montiel M. Guzmn D. Castro5181 (BM, MEXU). & J., PANAMA. Comarca de San Blas: UdirbiReserve, along et J. park boundary, F. McDonagh al 257 (BM,MO).PERU. M. Huanuco: Vic. ofTingoMaracliffs aboveRo Monzn, E. Mathias D. Taylor & 5343 (K). 288 Botteri (BM). PANAMA. Chiriqui: 3 kmNW ofCerro 1363 rd. Punta, alongdirt on rte.to Las Nubes,B. Hammel & (BM,MO,NY). PERU. San Martin:T. D. Pennington A. Daza 16676 (K, MOL). Cited Literature
A. Badilla,B., G. Mora, J.Lapa & J.A. SilvaE. 1999.Antiin of (Urticaceae) activity Urerabaccifera inflammatory Biol. Trop.47: 365-371. rats. Sprague-Dawley Revista of Rico and N. Britton, L. & P. Wilson.1924. Botany Porto Rico & Virgin Islands Islands.Sci. Surv.Porto theVirgin 5: 243. in W. Fieldiana, Burger, 1977. Urera. FloraCostaricensis. Bot.40: 276-280. 1994.Introduccin Davidse, M. S. Sousa& A. 0. Chater. G., in Pp. general. xiii-xiv G. Davidse,M. S. Sousa & A. 0. Vol. 6, AlisChater (editors),Flora Mesoamericana, a Universidad NacionalAutnoma mataceae Cyperaceae. St. MexicoD. F.; Missouri Botanical de Mexico, Garden, London. Museum, Louis;The Natural History & de Rooij,M. J.M. 1975. Urera. 301-309 inJ.Lanjouw Pp. Vol. A. L. Stoffers FloraofSuriname, 5(1). E. J. (editors), 10. Urera verrucosa (Liebm.) W. Steinm., V. Acta Leiden. Brill, Bot. Mex. 71: 39. 2005. Basionym:Unica Field Museum of Natural History.2006. The Botany of Collections Database. Department Botany, Field verrucosa Liebm., Kongel. Danske Vidensk. Museumof NaturalHistory, Chicago. <http://emuweb. Selsk.Skr.,Naturvidensk. Math.Afd., ser. 5, 2: php>, accessed3 Septem295. 1851. Urera caracasana var. tomentosa fieldmuseum.org/botany/Query. ber2008. Prodr. 90. 1869,nom. 16: (Wedd.)Wedd., illeg., Friis, 1989.TheUrticaceae: systematic A I. review. 285Pp. Evolu308 in P. R. Crane & S. Blackmore superfl.TYPE: Costa Rica. Cartago: Irasu, (editors), of and tion,Systematics, Fossil History the HamameOersted 14284 (holotype, C!). Association lidae, Vol. 2. Systematics Special Volume United 40B. Oxford Science Publications, Oxford, KingLocalnames. Chichicaste E. 333, (Mexico: Kerber dom. BM). C. Gaudichaud-Beaupr, 1830. Urera. Pp. ^96-497/in par ordre du Voyage autour du Monde, entrepris Habitatand distribution. Montane shade forest, . de roi,. . excutsur les corvettes S.M. VUraieet la coffee farms, forest, Pinus-Quercus-Liquidambar from Physicienne.par M. Louis de Freycinet. .. Botanique. 500-2800 m. Mexico (Chiapas, Veracruz),GuateParis. Pp. 496-^97 Pillet-ain,
283
V. Etnobo- 4. U. glabriuscula W. Steinm. J. Gonzales, C. 1994. BotnicaMedicinal Popular, BotnicoLa 5. U. guanacastensis K. Monro Al. Rodr. A. tnica Medicinalde El Salvador.Jardn & El 6. U. killipiana & Standl. Steyerm. Laguna, Cuscatln, Salvador. accessed12 7. U. laciniataGoudot. Wedd. ex Earth. 2008. <http://earth.google.com>, Google 8. U. lianoides K. Monro Al. Rodr. A. 2008. & April de 9. U. simplex Wedd. F. amaznicas usomedicinal Gunchez, J.1999.Plantas y V. (Liebm.) W. Steinm. para el 10. U. verrucosa mgico.FundacinPolar, ServicioAutnomo Caracas. Desarrollo Ambiental Amazonas, del L 480 (3),486 (3); Aguilar, 387 (10);Aguilar, I. R. Acosta, T. P. L. House, R.,S. Lagos-Witte, Ochoa,C. Torres, Meja& et al. 3591 (8); AldosS., J. 92 (9); Akrasquilla, 180 (10); L M. Rivas. 1995. Plantas Medicinales Comunes de E. P. Alfaro, 329 (8), 745 (3), 1260 (3), 1569 (3); Allen, H. Honduras.Litografa Lpez, S. de R. L. Universidad 911 (cf.2), 926 (1), 1471 (10), 2008 (8), 3617 (9), 6315 (1), de de Internacional Autnoma Honduras/Comit Nacional 6325 (9), 6330 (2), 6946 (10), 855 (cf.2); Anderson, & R. en Internacional Natural Honduras/Cooperacin Medicina S. L T. Mori, 147 (1); Angulo, 88 (5); Antonio, 611 (1), 1118 de Tcnica (Programa Cooperacin para el Desarrollo 1738 (9), 3528 (2), 4988 (9), 5111 (3); Araquistain, M. fr en Gessellshaft Tech- (9), Britnica Honduras)/Deutsche 3182 (9); Araquistain, & Castro, 1905 (1); Araquistain, M. D. nischeZusammenaerbeit, Tegucigalpa. M. & Moreno, P. 1502 (2), 2009 (1), 2235 (1), 2285 (1), P. Plant NamesIndex.2008. <http://www.ipni. International 2358 (9),2473 (9),2605 (9),2790 (9),2863 (1); Atwood, T. J. 2008. ore>, accessed 19 June A. S. & Moore, D. 304 (2), 489a (1); Aviles, 32 (9), 706 (8). Version and IUCN.2001. IUCN Red ListCategories Criteria, R. C. F. 2011 (2); Baker, R53 (2); Bangham, N. W. Commission. Baker, 3.1. Prepared theIUCN SpeciesSurvival by F. E. 211 (9); Barkley, A. & Barkley, D. 40130 (1); Barkley, and Cambridge,United IUCN, Gland, Switzerland, F. F. A. & Hernandez J.40263 (1); Barkley, A. & LeivaR., Kingdom. F. M. L 39579 (2); Barkley, A. & Smith, L 40859 ofthe Welchez, food D. Janzen, H. & W. Hallwachs. Caterpillar plants M. 11 (9); Bello,E. 2237 (5), 3143 (2), 5181 Costa (9); Barrelier, northwestern Area de Conservacin Guanacaste, G. S. J. (8); Bittner, 1882 (3), 1894 (3); Blackmore, & Heath, Rica. <http://janzen.sas.upenn.edu/Wadults/searchfood. B. M. L A. 1916 (9), Botteri, 288 '854' (10); Boyle, 2450 (3); 2005. lasso accessed9 November >, D. Breedlove, E. 9107 (1), 9994 (4), 10003 (5), 79906 (8), ed. C. Linneaus, 1763. SpeciesPlantarum, 2. Stockholm. 20207 (9),23388 (4),24709 (9),24249 (4),24998 (9),34570 in Flora Miquel, F. A. W. 1853. Urera.Pp. 188-193 D. P. (4), 51467 (5); Breedlove, E. & Raven, H. 13559 (4), Vol. Brasiliensis Leipzig. (Martius), 4(1). Fleischer, D. A. 13583 (9); Breedlove, E. & Smith, R. 19905 (9), 21681 A of A. Monro, K. 2006. The revision species-rich genera: D. R. 31520 (4); Breedlove, E. & Thome, F. 30986 (10); of revision Pilea (9), for framework thestrategic phylogenetic A. Brenes, M. 136 (5544) (3), 220 (4208) (3), 339 (9), 3480 and morphology. based on cpDNA,nrDNA, (Urticaceae) (9),3596 (9),5524 (2),5793 (8),6039 (2),6308 (8),6764 (9), Amer. Bot.93: 426-441. J. L E. G. 23063 (8); Brown, 6 (2); Bunting, S. & Licht, 1137 C. In Pool,A. 2001. Urera. W. D. Stevens, Ulloa,A. Pool & W. W. D. (1); Burch, 4603 (9); Burger, C. 4161 (1); Burger, C. Florade Nicaragua. 0. M. Montiel Syst. & Antonio, 10956 (9), 10993 (1); Burger, C. & Baker, Monogr. (editors), W. T. Bot. Bot.Missouri Gard.85: 2492-2495. W. M. R. 10013 (1); Burger, C. & Burger, 7678(9), 8496 (9); 2nd 1985.Taxonomic F. Literature, Stafleu, A. & R. S. Cowan. W. J. Burger, C. & Gentry, L 8639 (3), 8752 (3), 8836 (9), Utrecht/Antwer& Scheltema Holkena, ed., Vol. 5. Bohn, R. W. 8984 (9), 9092 (7); Burger, C. & Liesner, L 6605 (2), The b.v. pen,and Dr.W. Junk Publishers, Hague/Boston. 6879 (1), 6952 (8); Burger, C. & MattaU., G. 4819 (9); W. & H. A. Weddell. 1988. Taxonomic , R. W. Burger, C. & Stolze, G. 4912 (3), 5608 (9), 5904 (2); & Scheltema Holkena, 2nd Literature, ed., Vol. 7. Bohn, W. Burger, C. et al. 10502 (1), 70565 (1). The b.v. and Dr. Utrecht/Antwerpen, W. Junk Publishers, A. S. E. Cabrera, etal. 2666 (4); Cafferty,& Monro, K. 16 Hague/Boston. S. (9); Caldern, 727 (5), 850 (2), 7570 (2), 7539 (1), 7687 In 1952. Urera. Floraof P. Standley, C. & J.A. Steyermark. S., (2), 7775 (2), 7809 (2); Calonico J.etal. 21101 (4),27242 Bot.3: 424^28. Guatemala. Fieldiana, M. (4); Campos,P. (8); Carlson, C. 391 (10), 2056 (1); and a V. W. 2005. Fournewneotropical Steinmann, species P. U I. Chacon, A. 1881 (3); Chavarria, 303 (2); Chvelas, et ActaBot.Mex.71: al. ES-2280 of newcombination Urera (Urticaceae). C. J. (9); Chavez, 95 (5); Chavez, L 874 (8); 19-43. 88 M. A. Chickering, M. 126 (9); Chorley, & R. Atkinson (2); 10. Vellozo,J. M. C. 1827 [1831]. Florae Fluminensis M. E. Christopherson,D. 178 (10); Chrysler, A. 5612 (2); de Janeiro. Rio Nationali, Typographia G. G. J. Conrad, etal. 2796 (4),2875 (9); Cooper, P. & Slater, & H. Weddell, A. 1856. Urera. 143-162 in G. Baudry J. M. 178 (9); Cordero, 95 (9); Correa, D. etal. 2368 (2); Pp. M. G. de (editors), Monographie la Familledes Urtica- Cosentino, 83 (9); Cowan, P. 1721 (9), 3064 (9), 3709 Baudry C. K. Paris. ces. G. and J.Baudry, C. J. (9), 3757 (9); Cowan, P. etal. 3951 (9); Cowell, F. 257 . 1869. Urera.Pp. 199-203 in A. de Candolle (1); Croat, B. 10593 (3), 10597 (10), 12333 (9), 12466 (8), T. Prodromus (DC), Vol. 16. Paris. (editor), 15895 (9), 15897 (9),23728 (9),24254 (9),26599 (9),35138 (9), 36757 (3), 36777 (9), 40192 (8), 40230 (8), 40599 (5), are listed 40862 (9), 40956 (8 aff.), Appendix Index to Exsiccatae. Collections 1. 43488 (3), 43886 (9), 47437 (10), by alphabetically collectorname and then in ascending 49785 (7), 66252 (9), 66612 (2), 66795 (2), 68307 (1),68377 as order.Species are numbered in the list (9), 78493 (2), 78524 (9), 78545 (9); Croat, B. & Hannon, numerical T. here. D. P. 63356 (cf.2), 64732 (10), 65180 (8), 65332 (9), 65354 provided T. D. T. (4); Croat, B. & Porter, M. 15663 (9); Croat, B. & R. (10). Zhu, G. 76527 (9); Cruz, 31 (1), s.n. 'WB-1176' List of Species W. W. D'Arcy, G. 4239 (9), 10767 (9), 10998 (10); D'Arcy, K. W. B. G. & Hammel, 12235 (9); D'Arcy, G. & Sytsma, W. 14516 (cf.2); D'Arcy, etal. 12658 (3); Daro,M. 461 (9); ex 1. Urera (L.) baccifera Gaudich. Wedd. G. S. Darwin, etal. 2148 (1); Davidse, etal. 28277 (8),37383 2. U. caracasana(Jacq.)Griseb. & A. 3. U.fenestrata K. Monro Al. Rodr. (10), 37493 (1); Davidson,C. & Donahue,J. 8353 (9);
284
C. M. Davidson, 7202 (9); Davidson, E. 349 (3), 484 (9), 487 R. (10); Deam,C. C. 85 (1); Delgado, 24 (5), 76 (9); Delprete, P. 5150 (9); Dbbeler, 5162 (3); Douglas,W.& Krukoff, B. P. A. 3522 (2), 4001 (2); Dryer, J. 179 (3), 233 (3), 234 (3), V. 7646 (10); Duke, A. 9020 (10), 9266 (7), 77844 (9), 72008 J. D. (7); Dunlap,V.C. 174 (2), 247 (9); Dunn, etal. 23234 (2); J. Dwyer, D. 2414 (2),2895 (9), 75338 (5), 15365 (2); Dwyer, M. J. D. & Correa, 7923 (8), 14670 (1) (8); Dwyer, D. & J. Correa M D. 7503 (10), 7964 (9); Dwyer, D. etal. 4837 J. A., (9). J. C, Ebinger, E. 812 (9), 967 (2); Echevarra J. A. 207 J. (10), C. 268 (cf. 2); Edwards, B. 107 (1), P-707 (1); R. Espinoza, 771 (5). K. N. R. Fernndez, & Acosta-Zamudio, 2204 (4); Flores, & Flores, 130 (8); Folsom, P. 5932 (1); Folsom, P. et M. J. J. al. 2240 (3),5577 (9); Fonseca A. 79 (2), 773 (9); Fosberg, Z., F. R. 27335 (7); Frankie, W. 79c (2), 738c (9). G. A. E. B. Gamboa, 45 (5), Garcia, R. & Martinez, 47 (1); G. N. Garwood, C. etal 386 (3), 7734 (1), 2728 (9); Gaumer, F. 501 (1), 936 (1); Gentle, H. 2118 (9),2787 (1),2808 (9), P. 2879 (9); Gentry, 2043 (3), 6246 (9), 6648 (1), 7897 (9), A. 8427 (9); Gentry, et al. 43950 (8), 43976 (9), 79328 (9); A. L C. X., Gi//r, L & Hernandez E. 320 (5); Gmez 5535 (3), 8078 (3); Gome* 2207 (9),27040 (9),23729 (8),23772 (8); P. M. C. J. Gonzlez, & Aragn, 2327 (8); Grayum, H. & R. M. Evans9866 (9); Grayum, H. 7237(cf.3); Grayum, H. et M. A. al 5878 (8); Grijalva, & Burgos, 1547 (9); Grijalva, A. F. 1916 (2); Grijalva, etal 2906 (2), 2924 (8); Guadalupe A. J., S. & Lessette F. 33 (2). F., W. W. Haber, exBello,E. 6403 (1); Haber, & E. Bello7403 W. W. (8); Haber, & Zuchowski, 9293 (9), 9294 (2), 9377 (8), 10889(10), 10907 (9); Haber, etal. 11279 (9), 77296 (cf. W. C. S. 2); Hall, J. S. & Bockus, M. 7564 (2); Hamilton, & B. H C. Stockwell, 3537 (9); Hamilton, 558 (8); Hammel, & E. B. Chavarra, 17536 (5); Hammel, 1363 (10), 7578 (10), 2749 (2), 2676 (9), 3076 (10), 4769 (9); Hammel, et al B. 6840 (9); Hampshire, J.& Whitefoord, R. C. 125 (9), 733 (1), 203 (3),274 (3); Hampshire, J.etal. 632 (9); Hancock, W. R. s.n. (678139) (10); Harmon, E. & Dwyer, D. 3384 (9); W. J. W. W. C. Hatch, R. & Wilson, L s.n.(864375)(9); Hatheway, H. 1354 (10); Hawkes, G. etal. 2150 (9); Hawkins, 983 T. J. (9), 7779 (1), 7732 (2); Hayes,S. 84 (1), 86 (1), 750 (9); M. A. J. Heath, & Long, MA44(9), MA53(9); Henrich, E. & W. N. Stevens, D. 345 (2); Hensold, 1008 (9); Herrera, C. G., 354 (9), 360 (1), 7293 (8),2326 (8), Herrera, C. etal. 542 G., H. H. 0. (3),2926 (2); Herrera, & Guillen, 627 (9); Herrera, & H. H. J. Polanco, 795 (8); Herrera, 916 (9); Heyde, T. & Lux, D. 4067 (10); Holm, W.& Iltis, H. 124 (2); Howard, R. H. R. et al 460 (2); Huft, 1933 (9). M. IbezG.,A. 30 (9); II INBio 183 (5). Jimnez A. 1067 (8), 2275 (2); Jimnez A. & M., M., R. L Rodriguez, 323 (3); Jimenez, & Elizondo, H. 746 (9); Q. I. Jimenez, 876 (9); Johnston, M. 1600 (8), 7706 (8); Q. J. Johnston, R. 1279 (10). W. H. Kellerman, A. 6553 (10); Kennedy, 1939 (9); Kernan, C. 430 (9); Khan, R. et al. 915 (1), 959 (10), 7378 (3); Kirkbride J. H. 47 (10), 752 (2); Kirkbride J. H. & Jr., Jr., S. S. J. Duke, A. 770 (3); Knapp, 1016 (8), 7435 (9); Knapp, & S. A. J. Mallet, 9169 (2); Knapp, & Monro, K. 9255 (9); Knees, S. G. 2706 (9). A. Lao, E. A. & Gentry, 450 (9); Lems,K. 640913 (9); P. R. Lemus, s.n. WB-1178'(2), s.n. 'WB-1217' Lent, W. (2); 228 (2),237 (8), 806 (2), 2067 (9),2500 (3),2605 (10),2922 B. J. (8), 3648 (9), 3803 (3); Len, 796 (9), 939 (9); Lewis, B. 265 (1); Lewis, H. etal 867 (8), 947 (1),2087 (9); Liesner, W. R.L& Lockwood, 2443 (1); Liesner, L 1706 (9), 7733 R. R. R. (1), 7929 (9), 3038 (2), 3727 (9); Liesner, L. et al 15437
38 C. (9), 26275 (1); Livingston (9); Llodge, W.s.n. (13 July L. C. S. 1936)(2); Lobo, 661 (3); Long, E. 130 (1); Lundell, L. H. 6475 (2); Luteyn, L & Kennedy, 4149 (10). J. J. Maas, P. J.M. 2736 (1); MacDougal, M. etal 3217(9); T. MacDougall, s.n. 11 Dec. 1952' (10), s.n. '28 Apr.1964' S., (1); Martnez E. M. 8491 (1), 8492 (8), 15439 (8), 16642 0. S., (8), 20777 (2); Martnez E. M. & Tllez, 12839 (9); Martnez E. M. et al. 1755 (2), 20684 (9), 34699 (4), S., 0. 34824 (cf.4); Martnez, 31 (9), s.n. (ISF225) (2); Matuda, E. 115 (9), 3699 (4), 3956 (10), 4772 (9), 5097 (10), 15347 (9), 15409 (2), 15547 (5), 76687 (9), 16825 (2), 17637 (1); W. J. Maxon, R. 7123(2); McDonagh, F. etal 257 (9),550 (9 G. P. 605 McGillivary,11 (1); McPherson, aft), (9),609 (9 aff.); 15962 (9); Mndez A. 8945 (2), 9054 (4); Mndez A. T., G., 4863 (9), 6238 (9), 6738 (9), 7022 (9), 7267 (4), 9066 (9); G. J. J. Miller, S. 2001 (1); Miller, S. & Sandino, C. 1094 (8), 7743 (2); MolinaR.,A. 868 (1),3782 (9), 5520 (9), 6928 (9), 10875(2),12900(2),75557(9),27969 (cf.1); Molina A.& R., Molina A.24719 (2),26673 (2),30730 (2); Molina A.& R., R., E. Montalvo, 21603 (10); MolinaR., A. et al 16046 (10), E. A. A. Monro, K. 671 (1); Monro, K. & Alfaro, 4241 (2),4346 A. (2),4407 (8),4425 (2), (8); Monro, K. etal. 2997 (9),3076 E. (1), 3520 (3), 3679 (1), 3685 (1); Montalvo, A. 3850 (2); M. Mora,G.37 (5), 543 (3), 643 (3); Moraga, 60 (5); Moraks, J. J.F. 799 (10), 727 (8),5866 (3), 7383(8); Mora/es, F. etal. P. 9937 (8); Moreno, P. 167 (1),238 (1),437 (2),473 (2),544 (2), 978 (2), 7437 (9), 7503 (2), 2602 (8),2689 (2), 2775 (9), 2858 (2),2925 (2),3366 (9),4095 (2),4703 (8),6280 (9),6327 (9),8009 (1),8748 (1), 8377 (1), 10584(2), 70707(2), 77047 (2), 77707 (2), 12397 (8), 13525 (2), 15976 (1), 76468 (2), 16482 (2), 76525 (2), 76972 (2), 17047 (2), 77727(2), 17265 (9), 77297 (9), 77873 (2), 78747 (9), 19006 (9), 19078 (9), 79765 (9), 19582(2), 79827 (2),27469 (2),22093 (9),24724 P. J. (1),24263 (2); Moreno, P. & Henrich, 8410 (2),8883 (1); P. J. More/io, P. & Sandino, C. 14901 (cf.2). E. J. Navarro, 211 (8); Nee,M. & Miller, 27693 (8); Ne, D. M. & Vega, 27906 (2); Neill, A. N198 (2), 825 (2), 7907 S. (9), 7967 (1), 7972 (9), 3752 (2), 4758 (9); Neill,D. A. & P. Viricela, C. 3243a (1), 3593 (2), 3589 (1), 7477 (9); E. C. C. Nelson, & Romero, 4735 (2), 4738 (8); Nelson, 5200 V. C. (9); Nelson, etal 2926A(8),3535 (2), 3955 (2); Nilson, R. & Manfredi, 392 (3). P. A. Oersted, 15112 (7), 21740 (10); 0/>/er, A. 179 (2); J. Ortiz, J. 989 (4). M. 868 PecA;, E. 504 (1), PecA; (9); Penate,V. et al 1307 M. D. (2); Pennell, etal. 58 (10); Penneys, S. 19 (3); Penneys, D. S. & Haber,W. 710 (8); Pipoly, J. 3731 (1), 5036 (1); J. H. Pittier, 2939 (9),3804 (2),3899 (2); PoiN^a,L J.306 (3), 7707 (9), 7759 (7); Purpus, A. 7039(9), 7354(9), 7356(9). C. Quesada,A. 1161 (3); Quesada,F. 167 (8), 275 (5). K C. amirez, 794 (9); RamosE., G. & Cowan, 2685 (9); P. RamosE., G. etal. 2859 (4); Raven, H. 20901 (9), 21623 M. C. (8); Renderos, A. 494 (2); Renson, 204 (1), 279 (1); G. Ttoera, 258 (3), 259 (3), 432 (cf.3), 657 (5), 7752 (5), 7485 (8), 7563 (8); Robles, 1119 (1), 7762 (9), 2097 (9); R. W. Robleto, 153 (2), 935 (cf.9 [cf.1]),992 (cf.9 [closeto 1]); V. A. K. oc/ia, 6 (2); Rodrguez, et al. 6254 (8); 7?oe, et al. 731 (4), 898 (4), 7259 (2); %as, S. & Rojas,L M. 84 (9); J. Rosales, M. 840 (2), 7338 (2), 7427 (2), 7496 (2); Rossbach, G. B. 3690 (8), 3874 (4). A. J. Snchez, 10 (2); Sandino, C. 124 (2),220 (9),237 (2), 570 (1), 7273 (8), 7297 (2), 7369 (9), 7749 (1), 2035 (9), 2060 (2), 2543 (1), 2736 (1), 3307 (8), 3384 (2), 3400 (9); Sandino,J. C. et al 3569A (9); Sandoval,E. 1854 (2); E. R. E. Sandoval, & Chinchilla, 504 (2), 7782 (2); Sandoval, H. E. M. & Rivera, 1252 (9); Sandoval, & Sandoval, 1373 (9); W. J. Saunders, 1204 (7); Schipp, A. 400 (9),8777 (1); Schott,
285
B. D. A. 796 (1), 796a (1); Schubert, G. & Rogerson, L 846 R. 7277(9); Terry, E. & Terry, A. 1403 (7); Todzia, et M. R. C. B. M. S. A. (9); Seemann, C. 146 (9), 494 (7), 495 (1); Segura, 4 al. 2013 (9); Tomlin, 10 (2); Tonduz, 7167 (2), 77770 F. P. R. J. (8); Seibert, J.608 (2); Seymour, C. 3101 (9); Shank, (10); Tucker, M. 965 (1); TnO., R. R. 383 (1), 7080 (9), E. J. & MolinaR., A. 4265 (8), 4405 (8); ShilomT., A. 2579 7323 (8), 2689 (9); Tyson, L 882 (9), 7747 (2), 6869 (cf. K. E. (10),2582 (4), 2585 (5), 4173 (4); Sidwell, etai 435 (10), 9); Tyson, L et al. 4478 (8). A. 585 (2); Skutch, F. 966 (9),982 (5),1479(8),2863 (8),3565 K. J. Utley, & Utley, 635 (2), 7773 (8), 2379 (3), 2352 (3), /I. (3),3602 (3),3750 (9),4266 (7),4842 (9); 5m/i, A397(10), 2477 (9), 2892 (2), 5344 (2). 4439 (9), A597 (9), H408 (9), 7/477 P1988 (9), 700 (3), (9), M. H. Valerio, 1014 (10); van der Werff, & Herrera, J. C. 158 (9), 1082 (9), 7256 (9), 7487 (9),2788 (9); Snut, E. & 6187 (9); van Severen, L 113 (2); Vanderveen, D. 605 M. B. F. H. M. 3380 (9); Solano,J.23 (9), 77 (3); So/is, 187 Smith, E. J. (9); Vargas, et al. 392 (9); Vaughan, et al. 605 (9); P. (1); So2a, D. & Moreno, 76 (2); Soza, D. et al 265 (2); Vega,S. & Quezada,B. 167 (2); Ventur, 289 (5) (9); P. P. Standley, C. 8354 (2), 70772 (2), 77200 (9), 77202 (1), Ventura F. 20686 (4), 20944 (cf. 4); Ventura E. & A., L, 20526 (1),27880 (1),22344 (1),22394 (1),22764 (2),23982 E. 1901 (2); Verhoek, E. 5493 (7); vonHagen,C. & S. 30536 (1), 4736 (1), 52857 (2), 52978 (9), Lpez, (1), 24774 (2), vonHagen,W.2083 (2); von Tuerckheim, 1243 (9); von H. 53265(1),54057(9),60769(10),64714(5),67884(5),68232 H. Wedel, 842 (1). (9), 75549 (5), 75779 (1), 76507 (10), 76980 (5), 78294 (2), A. G. Webster, L et al. 12326 (2), 12688 (9); Weston, S. 79579 (1), 79832 (10), 87009 (10), 82128 (10), 84677 (4), A. 2991 (10),3782 (10),4742 (10),5080 (9); Weston, S. etal. M.37 (5); Semw, 86272(5),90675(9),91395(9); Steggerda, 3104 (8); Whitefoord,1081 (9), 7673 (1), 7887 (9), 3248 C. IT. IT.D. e al. 16650(7), 76979 (2),25612 (1); Seve/w, & & C. & A. R. 3250 (9); Whitefoord, Eddy, 249 (1); Wilbur, L B. A.4744 (2),6428 (9),6607 (1),8696 (2),8700 (1), (9), Krukoff, 13014 (10); Williams, 0. & Cordoba, J.4668 (3); L J. 9284(2),9623(2),72473(1), 12993(1),22708 (9); Severoon, et al. L N. S. 109 (1); Steyermark, 29936 (9), 33332 (4), 33421 Williams, O. et al. 21816 (10), 25249 (10), 26295 (4), J.A. R. 28917 28972 (3), 29009 (2); Williams, S. 692 (1), (9), 33545 (9), 34292 (4), 36257 (5), 36677 (10), 36619 (4), 776 (10), M. (10), 728 (9), 794 (9), 875 (10); Wilson, R. 40944 37409 (4),37516 (5),38770 (1),46672 (4),47937 (9),48785 R. & P. C. 48959 (5), 49249 (9), 57708 (5); Stocker, L 21 (1); (10); Wilson, 97 (9), 579 (1); Woodson, E., Jr. Schery, (9), R. et R. W.257 (3), 593 (3),863 (9); JfWson, E., Jr. al. 1005 H. Stork, E. 1753 (9), 2673 (2); S^ma, /CJ.etal. 1617 (8), C. (9), 7923 (8); Wright, s.n. (1). 7697 (9), 4982 (cf.8). T. T. Yuncker, G. 4514 (9), 5044 (2); Yuncker, G. et al. C. J. Tate,R. 395 (315) (1); Taylor, & Taylor, 11575 (9), R. K. 77672 (9); Taylor, 118 (8); Taylor, J.4536 (8); Tllez, 8025 (9), 8454 (8), 8487 (9). 0. 0. etal. 5181 (9), 7477 (4), 7827(9); Tllez, & Pankhurst, Zamora,N. 625 (3); Zuniga,R. etal. 180 (9).

Three New Species and A Nomenclatural Synopsis of Urera (Urticaceae) From Mesoamerica DCJMB4038

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Three New Species and a Nomenclatural Synopsis of "Urera" (Urticaceae) from Mesoamerica Author(s): Alexandre K.

THREENEWSPECIESANDA NOMENCLATURAL SYNOPSIS OF URERA FROM (URTICACEAE) MESOAMERICA1

AlexandreK. Monro2and AlexanderRodrguez3

Ann. Missouri Bot. Gard. 96: 268-285. Published on 7 July 2009.

Volume 96, Number2 2009

Monro& Rodrguez New Species and Synopsis of Urera

manni Scepocarpus Wedd.

Annals of the Missouri Botanical Garden

Volume 96, Number2 2009

Monro& Rodrguez New Species and Synopsis of Urera

Annals of the Missouri Botanical Garden

Volume 96, Number2 2009

Monro& Rodrguez New Species and Synopsis of Urera

Annals of the Missouri Botanical Garden

Volume 96, Number2 2009

Monro& Rodrguez New Species and Synopsis of Urera

Annals of the Missouri Botanical Garden

3 June1990, R. Delgado 24 (holotype, INB!;

Volume 96, Number2 2009

Monro& Rodrguez New Species and Synopsis of Urera

P. P. C. Standley 64714, F), nigita (Guatemala: C. 75549,F). Standley

Annals of the Missouri Botanical Garden

Volume 96, Number2 2009

Monro& Rodrguez New Species and Synopsis of Urera

Annals of the Missouri Botanical Garden

Santa Clara, Cantnde San Carlos,A. S. Agropecuario,

Volume 96, Number2 2009

Monro& Rodrguez New Species and Synopsis of Urera

Annals of the Missouri Botanical Garden

Volume 96, Number2 2009

Monro& Rodrguez New Species and Synopsis of Urera

Annals of the Missouri Botanical Garden

Volume 96, Number2 2009

Monro& Rodrguez New Species and Synopsis of Urera

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