Three New Species and a Nomenclatural Synopsis of Urera (Urticaceae) from Mesoamerica1

Alexandre K. Monro; Alexander Rodríguez

Uraria picta (Jacq.) DC. (Family Leguminosae, Papilionoidae) is an important plant species in Ayurvedic medicine and one of the most important constituents of the 10-herb formulation called 'Dashmula'. The ayurvedic name of the species is Prishni parni while the trade name is Dabra. The species is endangered, hence requires special attention and an overview was conducted involving the various aspects of U. picta to provide necessary information and to induce interest among researchers for its conservation and utilization in traditional as well as modern systems of medicine.

THREE NEW SPECIES AND A NOMENCLATURAL SYNOPSIS OF URERA (URTICACEAE) FROM MESOAMERICA1 Alexandre K. Monro2 and Alexander Rodrı́guez3 ABSTRACT Urera Gaudich. is unique among Mesoamerican Urticaceae in having bright, fleshy fruits. Within Mesoamerica, there is significant confusion over the application of many names, especially U. corallina (Liebm.) Wedd., U. elata (Sw.) Griseb., and U. eggersii Hieron. Three new species, U. fenestrata A. 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 staminate flowers, staminate inflorescences, stem, leaf morphology, and habit. The affinities of the new species are discussed. The first record of ant associations for the genus is documented in relation to U. fenestrata. In addition, a key is presented to the 10 species of Urera recognized for Mesoamerica; nomenclatural review is given in which U. mitis (Vell.) Miq. is lectotypified; U. baccifera (L.) Gaudich. ex Wedd., U. caracasana (Jacq.) Griseb., U. mitis, and Urtica nitida Vell. are epitypified; and Urera denticulata Miq., U. eggersii, U. subpeltata Miq., and U. subpeltata var. morifolia Miq. are neotypified; and a list is provided of more than 900 exsiccatae from 13 herbaria. RESUMEN Urera Gaudich. es un género único entre las Urticáceas de Mesoamérica por presentar frutos lustrosos y suculentos. En Mesoamérica existe una confusión significativa en la aplicación de muchos de los nombres, especialmente en las especies U. corallina (Liebm.) Wedd., U. elata (Sw.) Griseb. y U. eggersii Hieron. Tres nuevas especies, U. fenestrata A. 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 ilustradas sobre la base de sus flores e inflorescencias estaminadas, tallo, morfologı́a de hojas y hábito. Las afinidades de las nuevas especies son discutidas. Se registra por primer vez la asociación de una hormiga con el género, especı́ficamente en U. fenestrata. Adicionalmente se presenta una clave para las 10 especies reconocidas por Urera en Mesoamérica; se entrega una revisión en la cual U. mitis (Vell.) Miq. es lectotipificada; U. baccifera (L.) Gaudich. ex Wedd., U. caracasana (Jacq.) Griseb., U. mitis y Urtica nitida Vell. son epitipificadas; Urera denticulata Miq., U. eggersii, U. subpeltata Miq. y U. subpeltata var. morifolia Miq. son neotipificadas; y se suministra una lista de más de 900 especimenes existentes en 13 herbarios. Key words: Flora Mesoamericana, IUCN Red list, Mesoamerica, Urera, Urticaceae. The genus Urera Gaudich. comprises shrubs, trees, and vines that occur most frequently in riparian and disturbed vegetation in Mesoamerica. Within the Urticaceae, Urera is characterized by fleshy fruits (formed by the inflation of the tepals), penicillate or capitate stigmas, glabrous pistillodes, and hairs with bulbous bases that are stinging in some species. Urera 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, according to IUCN Red List criteria (IUCN, 2001). Within Mesoamerican Urticaceae, Urera is unique in having bright fleshy fruits. It is also characterized by stems that frequently release a watery latex when cut (a trait shared with Myriocarpa Benth.) and, in some species, stinging, bulbous hairs. It is for these stinging hairs that the genus is most widely known, hence the widespread local name of ‘‘chichicaste,’’ which is derived from the Nahuat word ‘‘tsijtsı́kast’’ meaning ‘‘to vibrate’’ (Bonilla A., pers. comm.). It is 1 We are grateful to Norman Robson (BM) for help with the Latin diagnoses; Charlie Jarvis (BM) and Sandra Knapp (BM) for comments on the manuscript; Roy Gereau (MO) and Melanie Wilmot-Dear (K) for reviewing the manuscript; Rosemary Wise (OXF) for the illustrations; Victor Steinmann for sending images of type material at IEB; and the curators at BM, C, F, GH, INB, K, LL, MEXU, MO, NY, P, PMA, TEX, and US for the loan of, and access to, collections. Some of the paratype material was collected with support from Darwin Initiative grant 15/027. 2 Department of Botany, The Natural History Museum, London, SW7 5BD, United Kingdom. a.monro@nhm.ac.uk. 3 Instituto Nacional de Biodiversidad, Apartado Postal 22-3100, Santo Domingo de Heredia, Costa Rica. arodrig@inbio.ac.cr. doi: 10.3417/2006121 ANN. MISSOURI BOT. GARD. 96: 268–285. PUBLISHED ON 00 MONTH 2009. Annals of the Missouri Botanical Garden mobt-96-02-05.3d 13/5/09 20:09:53 268 Volume 96, Number 2 2009 Monro & Rodrı́guez New Species and Synopsis of Urera also from these stinging hairs that Urera derives its limited economic and medical importance: U. baccifera (L.) Gaudich. ex Wedd. is used as ‘‘living fences’’ 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 treat arthritis (Gonzáles, 1994; House et al., 1995), fever (House et al., 1995), hemorrhage (Guánchez, 1999), erysipelas (Guánchez, 1999), and syphilis (House et al., 1995), and species that are of moderate importance as food for Lepidoptera (Janzen & Hallwachs, 2005). The genus Urera was first described by Gaudichaud-Beaupré 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 monospecific genus Scepocarpus Wedd. by Friis (1989), and chloroplast DNA sequence data (trnL-F) suggests that Urera could be sister to Poikilospermum Zipp. ex Miq. within the Urticaceae (Monro, 2006). Urera has attracted no monographic attention since Weddell (1856, 1869), and, to date, no subgeneric classification has been published (although Weddell did divide the species into unnamed groups according to inflorescence structure and distribution). Systematic work in Urera has largely resulted from localized floristic treatments in and adjoining Mesoamerica (Standley & Steyermark, 1952; Burger, 1977; Pool, 2001; Steinmann, 2005). There are currently 138 published species epithets for the genus (The International Plant Names Index, 2008), of which 133 appear legitimate. Of these epithets, 16 have since been transferred to Laportea Gaudich., Gyrotaenia Griseb., Dendrocnide Miq., Boehmeria Jacq., and Girardinia Gaudich. by subsequent authors. Of the remaining 117 names, Friis (1989) estimates that there are ca. 35 good species and Pool (2001) estimates 35 to 75. Within Mesoamerica, there are 25 specific epithets for 10 recognized species of Urera, and there is significant confusion over the application of many names, especially U. corallina (Liebm.) Wedd., U. elata (Sw.) Griseb., and U. eggersii Hieron. This, combined with a lack of regional keys, the similarity of form and habit, and the extent of overlapping variation in characters between species, has made the determination of collections difficult. The result is that a significant proportion of herbarium material from Mesoamerica is misidentified. MATERIALS AND METHODS 269 This work was undertaken as part of the revision of Mesoamerican Urticaceae for the Flora Mesoamericana project. The definition of Mesoamerica is as given in the Flora (Davidse et al., 1994): a region bounded to its north by the Mexican states of Yucatán, 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) from BM, 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 revision was based on the examination of the original published descriptions for all 24 accepted names, as well as type material. The macro-morphological characters used most frequently by previous authors for the delimitation of species are leaf shape, leaf margin morphology, inflorescence morphology and structure, trichome distribution and morphology, fruit size and color, cystolith shape, and stigma shape. In this study, emphasis was also placed on stem morphology and habit, together with personal observations in the field that some species release a thin, watery latex when cut. Although no reference to 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 collections may have released a watery latex, and it is for this reason that this was noted in the observations made and in the descriptions below. All material was examined using a stereomicroscope at 364 to 3400 magnification, and up to 138 observations were made for each specimen sampled. These observations were then used as a guide to delimit taxa within a morphological species concept and in the preparation of a key to the identification of the Mesoamerican species. Once species were delimited, they were then matched to type material. TAXONOMIC TREATMENT Urera Gaudich., Voy. Uranie 496. 1826 [1830]. TYPE (designated by Britton & Wilson, 1924: 243): Urera baccifera (L.) Gaudich. ex Wedd., Ann. Sci. Nat., Bot., sér. 3, 18: 199. 1852. Scepocarpus Wedd., Prodr. (DC.) 16(1): 98. 1869. TYPE: Scepocarpus manni Wedd. Annals of the Missouri Botanical Garden mobt-96-02-05.3d 13/5/09 20:10:00 269 270 Annals of the Missouri Botanical Garden KEY TO THE MESOAMERICAN SPECIES OF URERA Distributions for each species are given to country level within Mesoamerica, with the exception of Mexico and species that are only known from a single country, in which case they are given to state level. Global distributions in the key are given to regional level (e.g., South America, West Indies), following Flora Mesoamericana protocols (Davidse et al., 1994). Some of the characters in the following keys, e.g., cystoliths, are only visible in dried material. Several types of hairs (i.e., bulbous, straight, and curved) are found on the surface of the leaves and young stems of Mesoamerican Urera. Here I define bulbous hairs as those with a swollen and inflated base giving the base a bulbous or bottle-shaped appearance. 1a. Leaves lobed; stem releasing white latex when cut . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. U. laciniata 1b. Leaves not lobed; stem releasing latex or not; when released, latex gray (never white). 2a. Stem and leaves with spines; leaves coarsely dentate, the teeth spaced every ca. 0.5–1 cm; achene 2–3.2 mm; tepals covering the basal 1/4 or less of achene prior to the inflation of tepals . . . . . . . . . . . . . . . . . . . . . . . 1. U. baccifera 2b. Stem and leaves lacking spines; leaves dentate, crenate, or entire, when toothed the teeth spaced , 0.5 cm apart; achene 0.75–2 mm; tepals covering 3/4 or more of the achene prior to the inflation of the tepals. 3a. Adaxial surface of leaves sparsely pubescent or glabrous. 4a. Leaf margin shallowly crenate, crenate-serrate, sinuate, or entire; hairs never urticating. 5a. Shrub or small tree; leaf-bearing section of stem never hollow. 6a. Ratio of leaf length to width less than 4:1; abaxial leaf surface without domatia in the axils of the secondary veins; mature fleshy fruits pink or orange to orange-red . . . . . 4. U. glabriuscula 6b. Ratio of leaf length to width greater than 4:1; abaxial leaf surface with domatia in the axils of the secondary veins; mature fleshy fruits red . . . . . . . . . . . . . . . . . . . . . . 5. U. guanacastensis 5b. Shrub, lax shrub, or vine; leaf-bearing section of stem hollow, ca. 5–10 mm diam. 7a. Young shoots pubescent to densely pubescent, the hairs 0.125–1 mm; staminate flowers ca. 1.25 3 1.25 mm immediately prior to anthesis . . . . . . . . . . . . . . . . . . . . . . 8. U. lianoides 7b. Young shoots sparsely pubescent or glabrous, the hairs when present 0.125–0.25 mm; staminate flowers ca. 2 3 2.5 mm immediately prior to anthesis . . . . . . . . . 3. U. fenestrata 4b. Leaf margin prominently serrate, crenate, or dentate; hairs urticating or not urticating. 8a. Young stem glabrous or sparsely pubescent, the hairs # 0.25 mm; stem coarsely sulcate, frequently fenestrate; stems, leaves, and petioles lacking bulbous hairs; staminate inflorescence to 110 mm; mature fleshy fruits red-pink . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. U. fenestrata 8b. Young stem sparsely to densely pubescent, the hairs . 0.5 mm; stem coarsely sulcate but never fenestrate; stems, leaves, and petioles with or without bulbous hairs; staminate inflorescence to 80 mm; mature fleshy fruits orange or orange-pink. 9a. Tertiary venation of abaxial leaf surface cream to pale green, noticeably paler than lamina; stipules forked or not forked; stem sparsely pubescent . . . . . . . . . . . . . . . . 6. U. killipiana 9b. Tertiary venation of abaxial leaf surface darker or rarely paler than the lamina, where paler than the lamina pale brown to orange-brown in color; stipules not forked; stem densely pubescent. 10a. Leaves ovate or cordiform, never obovate or lanceolate; hairs urticating . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. U. caracasana 10b. Leaves obovate, lanceolate, elliptic, or ovate, never cordiform; hairs urticating or not . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. U. simplex 3b. Adaxial surface of leaves pubescent to densely pubescent. 11a. Leaves bullate; staminate peduncle unbranched at base for 40–80 mm; pistillate peduncle unbranched at base for 27–98 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. U. verrucosa 11b. Leaves not bullate; staminate and pistillate peduncles unbranched at base for , 25 mm. 12a. Leaves ovate or cordiform, never obovate or lanceolate; hairs urticating . . . . 2. U. caracasana 12b. Leaves obovate, lanceolate, elliptic, or ovate, never cordiform; hairs urticating or not . . . 9. U. simplex 1. Urera baccifera (L.) Gaudich. ex Wedd., Ann. Sci. Nat., Bot., sér. 3, 18: 199. 1852. Basionym: Urtica baccifera L., Sp. Pl., ed. 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, BM!). Urera armigera (C. Presl) Miq., Fl. Bras. (Martius) 4: 192. 1853. Basionym: Urtica armigera C. Presl, Bot. Bemerk. (C. Presl): 110. 1844 [1845]. TYPE: Brazil. ‘‘near Rio de Janeiro,’’ J. Lhotsky s.n. (lectotype, designated by De Rooij [1975: 302], PR not seen). Urera denticulata Miq., Fl. Bras. (Martius) 4: 192. 1853. TYPE: Brazil. Minas Gerais: Viçosa, rd. E from Chacha valley, Fazenda da Creciuma, 10 May 1930, Y. Mexia 4679 (neotype, designated here, BM!; isotype, MO not seen). Urera baccifera var. horrida (Kunth) Wedd., Arch. Mus. Hist. Nat. 9: 151. 1856. Basionym: Urtica horrida Kunth, Nov. Gen. Sp. [HBK] (quarto ed.) 2: 41. 1817. Urera horrida (Kunth) Miq., Fl. Bras. (Martius) 4: 192. 1853. TYPE: Colombia. ‘‘Santander, Magdalena prope Angostura de Carare,’’ Humboldt & Bonpland 1639 (lectotype, designated by De Rooij [1975: 302], P!). Urtica nitida Vell., Fl. Flumin. Icon. 10: t. 20. 1827 [1831]. TYPE: Fl. Flumin. Icon. 10: tab. 20. 1827. EPITYPE: Brazil. Mato Grosso do Sul: Mpio. Paraguai, Serra des Annals of the Missouri Botanical Garden mobt-96-02-05.3d 13/5/09 20:10:01 270 Volume 96, Number 2 2009 Monro & Rodrı́guez New Species and Synopsis of Urera Araras, Fazenda Currupira, 15u109S, 56u59W, 24 Jan. 1995, Dubs 1770 (epitype, designated here, K!; duplicates, E not seen, ESA not seen). Urtica grandidentata Liebm., Kongel. Danske Vidensk. Selsk. Skr. (Trondheim) ser. 5, 2: 296. 1851. TYPE: Costa Rica. Cartago: ‘‘Irasú,’’ Jan. 1848, A. Örsted 14283 (lectotype, designated by De Rooij [1975: 302], C!). US), nigua, niguilla (El Salvador: P. C. Standley 22394, GH), ortiga (Costa Rica: R. Anderson & S. Mori 147, F; Panama: P. C. Standley 30536, US), ortiga de los caballos (Mexico: A. Schott 796, BM, F), rascate bien (Honduras: A. Molina R. 868, F, GH). An epitype is selected for Urera baccifera because the type illustration, although accurate, is not sufficient to support the unambiguous fixing of the name to this species. Fawcett 7177 was chosen because it includes flowering and vegetative material that conforms to Linnaeus’ (1763) description ‘‘Urtica foliis alternis, cordatis’’ and is from the West Indies, as was the Plumier material that formed the basis of the type illustration. An epitype is selected for Urtica nitida because the type illustration, although accurate, is not sufficient to support the unambiguous fixing of the name to this species. Dubs 1770 was chosen because it includes flowering and vegetative material that conforms to 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, although de 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.) or BR (P. Stoffelen, pers. comm.). The collections database from the Field Museum’s Department of Botany (Field Museum of Natural History, 2006), which includes the negatives of photographs of European type material taken by Macbride, does, however, include a negative (no. 8845) cited as being from a Martius collection that represents type material of U. denticulata at M. Such a negative cannot be considered isotype material, but it does indicate the existence of a type specimen existing prior to World War II. It may be that de Rooij based his lectotypification on this photograph or 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 includes good leaf, stem, and fruiting material and is from the same country and state as the material cited in Miquel’s description. Local names. Bringa mosa (Panama: C. Whitefoord & A. Eddy 249, BM), chichicaste (Guatemala: J. A. Steyermark 38770, F; Honduras: P. C. Standley 20526, F; El Salvador: P. C. Standley 22344, F; Nicaragua: P. C. Standley 11202, F), chichicaste cuyanigua (El Salvador: P. C. Standley 21880, GH, US), chichicaste nigra (El Salvador: S. Calderón 1539, NY), cow itch (Belize: P. H. Gentle 2781, F, GH, NY, 271 Habitat and distribution. Evergreen or seasonal forest, riparian vegetation, from sea level to 1400 m, Mexico to Panama, Colombia, Peru, Bolivia, Brazil, Paraguay. Comments. This species most closely resembles Urera laciniata Goudot. ex Wedd. These species can be distinguished from each other based on the presence of latex and spines and leaf margin morphology as 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 margin deeply lobed or laciniate. Selected specimens examined. BELIZE. Cayo: Chiquibul, Las Cuevas, 16u439N, 88u599W, A. K. Monro 671 (BM, BRH, MO). BOLIVIA. Cochabamba: A. M. Bang, N. L. Britton & H. H. Rusby 1209 (A, BM, GH, MO). COSTA RICA. Alajuela: Rı́o Sarapiquı́ at bridge on rd. to Colonia Virgen del Socorro, 9 mi. SE of San José–Puerto Viejo hwy., 10u169N, 84u119W, T. B. Croat 68307 (BM, MO). EL SALVADOR. La Libertad: Cordillera de Balsamó, San Julián rd., towards the Pacific, 13u419000N, 89u389320W, A. K. Monro et al. 3676 (BM, ITIC, LAGU, MO). GUATEMALA. Izabal: Montañas del Mico, 4–5 km W of Santo Tomás de Castilla, W. D. Stevens et al. 25612 (BM, MO). HONDURAS. Atlántida: 15u429N, 86u519W, R. L. Liesner 26275 (MO). NICARAGUA. Rı́o San Juan: Near Caño Chontaleño 20 km NE of El Castillo, D. Neill & P. C. Vincelli 3589 (BM, MO). PANAMA. Darién: Rı́o Cocalito, C. Whitefoord & A. Eddy 249 (BM, MO). PARAGUAY. Canindeyú: Karapa, Salto a 5 km del puesto, G. Marı́n, B. Jiménez & M. Chocarro P. 763 (BM, FMB). PERU. Pasco: Oxapampa, Distr. Huancabamba, Sector Grapanazu, limite Parque Nac. Yanachaga-Chemillen, R. Rojas, K. Meza, J. Lingan, E. Camavilca & M. Villaran 1832 (BM, MO). 2. Urera caracasana (Jacq.) Griseb., Fl. 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!). Urera alceifolia (Poir.) Gaudich., Voy. Uranie, Bot. 497. 1826 [1830]. Basionym: Urtica alceifolia Poir., Encycl. (Lamarck) Suppl. 4: 227. 1816. TYPE: French Guyana. Cayenne: s.d., Martin s.n. (lectotype, designated by de Rooij [1975: 304], P!). Urtica tiliifolia Kunth, Nov. Gen. Sp. [HBK] (quarto ed.) 2: 34. 1817, as ‘‘tiliaefolia.’’ TYPE: Colombia. Bolı́var: Rı́o Magdalena, Humboldt & Bonpland 1633 (lectotype, designated by de Rooij [1975: 306], P!). Urera jacquinii var. ulmifolia (Kunth) Wedd., Arch. Mus. Hist. Nat. 9: 145. 1856. Basionym: Urtica ulmifolia Annals of the Missouri Botanical Garden mobt-96-02-05.3d 13/5/09 20:10:01 271 272 Annals of the Missouri Botanical Garden Kunth, Nov. Gen. Sp. [HBK] (quarto ed.) 2: 141. 1817. TYPE: Colombia. Bolı́var, Humboldt & Bonpland 1427 (lectotype, designated by de Rooij [1975: 306], P!). Urera mitis (Vell.) Miq., Fl. Bras. (Martius) 4(1): 191. 1853. Basionym: Urtica mitis Vell., Fl. Flumin. 10 tab. 19. 1827 [1831]. Urera caracasana var. mitis (Vell.) Wedd., Prodr. (DC.) 16: 90. 1869. TYPE: Fl. Flumin. Icon. 10: tab. 19 (lectotype, designated here, tab. 19!). EPITYPE: Brazil. Amazonas: Marapata, Municipality of Carauary, 25 May 1933, B. Krukoff 4568 (epitype, 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, Örsted 14282 (lectotype, designated by de Rooij [1975: 304], further designation here, C sheet ‘‘26/2003/2’’!). Urera capitata Wedd., Ann. Sci. Nat., Bot., ser. 3, 18: 201. 1852. TYPE: Bolivia. Yungas: Dec. 1846, Weddell 4317 (holotype, P!). Urera subpeltata Miq., Fl. Bras. (Martius) 4(1): 189, pl. 66. 1853. Urera jacquinii var. subpeltata (Miq.) Wedd., Arch. Mus. Hist. Nat. 9: 145. 1856. Urera caracasana var. subpeltata (Miq.) Wedd., Prodr. (DC.) 16: 90. 1869. TYPE: Brazil. Bahia: 1839. Blanchet 927 (neotype, designated here, BM!; isotype, G not seen). Urera subpeltata var. morifolia Miq., Fl. Bras. (Martius) 4(1): 190. 1853. TYPE: Brazil. Pará: Santarem, July 1850, R. Spruce s.n. (neotype, designated here, BM!). Urera jacquinii var. miquelii Wedd., Arch. Mus. Hist. Nat. 9: 145. 1856. Urera caracasana var. miquelii Wedd., Prodr. (DC.) 16: 90. 1869. TYPE. Peru. Gay s.n. (holotype, P not seen). Urera acuminata Miq., Fl. Bras. (Martius) 4(1): 190. 1853, nom. illeg. non Urera acuminata (Poir.) Gaudich. and is from the same country and region as the holotype. A neotype is selected for Urera subpeltata var. morifolia because although de Rooij (1975: 306) cites Spruce 633 (M) as lectotype, no Spruce collection fitting this description could be traced either at M (M. Esser, pers. comm.) or BR (P. Stoffelen, pers. comm.). Miquel (1853: 190), in his original description, cites two collections from the Brazilian Amazon, ‘‘Martius in silvis amazonicis, ad Barra do Rı́o Negro’’ and ‘‘Spruce ad Santarem,’’ neither of which could be traced either at M (M. Esser, pers. comm.) or BR (P. Stoffelen, pers. comm.). A Spruce collection was, however, located at BM with the annotation, ‘‘In vicinibus Santarem, Prov. Pará.’’ This is selected as neotype on the basis that the collection is from the same collector, country, and state and includes good leaf, pistillate, and immature fruit material. A neotype is selected for Urera subpeltata because although de Rooij (1975: 306) cites Martius s.n. at M as lectotype for U. denticulata, no such collections could be traced either at M (M. Esser, pers. comm.) or BR (P. Stoffelen, pers. comm.). The specimen designated as neotype was selected because it includes good leaf, stem, and pistillate and staminate material and is from the same country and state as the material cited in Miquel’s 1853 description. De Rooij (1973: 306) cites Makin s.n. (not traced) as type (holotype?) for Urera jacquinii var. miquelii. Weddell (1856), however, cites only Claude Gay’s collection from Peru in his description and this is maintained as holotype. Jacquin’s original description of Urera caracasana is based solely on staminate material. Original material for the name has not been located at BM or LINN, and any material that may have been at W has probably been destroyed (material could not be traced at W). Based on the type illustration, it is not possible to distinguish between U. caracasana and U. corallina (Liebm.) Wedd. on sterile characters alone. Material examined that had been determined as U. corallina (including the holotype) by Weddel and as U. caracasana did not uncover any significant morphological differences. An epitype is selected because the type illustration, although accurate, is not sufficient to support the unambiguous application of the name to this species. A lectotype is designated for Urtica mitis Vell. because Vellozo (1827) does not refer to the plate as type material as his description predates the type concept. Tabulae 19 can be considered original type material. An epitype is selected for Urera mitis because the type illustration, although accurate, is not sufficient to support the unambiguous fixing of the name to this species. The epitype was selected because it includes both leaves and inflorescences Local names. Chichicaste (El Salvador: E. Sandoval 1854, BM; Nicaragua: P. C. Standley 10712, F), chichicaste blanco picante (El Salvador: E. Sandoval & R. Chinchilla 504, MO), chichicaste cujanigua de altura (El Salvador: E. Sandoval & R. Chinchilla 1182, MO), chichicaste rojo picapica (El Salvador: O. Martinez s.n. (ISF225), MO), pan caliente (Honduras: C. Nelson et al. 3955, BM), migirillo (Costa Rica: A. Sanchez 10, F), miguito (Costa Rica: J. A. Echeverria C. 268, F), ortiga (Costa Rica: J. A. Echeverria C. 268, F), pan caliente (El Salvador: M. L. van Severen 113, F), zulsimtezla (Mexico: A. Méndez G. 8945, BM). Habitat and distribution. Cloud forest, shade coffee forest, pine forest, Quercus L.–Liquidambar L.–Pinus L. forest, from sea level to 2300 m. Mexico (Chiapas), Belize, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, and Argentina. Comments. Herbarium material of this species is commonly determined as Urera corallina. This species Annals of the Missouri Botanical Garden mobt-96-02-05.3d 13/5/09 20:10:01 272 Volume 96, Number 2 2009 Monro & Rodrı́guez New Species and Synopsis of Urera most closely resembles U. verrucosa (Liebm.) V. W. Steinm. The two species can be distinguished from each other based on leaf texture and inflorescence peduncle size as follows: (1) for U. caracasana, leaves chartaceous, staminate peduncle branched to base or unbranched at base for 2–13 mm, pistillate peduncle branched to base or unbranched at base for 2–20 mm; (2) for U. verrucosa, leaves bullate, staminate peduncle unbranched at base for 40–80 mm and densely pubescent, pistillate peduncle unbranched at base for 27–98 mm. pubescent, the hairs 0.25–0.5 mm; leaf blade 103– 320 3 64–220 mm, ovate, broad-ovate, or broadelliptic, chartaceous; adaxial surface sparsely pubescent or glabrous, the hairs 0.375–0.675 mm, weakly appressed, weakly curved or crooked; the cystoliths punctiform to 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, primary to quinternary or hexternary veins visible to the naked eye, the lateral primary veins visible for 2/3 of the leaf length; domatia not present in the axils of the secondary veins; base cordate or obtuse; margins entire or weakly crenate to serrate; apex subcuspidate to cuspidate. Peduncular bracts 2–3 mm; bracteoles 0.375–0.5 mm; staminate inflorescences 1 to 16 per stem, peduncle branched to base or unbranched from base for 2–21 mm, pubescent, the hairs to 0.125 mm, the whole inflorescence 15–110 mm, bearing 160 to 350 flowers in a symmetrical cyme with 3 or 4 orders of dichotomous branching; the flowers borne in clusters of 10 to 35, pedicellate to subsessile, the pedicels when present to 0.5 3 0.125 mm. Pistillate inflorescences 1 to 4 per stem, the peduncle branched to base or unbranched at base for 2–26 mm, pubescent, the hairs ca. 0.125 mm, erect, straight, the whole inflorescence 10–80 mm (47–80 mm in fruit), broader than long (or as long as broad), bearing 224 to 750 flowers in a symmetrical cyme with 5 orders of dichotomous branching, the flowers borne in clusters of 3 or 4, pedicellate to subsessile, the pedicels when present to 0.25 3 0.125 mm, glabrous. Staminate flowers 2–2.5 3 1.25–1.5 mm immediately prior to anthesis; tepals 4, ca. 3 mm; stamens 2.5– 3 mm; pistillode ca. 0.5 mm in diam., glabrous. Pistillate flowers ca. 1 3 0.5 mm, the lateral tepals 0.5–0.675 mm, asymmetrically ovate; the dorsal tepal 0.50–0.675 mm, ovate, with a subapical dorsal thickening, the ventral tepal ca. 0.375 mm, ovate; stigma penicillate, erect. Fruit prior to inflation of tepals ca. 1.25 3 1 mm; basal 3/4 of achene obscured by tepals, the laterals ca. 1 mm, the dorsal tepal 0.675–0.75 mm, the ventral ca. 0.5 mm; achene 0.75–1 3 0.75 mm, asymmetrically elliptic, keelshaped, surface smooth. Fruit when ripe with tepals inflated and berry-like, ca. 1.5 3 1 mm, red-pink, red-violet, pink, or orange when fresh. Selected specimens examined. ARGENTINA. Jujuy: San Pedro, Sierra Sta. Barbara, S. Venturi 9633 (K). BELIZE. Cayo: Caracol Maya ruins, 14 km W of Las Cuevas, 16u459N, 88u079W, T. Hawkins 1132 (BM, MO). BOLIVIA. Cochabamba: Carrasco, ca. 11 km below Sehuencas, J. R. I. Wood 10275 (BM, K). BRAZIL. Roraima: Ilha de Maracá, SEMA Ecological Reserva, S of Cachoeira de Fumaça, W. Milliken 546 (K). COSTA RICA. Puntarenas: W. Haber & W. Zuchowski 9294 (MO). ECUADOR. s. loc., Eggers 15611 (K). Pichincha: new Alluriquı́n–Quito rd., Km 4, P. J. M. Maas & L. Cobb 4770 (K). EL SALVADOR. La Libertad: Mpio. Antiguo Cuscatlan, ‘‘Laderas de La Laguna,’’ P. Lemus s.n. ‘‘WB-01217’’ (BM, LAGU, MO). GUATEMALA. Jutiapa: D. Dunn et al. 23234 (MO). HONDURAS. Ocotepeque: Alrededores de Belén Gualcho, C. H. Nelson, E. Romero, A. Rubio & M. Pereira 3955 (BM, MO). MEXICO. Chiapas: Rancho Puy Ukum, sobre la carr. a 2 km de Bochil, Mpio. de Bochil, A. Méndez G. 8945 (BM, MEXU). NICARAGUA. Jinotega: ‘‘El Recreo,’’ 4 km al N de Sta. Gertrudis, P. Moreno & J. C. Sandino 7897 (BM, HNMN). PANAMA. Chiriquı́/Bocas del Toro: Along Continental Divide on trail in Zona Palo Seco, 08u479N, 82u139W, S. Knapp & J. Mallet 9169 (BM, MO, PMA, SCZ). PERU. Loreto: Rı́o Itaya, T. B. Croat 18829 (K). VENEZUELA. Falcón: Sierra de San Luis, ca. del Puente de Jobo entre Curimagua y San Luis, J. A. Steyermark 99249 (K). 3. Urera fenestrata A. K. Monro & Al. Rodr., sp. nov. TYPE: Costa Rica. Guanacaste: Monteverde, Cordillera de Tilarán, Pacific slope, above Quebrada Cuecha, 1540–1600 m, 6 May 1976, V. J. Dryer 179 (holotype, F!; isotype, CR!). Figure 1A–E. Species nova Urerae caracasanae (Jacq.) Griseb. similis, sed ab ea floribus staminatis tetrapartitis, ramulis petiolisque espinosis ramulis saepe fenestratis atque foliorum nervis lateralibus per 2/3 longitudinis visibilibus differt. Shrub, lax shrub, vine, or small slender tree, dioecious (?). Main stems arching 2–10 m, stems not releasing white latex when cut; without spines; young shoots glabrous or sparsely pubescent, the hairs 0.125–0.25 mm, erect, straight to weakly curved; internodes of leaf-bearing sections of stem 9–30 3 4– 10 mm, pale brown to red-brown, coarsely sulcate, hollow, and frequently fenestrate when $ 5 mm diam. Stipules 5–18 mm, lanceolate, not forked, pubescent; petioles 52–350 3 1.5–1.75 mm, glabrous or sparsely Local names. 273 Ortı́ga (A. Smith 100, F, NY). Habitat and distribution. Urera fenestrata is found in premontane, montane, and cloud forest, frequently in disturbed shaded areas close to streams or small rivers, at 800–3000 m. It occurs on the Pacific and the Caribbean slopes of the Cordillera de Tilarán in Costa Rica and the Cordillera de Talamanca in Costa Rica and Panama. Annals of the Missouri Botanical Garden mobt-96-02-05.3d 13/5/09 20:10:02 273 274 Annals of the Missouri Botanical Garden Figure 1. A–E. Urera fenestrata A. K. Monro & Al. Rodr. —A. Habit, with staminate 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, with staminate inflorescences. —G. Leaf adaxial surface. —H. Pistillate inflorescences. —I. Immature fruit. (F–I: Delgado 24 [MO]). Annals of the Missouri Botanical Garden mobt-96-02-05.3d 13/5/09 20:10:02 274 Volume 96, Number 2 2009 Monro & Rodrı́guez New Species and Synopsis of Urera IUCN Red List category. Conservation for Urera fenestrata 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 28 times in several localities in Costa Rica and Panama. 9 mi. from bridge over Rı́o Grande de Orosı́, A. Gentry 2043 (MO); a la vera del Rı́o Turrialba, camino entre Pacayas y Santa Cruz de Turrialba, J. Gómez-L. 8078 (CR); Tausito, S of Tres Rı́os, cerros de la Carpintera, R. Khan, M. Tebbs & A. R. Vickery 1318 (BM); ca. S of Tapantı́, along Rı́o Grande de Orosı́, R. W. Lent 806 (CR); Tausito, R. W. Lent 3803 (F, NY); Parque Nacional Tapantı́, Sector Dos Amigos, G. Mora 543 (INB); Reserva Forestal Rı́o Macho, Sector El Embalse, G. Mora 643 (INB); Reserva Forestal Rı́o Macho, sendero atráz da casa a direito, V. Nilsson & R. Manfredi 393 (CR); El Empalme, Interamericana Sur, L. J. Poveda 306 (CR); El Guarco, Parque Nac. Tapantı́–Macizo de la Muerte, Cerro de la Muerte, 600 m N de Chespirito, D. Solano 77 (INB); Turrialba, camino a Moravia, antes del Rı́o Pacuare, L. O. Williams & J. J. Córdoba 4668 (CR); Guanacaste: Palmira, fog zone, A. Smith 100 (F, NY). Heredia: Parque Nac. Braulio Carrillo, Zurqui station, just above Los Guarumos trail, B. Boyle 2450 (BM, CR); Parque Nac. Braulio Carrillo, sector Zurquı́, I. A. Chacón 1881 (CR); La Palma, Rı́o Bajo de Honduras, Braulio Carrillo Natl. Park, N. Garwood, M. Gibby, R. J. Hampshire & C. J. Humphries 386 (BM, CR); Quebrada el Mochote, borde de Cerro Zurquı́, L. D. Gómez, I. A. Chacón, G. Herrera, M. M. Chavarrı́a 21040 (CR); Porrosatı́ de Barva, A. Jiménez & R. Rodrı́guez 323 (CR); Cantón de Heredia, Cordillera Central, headwaters of Rı́o Santo Domingo, N slope of Volcán Barva, M. Grayum 7237 (INB); S slope of Volcán Barva, W. H. Hatheway 1354 (CR); Parque Nac. Braulio Carrillo, Estación Barva, G. Rivera 258 (INB), G. Rivera 259 (INB); Rı́o Porrosatı́, 50 m N de parada de buses de Paso Llano, G. Rivera 432 (INB); S slopes of Volcán Barba betw. Rı́o Cirueles & Sacramento, J. Utley & K. Utley 2319 (NY); Parque Nac. Braulio Carrillo, sendero hacia Quebrada Zrquı́, Zamora 625 (CR). Limón: Cantón de Talamanca, Cordillera Talamanca, camp Rı́o Lori, J. Bittner 1882 (INB), 1894 (INB). Puntarenas: Cantón de Buenos Aires, Estación Tres Colinas, finca Benito Acuña, E. Alfaro 745 (INB); Cantón de Coto Brus, Zona Protectora Las Tablas, Las Tablas, sendero Echandi, E. Alfaro 1260 (INB); Cantón de Coto Brus, Zona Protectora Las Tablas, I Camp. ACLA, camino a Cerro echando, E. Alfaro 1569 (CR); near the Continental Divide ca. 2–5 km E & SE of Monteverde, W. C. Burger & J. L. Gentry 8639 (CR), 8752 (CR). Puntarenas– Alajuela border: 2–5 km E & SE of Monteverde, W. C. Burger & R. L. Liesner 8639 (F); Reserva Santa Elena, Monteverde, D. S. Penneys 19 (CR). San José: Vásquez de Coronado, Parque Nac. Braulio Carrillo, sendero frente a Estación Zurquı́, L. Acosta 480 (CR, INB), L. Acosta 486 (CR, INB); valley of Rı́o Hondura (below La Palma), NE of San Jerónimo, W. C. Burger & R. G. Stolze 4912 (CR, F); Rı́o Claro valley (Rı́o La Hondura drainage), below La Palma, NE of San Jerónimo, W. C. Burger & R. G. Stolze 7646 (F); bosquecillos residuales entre las Nubes y Cascajal de Coronado, J. Gómez-Laurito 5535 (CR); near quebrada Bajo Máquina, 3 km NE of Cascajal, R. W. Lent 2500 (BM); Acosta Palmichal, Zona Protectora Cerros de Escazú, El Cedral, ca. de la casa, S. Lobo 661 (CR); Cantón de Aserrı́, Zona Protectora Cerros de Escazú, Cerros Escazú–La Carpintera, la cima del Cerro Daser, Alto Hierba Buena, J. F. Morales & L. Bohs 3787 (INB); Cerro Cedral, falda NW, J. F. Morales 5866 (CR, INB); Tarrazú, San Carlos, Bajos de La Virgen, confluencia rı́os Negro y Blanco, Albergue Rı́os Paraı́so, A. Quesada 1161 (CR); Turrubares, San Luis, San Rafael, Finca de Melvin Chavarrı́a, A. Rodrı́guez 9313 (INB); Alto de la Palma, ca. de 6 km N of San Jerónimo, J. Utley & K. Utley 635 (CR). PANAMA. Bocas del Toro: Chiriquı́ trail betw. quebrada Higuerón & Gutierrez, J. H. Kirkbride & J. A. Duke 770 (NY); Caribbean slopes of Cerro Fábrega at foot of ‘‘Falso Etymology. From the Latin ‘‘fenestratus,’’ meaning ‘‘windowed,’’ referring to the stems which, when greater than or equal to 5 mm diam., become characteristically windowed. Discussion. This species is characterized by the gnarled, windowed, leaf-bearing stem sections and the ovate entire or subentire to weakly indented leaves. Urera fenestrata has most frequently been determined as U. elata or U. caracasana and may be distinguished from them as follows: (1) for U. caracasana, stems when greater than or equal to 5 mm diam. hollow, but never fenestrate, lacking spines; petiole lacking spines; the leaf margins serrate, serrate-dentate, or crenate-serrate to dentate; staminate flowers 5(4)parted; (2) for U. elata, stems when greater than or equal to 5 mm diam. hollow, but never fenestrate, small spines present; petiole with small spines; the leaf margins serrate, serrate-dentate, or dentate; staminate flowers 4-parted; (3) for U. fenestrata, stems when greater than or equal to 5 mm diam. hollow, fenestrate with prominent ca. 5 mm windows, lacking spines; petioles lacking small spines; the leaf margins entire or weakly crenate to serrate; staminate flowers 4-parted. Both authors have observed ants of the genera Crematogaster Lund. and Pheidole Westwood (Formicidae, Myrmicinae) occupying the hollow stems of this species in the field. Collections of these ant species are available at the Instituto Nacional de Biodiversidad entomology collections. Paratypes. COSTA RICA. Alajuela: La Palma de San Ramón, A. M. Brenes 136 (5544) (CR, F, NY); La Palma de San Ramón, A. M. Brenes 220 (4208) (CR, F); rd. to Finca Los Ensayos off hwy. 15, ca. 7.5 mi. N of Zarcero, T. B. Croat 43488 (MEXU, MO); Cordillera de Talamanca, V. J. Dryer 44 (CR); Cordillera de Tilarán, V. J. Dryer 179 (CR), 233 (CR), 234 (CR); Cantón de San Ramón, Reserva Santa Elena, Cordillera de Tilarán, 100 m NW of Station, D. Penneys 19 (INB); Vara Blanca de Sarapiquı́, N slope of Central Cordillera, betw. Poás & Barba volcanoes, A. F. Skutch 3602 (NY); Cordillera Central near San Juan de Laja, ca. 15 km N of Zarcero, L. O. Williams, A. Molina R., T. P. Williams & D. N. Gibson 28972 (F, NY); Reserva Forestal de Grecia, cuenca superior del Rı́o Rosales, G. Herrera C., G. Umaño & H. Gómez 542 (BM, INB, MO); above Quaker settlement at Monteverde, J. Utley & K. Utley 2352 (MO). Cartago: Cantón de Paraı́so, Reserva Forestal Rı́o Macho, Rı́o Pejivalle, E. Alfaro 1799 (INB); Tapantı́ Hydroelectric Reserve along Rı́o Grande de Orosi, T. Croat 36151 (CR); Orosi, farhweg von Rı́o Macho, in westlicher Richtung zum Stausee, P. Döbbeler 5162 (CR); Tapanti I.C.E. Reservation, Annals of the Missouri Botanical Garden mobt-96-02-05.3d 13/5/09 20:10:04 275 275 276 Annals of the Missouri Botanical Garden Fábrega,’’ A. K. Monro & S. Cafferty 4855 (BM, INB, MO, PMA), 4856 (BM, INB, MO, PMA). Chiriquı́: vic. of Gualaca ca. 10.7 mi. from Planes de Hornito, La Fortuna rd. to dam site, T. Antonio 5111 (BM); betw. Bambito & Cerro Punta, T. B. Croat 10593 (MO); betw. Palo Alto & top of ridge (divide) near Cerro Pate Macho above Rı́o Palo Alto, W. D’Arcy et al. 12658 (BM); Boquete, Bajo Chorro, M. E. Davidson 349 (F); near Fortuna Dam, R. J. Hampshire & C. Whitefoord 214 (BM), 203 (BM); lower reaches of trail to Cerro Pando, A. K. Monro, S. Knapp, J. Mallet, A. Mallet, I. Mallet & V. Mallet 3520 (BM, MO); quebrada Velo, R. E. Woodson &R. W. Schery 257 (MO); vic. of Bajo Mona & quebrada Chiquero, R. E. Woodson & R. W. Schery 593 (GH, MO); Bajo Mona, mouth of quebrada Chiquero, along Rı́o Caldera, R. E. Woodson, P. H. Allen & R. J. Seibert 1005 (MO, NY). 4. Urera glabriuscula V. W. Steinm., Acta Bot. Mex. 71: 22. 2005. TYPE: Mexico. Veracruz: Mpio. San Andrés Tuxtla, Volcán San Martı́n, 1300 m, 2 Apr. 1985, Cedillo T. 3175 (holotype, IEB!; isotype, MEXU not seen). Local names. G. 9054, BM). K’anal zulzimtez (Mexico: A. Méndez Habitat and distribution. Montane forest, deciduous and evergreen forest, Liquidambar–Taxodium Rich. forest from 100–2800 m. El Salvador, Mexico (Chiapas, Oaxaca, Tabasco, Veracruz), Guatemala. Comments. Urera glabriuscula is most similar to U. lianoides in its glabrous or sparsely pubescent leaves that are entire or discretely divided. The two species can be distinguished from each other based on habit, stem, stipule, staminate flower, and achene morphology as follows: (1) for U. glabriuscula, shrub or small tree, leaf-bearing section of stem never hollow, with internodes less than 20 mm; stipule apex minutely forked; pistillode 0.75 mm diam.; achene surface verrucate; (2) for U. lianoides, vine or scrambling shrub, leaf-bearing section of stem hollow at ca. 5–10 mm diam. with internodes greater than 20 mm; stipule apex not forked; pistillode 0.375 mm diam.; achene surface smooth. Selected specimens examined. GUATEMALA. Quezaltenango: J. A. Steyermark 34292 (F). MEXICO. Chiapas: Mpio. Maragritas, Tenejapa, Colonia Maravilla, A. Méndez G. 9054 (BM, MEXU). Oaxaca: Héctor M. Hernández y A. Chacón 487 (MO). Tabasco: Mpio. Teapa, en Cerro Madrigal a 500 m al E de Puyacatengo, Universidad Autónoma Chapingo, E. M. Martı́nez S., J. Calonico Soto, A. M. Hanan-Alipi, M. A. Hernández, A. Martinez & N. Peregrino 34699 (BM, MEXU). Veracruz: A. Gentry, E. Lott & UNAM tropical botany class 32227 (BM, MO). 5. Urera guanacastensis A. K. Monro & Al. Rodr., sp. nov. TYPE: Costa Rica. Guanacaste: Parque Nacional Guanacaste, Cantón de Liberia, Estación Cacao, 10u559450N, 85u289150W, 1100 m, 3 June 1990, R. Delgado 24 (holotype, INB!; isotype, MO!). Figure 1F–I. Species nova Urerae simplici Wedd. similis, sed ab ea ramulis glabris, foliis angustioribus supra parce pubescentibus vel galbris subtus domatiis praeditis, pedicellis florum staminatorum breviorbus atque fructu rubro differt. Shrub to small tree, dioecious (?), main stems arching, 2–4 m, not releasing white latex, without spines; young shoots glabrous; internodes of leafbearing sections of stem 3–10 3 ca. 2 mm, pale greybrown, not hollow, lacking a dark stain on the cut portion of the stem. Stipules 5–12 mm, narrowly lanceolate, forked, sparsely pubescent; petioles 6–90 3 ca. 0.5 mm, glabrous to sparsely pubescent toward leaf base, the hairs 0.25–0.375 mm, strongly appressed, straight; leaf blade 55–230 3 8–55 mm, narrowly lanceolate, lanceolate to oblanceolate, chartaceous to subcoriaceous; adaxial surface sparsely pubescent to glabrous, the hairs most frequent toward the leaf base, hairs 0.375–0.5 mm, weakly appressed, straight; the cystoliths punctiform, inflated, densely scattered; abaxial surface glabrous; the cystoliths punctiform to oblong, occasionally appearing inflated, scattered and parallel to the veins; primary veins 3, primary to tertiary and occasionally quarternary veins visible to the naked eye, the lateral primary veins finer than midrib and visible for 1/3 of leaf length; domatia present in the axils of the secondary veins, composed of hairs; base obtuse; margins asymmetrically discretely crenate to serrate; apex pungent to subacuminate. Peduncular bracts 1–2 mm; bracteoles 0.25–0.5 mm, staminate inflorescences ca. 15 per stem, peduncle unbranched from base for 6–9 mm, pubescent, the hairs to 0.25 mm, the whole inflorescence 5–40 mm, bearing ca. 110 flowers in a weakly asymmetrical to symmetrical cyme with 4 or 5 orders of dichotomous branching; the flowers subsessile, borne in clusters of 5 to 7; pistillate inflorescences 2 to 21 per stem (4 to 7 in fruit), the peduncle branched to base or unbranched at base for 2–12 mm (6–17 mm in fruit), sparsely pubescent, the hairs ca. 0.1 mm, erect, straight; the whole inflorescence 5–40 mm (23– 30 mm in fruit), longer than broad (or as long as broad), bearing 30 to 250 flowers (59 to 67 in fruit) in a symmetrical cyme with 4 or 5 orders of dichotomous branching, flowers pedicellate, borne in clusters of 3, rarely 2 to 4, the pedicels subsessile to 0.25 3 0.175 mm, glabrous. Staminate flowers 1–1.25 3 1.50–1.75 mm immediately prior to anthesis; tepals 4, ca. 2 mm; stamens and pistillode not seen. Pistillate flowers 0.4–0.6 3 0.25–0.6 mm, lateral tepals ca. 0.4 3 0.25 mm, ovate; dorsal tepal ca. 0.4 3 0.25 mm, ovate, the ventral tepal ca. 0.4 3 0.25 mm, ovate; stigma penicillate, erect. Fruit prior Annals of the Missouri Botanical Garden mobt-96-02-05.3d 13/5/09 20:10:04 276 Volume 96, Number 2 2009 Monro & Rodrı́guez New Species and Synopsis of Urera to inflation of tepals with achene entirely obscured by tepals, the laterals ca. 1.75 mm, the dorsal ca. 1 mm, the ventral ca. 0.675 mm, achene 1–1.375 3 0.75– 1.25 mm, elliptic, surface smooth, fruit when ripe with tepals inflated and berry-like, 1–1.25 3 ca. 2 mm, red. domatia; pedicels of staminate flowers greater than 1 mm; mature, fleshy fruit orange. Habitat and distribution. Urera guanacastensis is found in montane and cloud forest, disturbed and undisturbed forest, at 820–1350 m, known only from the Cordillera de Guanacaste in the Cantón de Liberia, Guanacaste Conservation Area in Costa Rica. IUCN Red List category. Conservation for Urera guanacastensis is considered as Near Threatened (NT) according to IUCN Red List criteria (IUCN, 2001). This is based on an evaluation of the potential distribution of U. guanacastensis, the threat to its habitat within that area, and the number of existing records for the species. Extrapolating from the 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 forest localities (Google Earth, 2008; collection label data) that form part of Costa Rica’s protected areas’ network. Currently, ca. 50% of the potential distribution is deforested (Google Earth, 2008) and no records exist from deforested localities. We therefore 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 Protected Areas Network. Etymology. Urera guanacastensis is named after the Guanacaste Conservation Area, where all known records of this species have been collected. Discussion. Urera guanacastensis is characterized by narrowly lanceolate, lanceolate, or oblanceolate, glabrous to sparsely pubescent leaves that frequently possess domatia on the abaxial surface composed of a cluster of hairs in the axils of the secondary veins, and by the red color of the mature fleshy fruits (not unique within the genus). It is most likely to be confused with U. simplex Wedd., from which it may be distinguished by leaf width, pubescence, and the presence of domatia, as well as by the length of the staminate pedicels and color of the fruit as follows: (1) for U. guanacastensis, young stems glabrous; leaves 8– 38 mm wide, adaxial surface very sparsely pubescent to glabrous, abaxial surface with domatia in the axils of the secondary veins; pedicels of staminate flowers less than 0.5 mm; mature, fleshy fruit red; (2) for U. simplex, young stems pubescent, frequently densely so; leaves 37–210 mm wide, adaxial surface pubescent, frequently densely so, abaxial surface lacking 277 Paratypes. COSTA RICA. Guanacaste: Cantón de Liberia, Parque Nac. Guanacaste, Cordillera de Guanacaste, Estación Cacao, sendero Maritza, L. Angulo 88 (INB); Estación Cacao, bosques primarios y orillas de bosque, W, E. Bello 2237 (INB); Estación Cacao, C. Chávez 95 (INB); Parque Nac. Rincón de La Vieja, Cordillera de Guanacaste, Estación Las Pailas, R. Espinoza 771 (INB); Estación Cacao, sendero a casa de Fran, B. Gamboa 45 (CR, INB); Estación Mengo, Volcán Cacao, sendero entre Estación y potrero Los Naranjos, W, B. Hammel & E. Chavarrı́a 17536 (BM); Estación Mengo, sendero el Potrero, lado sur, II INBio 183 (BM, INB); Estación Cacao, cerro Cacao, sendero a Casa Fran, W, A. Mora 37 (INB); Estación Cacao, cerro Cacao, M. Moraga 60 (INB); Estación Cacao, Cerro Cacao, W, F. Quesada 275 (INB); Sector Las Pailas, Rı́o Colorado, Aguas Arriba, G. Rivera 651 (INB); Cordillera de Guanacaste, sendero a laguna Santa Marı́a, W, G. Rivera 1152 (INB). 6. Urera killipiana Standl. & Steyerm., Fieldiana, Bot. 24: 427. 1952. TYPE: Guatemala. Quetzaltenango: Volcán Junil, 1700 m, 8 Aug. 1934, A. F. Skutch 982 (holotype, F!; isotype, GH!). Local names. Chichicaste (Guatemala: P. C. Standley 76980, F), chichicaste común (Guatemala: P. C. Standley 64714, F), nigüita (Guatemala: P. C. Standley 75549, F). Habitat and distribution. Disturbed vegetation, montane forest, riversides, from 900–3000 m, Mexico (Hidalgo, Queretaro, Oaxaca, Tabasco, Chiapas), Guatemala, and El Salvador. Comments. Urera killipiana is most frequently determined as U. eggersii (5 U. simplex) or U. caracasana. These species can be distinguished from each other based on venation, stipule morphology, and stem indumentum as follows: (1) for U. killipiana, young stems sparsely pubescent, stipule apex forked or not forked, the secondary veins of the abaxial leaf surface without domatia in their axils, tertiary venation of abaxial leaf surface noticeably paler than the lamina; (2) for U. caracasana, young stems densely pubescent, stipule apex not forked; the secondary veins of the abaxial leaf surface frequently with domatia (flap-like or tufts of hairs) in their axils, tertiary venation of abaxial leaf surface darker or of the same color tone as the lamina; (3) for U. simplex, young stems densely pubescent, stipule apex not forked; the secondary veins of the abaxial leaf surface without domatia in their axils, tertiary venation of abaxial leaf surface darker or of the same color tone as the lamina. Selected specimens examined. EL SALVADOR. San Salvador: S. Calderon 727 (GH, NY). GUATEMALA. San Marcos: San Marcos, Finca Armenia, Dwyer 15338 (MO). MEXICO. Chiapas: Ocozocoautla de Espinosa, T. B. Croat Annals of the Missouri Botanical Garden mobt-96-02-05.3d 13/5/09 20:10:04 277 278 Annals of the Missouri Botanical Garden 40599 (MO). Hidalgo: Puerto Obscuro near Km 328 on hwy. betw. Santa Ana & Chapulhuacán, Jacala Distr., H. E. Moore 5049 (BM). Oaxaca: B. Hammel & M. Merello 15457 (MO). Queretaro: 3–4 km al S de La Parada, Mpio. de Jalpan, B. Servin 557 (BM). Tabasco: C. L. Gilly & E. Hernandez X. 320 (GH, MEXU). caulis 5–10 mm in diametro cava, stipulis apice integris non furcatis, pistillodio minore atque achenio laevi differt. Vine, scrambling shrub, dioecious (?); main stems cane-like, to 25 m, young shoots pubescent to densely pubescent, the hairs 0.25–1 mm, erect or weakly appressed, weakly curved or crooked, occasionally straight; internodes of leaf-bearing sections of stem 21–84 3 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 4– 12 mm, narrowly ovate or lanceolate, not forked, sparsely pubescent to pubescent; petioles 11–90 (–170) 3 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 3 29–110 mm, narrowly ovate, obovate, elliptic, or oblanceolate, always longer than wide, chartaceous and occasionally bullate; adaxial surface sparsely pubescent, the hairs 0.5–0.75 mm, appressed, weakly curved or crooked; the cystoliths punctiform, oblong or occasionally fusiform, occasionally inflated, randomly scattered and occasionally parallel to veins, rarely arranged radially around a hair base; abaxial surface pubescent, the hairs 0.125– 0.5(1) mm, appressed or erect, weakly curved, occasionally straight; the cystoliths fusiform, occasionally oblong, parallel to the veins, occasionally randomly scattered; primary veins 3, occasionally 5, primary to quinternary veins visible to the naked eye, the lateral primary veins visible for 1/2–2/3 of the leaf length; domatia not present in the axils of the secondary veins; base subcordate or obtuse, occasionally cuneate; margins dentate, 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 from the 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 in an asymmetrical cyme with 4 or 5 orders of dichotomous branching; flowers borne in clusters of 6 to 12, sessile; pistillate inflorescences ca. 6 per stem, unbranched base of peduncle 7–13 mm, 5–11 mm in fruit, densely pubescent, the hairs 0.125– 0.25 mm, erect, straight; the whole inflorescence 12– 40 mm (12–45 mm in fruit), broader than long, bearing 112 to 448 flowers in 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 3 0.125–0.25 mm, glabrous. Staminate flowers ca. 1.25 3 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 flowers 0.5–1 3 0.375–0.5 mm, white to 7. Urera laciniata Goudot. ex Wedd., Ann. Sci. Nat., Bot., sér. 4, 18: 203. 1854. TYPE: [Colombia] Nouvelle Grenade. ‘‘Quindui’’ [Quindı́o?], La Bolsa, 1844, J. Goudot s.n. (lectotype, designated by de Rooij [1975: 308], P #00281783!). Urtica girardinioides Seem., Bot. Voy. Herald 194. 1854. TYPE: Panama. 1846–1849, B. Seemann 494 (lectotype, designated by de Rooij [1975: 308], BM!). Both epithets, Urera laciniata and Urtica girardinioides, were published in 1854. According to Stafleu and Cowan (1988: 139), the probable month of publication for Weddell’s publication is March, while that for Seemann’s publication is July (Stafleu & Cowan, 1985: 476), thereby giving priority to Urera laciniata. Local name. 9266, MO). Pringamoza (Panama: J. A. Duke Habitat and distribution. Urera laciniata has been collected from riverside scrub and from sea level to 1200 m. Its range extends from 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 unlikely to be confused with any other. Selected specimens examined. BOLIVIA. La Paz: Franz tamayo, Parque Nac. Madidi, senda Azariamas–San Fermin sector Mutún, E Ticona, A. Araujo M., V. Torrez, C. Perez, G. Jove & A. Urbano 149 (BM, MO). COSTA RICA. San José: A. F. Skutch 4266 (GH, MO, NY). HONDURAS. Gracias a Dios: J. Saunders 1204 (NY). NICARAGUA. Jinotega: Caño Litutus, Rı́o Bocay, W. D. Stevens, J. H. Beach, J. Schal & O. M. Montiel J. 16650 (BM, MO). PANAMA. s. loc.: 1846–1849, B. C. Seemann 494 (BM). PERU. Junin: Cataract El Tirol, A. K. Monro, R. T. Pennington & A. Daza 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 Espı́ritu Santo hasta bosque residual en el flanco noreste del mismo, 10u059200N, 84u249200W, 1000–1200 m, 24 Nov. 1988, G. Herrera 2326 (holotype, INB!; isotypes, BM!, MO!). Figure 2A–F. Species nova Urerae glabriusculae V. W. Steinm. similis, sed ab ea habitu lianiformi, parte foliifera Annals of the Missouri Botanical Garden mobt-96-02-05.3d 13/5/09 20:10:05 278 Volume 96, Number 2 2009 Monro & Rodrı́guez New Species and Synopsis of Urera 279 Figure 2. Urera lianoides A. K. Monro & Al. Rodr. —A. Habit with staminate inflorescences. —B. Cluster of staminate flowers. —C. Staminate flower at anthesis. —D. Leaf abaxial surface with cystoliths. —E. Habit, with pistillate inflorescences. —F. Immature fruit. (A–D: Johnson 1600 [GH]; E, F: Shank & Molina R. 4405 [NY]). Annals of the Missouri Botanical Garden mobt-96-02-05.3d 13/5/09 20:10:05 279 280 Annals of the Missouri Botanical Garden yellow-green; lateral tepals 0.375–0.75 mm, asymmetrically ovate, dorsal tepal 0.25–0.75 mm, ovate, with a subapical dorsal thickening; ventral tepal 0.25– 0.75 mm, ovate, stigma penicillate. Fruit prior to inflation of tepals ca. 1.5 3 0.75 mm; basal 3/4 of achene obscured by tepals, lateral tepals ca. 1 mm, dorsal tepals ca. 0.75 mm, ventral tepals ca. 0.5 mm; achene 1.25–1.5 3 1–1.25 mm, asymmetrically elliptic, keel-shaped, surface smooth. Fruit when ripe, with tepals inflated and berry-like 1.5–2.25 3 1.25– 1.75 mm, orange when fresh. Provincia Arce, 108 km de Tarija hacia Bermejo, R. Ehrich 494 (K). COSTA RICA. Alajuela: Cantón de San Ramón, Reserva Biológica Monteverde, Cordillera de Tilarán, Refugio de Eladio, Valle del Rı́o Peñas Blancas, E. Bello 5181 (INB); La Palma de San Ramón, A. M. Brenes 5793 (F, NY); Mata Cartago, La Palma de San Ramón, A. M. Brenes 6308 (F, NY); quebrada Azul (San Carlos), A. M. Brenes 23063 (NY); Guatuso, Parque Nac. Volcán Tenorio, El Pilón, colectado en bosque primario a orillas de sendero Los Misterios del Tenorio, J. L. Chávez 874 (INB); Cantón de Los Chiles, Refugio Natural de Vida Silvestre Caño Negro, Rı́o Frı́o, Finca Betel, K. Flores & M. Flores 130 (INB); Reserva Biológica Monteverde, Rı́o Peñas Blancas, Finca Wilson Salazar, vertiente Atlántica, W. Haber & E. Bello 7403 (INB, MO); Reserva Monteverde, Poco Sol 13 km S Fortuna, W. Haber & W. Zuchowski 9377 (BM, INB); Upala, Bijagua, El Pilón, cabecera del Rı́o Celeste, G. Herrera 1293 (BM, INB); N side of Volcán Arenal, R. W. Lent 2922 (F); Upala, Zona Protectora Miravalles, 3 km W of Bijagua, ridges above Rı́o Zapote, D. S. Penneys & W. Haber 710 (INB); Parque Nac. Rincón de La Vieja, Colonia La Libertad, Finca Julio Soto, G. Rivera 1485 (INB); Colonia Blanca, Finca Rı́o Negro, G. Rivera 1563 (INB); San Carlos, Parque Nac. Arenal, Cerro Chato, sendero que lleva a la Laguna, A. Rodrı́guez, V. H. Ramı́rez, G. de La O & G. Soto 6254 (INB); Colegio Agropecuario, Santa Clara, Cantón de San Carlos, A. S. Weston, D. F. Weston & J. Weston 3104 (MO). Cartago: Cantón de Turrialba, Valle del Reventazón, Carolina, Moravia de Chirripó, P. Campos 176 (INB); 1/2 km S of Chitarı́a, near rd. #CR233, R. W. Lent 237 (NY). Guanacaste: Cantón de Tilarán, Cordillera de Tilarán, 1– 2 km W of Lago de Cote, 13 km N of Tilarán, continental divide on SE, slope of Volcán Tenorio, W. Haber & W. Zuchowski 11624 (INB). Heredia: L. R. Holdridge’s Finca La Selva, Rı́o Puerto Viejo at quebrada El Sura & quebrada El Salto, ca. 1 mi. above jct. Rı́o Sarapiqui, G. B. Rossbach 3690 (GH); roadside bank about 35 km NE of Alajuela, R. J. Taylor 4536 (NY). Limón: Level areas betw. Siquerres & Rı́o Pacuare, steep hills S of the railroad bridge over the Rı́o Pacuare, W. C. Burger & R. L. Liesner 6952 (NY); borde de lago Dabagri hasta Rı́o Llei, L. D. Gómez et al. 23129 (BM); Costado Oeste de lago Dabagri hasta Rı́o Llei, L. D. Gómez et al. 23172 (BM); vecino al puente sobre el Rı́o Toro Amarillo, 5.5 km al SO de Guápiles, A. Jimenez M. 1067 (F); pantanosos-yolillosos de Suerre y Dos Bocas, drenajes de los Rı́os Parismina y Reventazón, P. J. Shank & A. Molina R. 4265 (GH); pantanosos-yolillosos de Goldengrove, drenaje de Rı́o Reventazón, P. J. Shank & A. Molina R. 4405 (F). Puntarenas: Cantón de Osa, Refugio Forestal Golfo Dulce, Penı́nsula de Osa, Los Mogos, Bahı́a Chal, entrada Chocuaco, R. Aguilar 3591 (INB); foothills of Cordillera de Talamanca, just N of Santa Elena on Fila Cotón, S of Agua Caliente, G. Davidse, G. Herrera Ch. & M. H. Grayum 28277 (BM); Osa, Ballena, entrando por Puertecito, hasta el Cerro Escalera, J. González & C. Aragón 2327 (INB); Cantón de Coto Brus, Parque Internacional La Amistad, Cordillera de Talamanca, Estación Pittier, Agua Caliente, E. Navarro 211 (INB); cantón de Osa, Penı́nsula de Osa, Rancho Quemado, F. Quesada 167 (INB); deep forest near airport, 4 mi. W of Rincón de Osa, Osa Penı́nsula, P. H. Raven 21623 (MO); Canton de Osa, Refugio Forestal Golfo Dulce, Penı́nsula de Osa, Rancho Quemado, Fila Guerra, Rincón, finca del Grupo de Conservación, M. Segura 4 (INB); Reserva Biológica Monteverde, Cordillera de Tilarán, San Luis de Monteverde, bosques en la cuenca del Rı́o San Luis, K. Taylor 118 (INB); Rincón de Osa, area N of airport, J. Utley & K. Utley 1173 (F). San José: Canton de Pérez Zeledón, Cordillera de Habitat and distribution. The new species is found in premontane, montane, and cloud forest, in disturbed and undisturbed forest, from sea level to 1300(1900–2500) m. Mexico (Chiapas), Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Peru, and Bolivia. IUCN Red List category. Conservation for 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 times in several localities across Central and South America. Etymology. From the English, ‘‘liana’’ or ‘‘vine,’’ derived from the French, ‘‘lier,’’ which, in turn, is derived from the Latin ‘‘ligare’’ or ‘‘to tie,’’ referring to the liana-like habit of this species. Discussion. Urera lianoides is distinguished by its scandent habit, hollow stems, sparse non-urticating hairs, and orange, mature, fleshy fruit. Material of U. lianoides from Chiapas differs from other collections of this species in having densely pubescent young stem and abaxial leaf surfaces, with very short hairs (0.5 mm or less). This species corresponds to ‘‘sp. A’’ in the treatment of Urera in the Flora de Nicaragua (Pool, 2001: 2495). Urera lianoides most closely resembles U. glabriuscula with which it shares a distribution in Chiapas, Mexico. The two species can be distinguished from each other as follows: (1) for U. lianoides, a vine or scrambling shrub, ca. 5–10 mm diam., leaf-bearing stem section hollow, with internodes greater than 20 mm; stipule apex not forked; pistillode 0.375 mm diam.; achene surface smooth; (2) for U. glabriuscula, a shrub or small tree, ca. 5– 10 mm diam., leaf-bearing stem section never hollow, with internodes less than 20 mm; stipule apex minutely forked; pistillode 0.75 mm diam.; achene surface verrucate. Paratypes. BOLIVIA. Beni: Provincia Ballivian, Pilón Laja, 130 km de San Borja, M. Moraes 670 (K). La Paz: Provincia Nor Yungas, ca. 2 km al S de Coroico, S. G. Beck 21926 (K); Santa Fé Tacana, al lado de la parcela permanente de Santa Fé, G. Bourdy 1746 (K). Tarija: Annals of the Missouri Botanical Garden mobt-96-02-05.3d 13/5/09 20:10:07 280 Volume 96, Number 2 2009 Monro & Rodrı́guez New Species and Synopsis of Urera 281 Talamanca, Las Nubes, Santa Elena, E. Alfaro 329 (INB); W part of montañas Jamaica, ca. 3 km NE of Bijagual de Turrubares, Carara reserve, M. H. Grayum et al. 5878 (BM); Zona Protectora La Cangreja, Mastatal de Puriscal, J. F. Morales 727 (INB); Acosta, Colorado, Fila San Jerónimo, Rı́o Colorado, J. F. Morales 7383 (INB); Acosta, Aserrı́, Agua Buena, cabeceras Quebrada Laja, J. F. Morales et al. 9937 (INB); vic. of El General, A. F. Skutch 2863 (GH, MO, NY). GUATEMALA. Petén: Dolores, ca. 100 m del cementario, lado N, R. Tún O. 1323 (BM, F). Suchitepequez: Finca Mocá, A. F. Skutch 1479 (GH). HONDURAS. Atlantida: Vic. of La Ceiba, bank of Danto river, slopes of Mount Cangrejal, T. G. Yuncker, J. M. Koepper & K. A. Wagner 8454 (GH, MO, NY). Cortés: Aldea de Corinto y alrededores frontera con Guatemala, 55 km al O de Puerto Cortés, C. Nelson, E. Vargas, M. Erazo, M. Garcı́a & M. Sierra 2926A (BM, MO). Olancho: Vaguada del Rı́o de la población de Culmı́, C. Nelson & E. Romero 4738 (MO). MEXICO. Chiapas: 2–4 km below Ixhuatán along rd. to Pichucalco, Muni. of Solosuchiapa, D. E. Breedlove 19906 (MO); long gravel rd. betw. Palenque & Bonampak, 60 mi. SE of Palenque, T. B. Croat 40192 (MEXU, MO); along gravel rd. betw. Palenque & Bonampak, 88–90 mi. SE of Palenque, T. B. Croat 40230 (MO); along hwy. 195, betw. Chiapa de Corzo & Pichucalco, ca. 6 mi. NW of Pueblo Nuevo Solistahuacán, along mirador overlooking Caribbean slope, Selva Negra lookout point at trail to Colonia Pinabeto, vic. Km 99, T. B. Croat & D. P. Hannon 65180 (MO); Crucero Corozal, camino Palenque–Boca Lacantum, Mpio. Ococingo, E. Martı́nez S. 15439 (BM); a 14 km al NW de Crucero Corozal sobre el camino Palenque–Boca Lacantum, Mpio. Ococingo, E. Martı́nez S. 16642 (BM). Veracruz: camino Bastonal– Tebanca, Mpio. Catemaco, R. Cedillo T. 3439 (BM); Estación Biológica Los Tuxtlas, B. Hammel, M. Merello & S. Sinaca 15492 (MO). NICARAGUA. Chontales: On ridge top of Cordillera Amerisque, A.Gentry, W. D. Stevens, A. Grijalva P. & P. P. Moreno 43950 (MO). Granada: Volcán Mombacho, Hacienda UPE-Pancasán (antes del cráter), A. Grijalva, O. Vanegas & R. M. Rueda 2924 (BM); NE del Volcán Mombacho, en el emplame de los caminos a Sta. Isabel y a Cutirre, P. Moreno 2602 (BM, MO); Volcán Mombacho, subiendo por la Finca Las Delicias, 1.5 km de la hacienda, P. P. Moreno 4103 (MO); NW slopes of Volcán Mombacho, 10 km S of Granada, M. Nee & J. Miller 27693 (MO); NW de Volcán Mombacho, cafetales de Finca Cutirre y camino que lleva S del volcán, J. C. Sandino 1273 (BM, MO). Zelaya [Región Autónoma del Atlántico Sur]: along the rd. betw. Nueva Guinea & Verdún, J. S. Miller & J. C. Sandino 1094 (MO); Monkey Point, Caño el Pato, 1.5 km sobre la ribera del Caño, P. P. Moreno 12397 (MO). PANAMA. s. loc.: S. Hayes 683 (K); Canal area, vic. of Madden Dam, P. H. Allen 2008 (F, NY); Barro Colorado Island, S. Aviles 106 (F); rd. S-11, NW of Escobal, T. B. Croat 12466 (BM, NY); rd. along Rı́o Piña–Rı́o Media divide, NW part of Canal Zone (area W of Limon Bay, Gatun Locks & Gatun Lake), I. M. Johnston 1600 (GH); rd. along W side of Gatun lake, NW part of Canal Zone (area W of Limón Bay, Gatun Locks & Gatun Lake), I. M. Johnston 1706 (GH); 12 mi. S of Colón, E. L. Tyson et al. 4478 (GH). Bocas del Toro: Ridge N of Campamento Luchio, A. K. Monro & E. Alfaro 4506 (BM, INB, MEXU, MO, PMA); Rı́o Cricamola, betw. Finca St. Louis & Konkintoë, R. E. Woodson, P. H. Allen & R. J. Seibert 1923 (GH, NY); 10–15 mi. inland S from mouth of Changuinola River, W. H. Lewis, J. D. Dwyer, T. S. Elias & K. R. Robertson 867 (GH, K, UC, US). Chiriquı́: Bajo Chorro, Boquete Distr., M. E. Davidson 349 (MO). Coclé: Foothills of Cerro Pilón, near El Valle, J. Duke & M. Correa 14670(1) (MO); El Valle from potato farm above village to Cerro Pilon, J. Dwyer & M. Correa 7923 (BM). Colón: From Portobelo hwy. to 4 km up Rı́o Guanche, S. Knapp 1016 (BM); Rı́o Guache, K. J. Sytsma 1617 (BM). Darién: Parque Nac. Darién, Serranı́a de Sapo, lı́mite del Parque hasta la cima, H. Herrera & J. Polanco 795 (BM). Panamá: Piriatı́, S of Pan-American Hwy., C. Hamilton 558 (BM). PERU. Madre de Dios: Parque Nac. del Manu, Rı́o Manu, Cocha Cashu Station, J. Terborgh & R. B. Foster 6488 (K). 9. Urera simplex Wedd., Prodr. (DC.) 16(1): 90. 1869. TYPE. Colombia. Cundinamarca: ‘‘ad salto de Tequendama,’’ Mar. 1856, Triana s.n. (holotype, P 00281785!). Urera eggersii Hieron., Bot. Jahrb. Syst. 20: 3. 1895. TYPE: Ecuador. Pichincha: 20 km W of Santo Domingo de los Colorados, 1000 ft., 25 Nov. 1961, P. C. D. Cazalet & T. D. Pennington 5150 (neotype, designated here, K!; isotypes, B not seen, NY not seen). Urera rzedowskii V. W. Steinm., Acta Bot. Mex. 71: 37. 2005. TYPE: Mexico. Veracruz: Mpio. San Andrés Tuxtla, 8 km al N de San Andrés Tuxtla, Laguna Encantada, 18u289N, 95u109W, 350 m, 4 Apr. 1981, J. I. Calzada 8105 (holotype, IEB not seen; isotype, ENCB not seen). Urera tuerckheimii Donn. Sm., Bot. Gaz. 23(1): 14. 1897. TYPE: Guatemala. Alta Verapaz: Pansamalá, 1160 m, May 1887, H. von Tuerckheim 1243 (holotype, US not seen; isotype, NY!). A neotype was selected for Urera eggersii because the type collection cited by Hieronymus, Eggers 14466 (B), was destroyed in enemy action during World War II and only photographs of the holotype could be located (F, MO). The neotype was selected on the basis that it includes good leaf and fertile material and was from the same country as the holotype. Local names. Bilsimtezla (Mexico: A. Méndez T. 6738, BM), chenek’mut (Mexico: A. Méndez T. 4863, BM), chichicaste (Guatemala: P. C. Standley 68232, F; Mexico: M. Heath & A. Long MA 44, BM), chichicaste huevo de cangrejo (El Salvador: E. Sandoval & H. Rivera 1252, MO), sakil zulsimtez laa (Mexico: A. Méndez T. 6238, BM), tzotzniz zul simtez (A. Méndez T. 9066, BM), zulsimtezla (Mexico: A. Méndez T. 7022, BM). Habitat and distribution. Disturbed and undisturbed forest, cloud forest, and humid scrub from sea level to 2500 m. Mexico (Chiapas, Tabasco, Veracruz), Belize, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Peru, Bolivia, Brazil. Comments. Material of this species has frequently been determined and referred to (Flora of Guatemala, Flora de Nicaragua, and Flora Costaricensis) as Urera elata, U. eggersii, or U. tuerckheimii. Examination of type material of these species indicates that U. elata is a species endemic to Jamaica, while U. eggersii and Annals of the Missouri Botanical Garden mobt-96-02-05.3d 13/5/09 20:10:08 281 282 Annals of the Missouri Botanical Garden U. tuerckheimii are conspecific with U. simplex. Pool (2001) indicates U. tuerckheimii (5 U. simplex) may correspond to U. aurantiaca Wedd. from South America (Argentina, Bolivia, Paraguay); however, comparison of the holotypes suggests that the two species are distinct, with U. aurantica characterized by ovate or cordiform leaves and relatively short pistillate inflorescences and known only from South America (Argentina, Bolivia, Brazil, and Paraguay). Some collections of U. simplex from Costa Rica and Panama, e.g., Kennedy 1939 (GH) and Folsom y Page 5984 (PMA), are unusual in the possession of narrowly oblanceolate pubescent leaves, while some material from Chiapas (Purpus 7039, NY) is characterized by densely pubescent leaves. Urera simplex most closely resembles U. elata. The two species can be distinguished from each other based on leaf texture and inflorescence peduncle size as follows: (1) for U. simplex, petiole lacking small spines; staminate flowers 5-parted, occasionally 4-parted; (2) for U. elata, petiole with small spines; staminate flowers 4-parted. mala, Honduras, El Salvador, Costa Rica, Panama, Peru, and Bolivia. Selected specimens examined. BELIZE. Stann Creek: Middlesex, W. A. Schipp 400 (BM, F, GH, NY). BOLIVIA. La Paz: N Yungas, valle de Huarinillas, Estación Biológica Tunquini, S. G. Beck 24612 (K, LPB). BRAZIL. Amazonas: E. Ule 5465 (K). COLOMBIA. Putumayo: Umbrı́a, G. Klug 1741 (K). COSTA RICA. Puntarenas: Cantón de Golfito Dos Brazos de Rı́o Tigre, Jiménez, orilla de Quebrada Pizote, G. Cordero 95 (BM, INB, MO). ECUADOR. Pichincha: P. C. D. Cazalet & T. D. Pennington 5073 (K). EL SALVADOR. Ahuachapán: Finca L’Esperanza, Jujutla, A. K. Monro et al. 2997 (BM, ITIC, LAGU, MO). HONDURAS. Olancho: Catacamas, Rı́o Catacamas, slope of Sierra de Agalto, S. Blackmore & G. L. A. Heath 1916 (BM). MEXICO. Chiapas: Finca Mexiquito, C. A. Purpus 7039 (BM, F, MO, NY). NICARAGUA. Matagalpa: Macizo de Peñas Blancas, Finca San Sebastian, O. Téllez V., R. Riviere, W. D. Stevens, O. M. Montiel J., M. Guzmán & D. Castro 5181 (BM, MEXU). PANAMA. Comarca de San Blas: Udirbi Reserve, along park boundary, J. F. McDonagh et al. 257 (BM, MO). PERU. Huanuco: Vic. of Tingo Marı́a cliffs above Rı́o Monzon, M. E. Mathias & D. Taylor 5343 (K). 10. Urera verrucosa (Liebm.) V. W. Steinm., Acta Bot. Mex. 71: 39. 2005. Basionym: Urtica verrucosa Liebm., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 5, 2: 295. 1851. Urera caracasana var. tomentosa (Wedd.) Wedd., Prodr. 16: 90. 1869. nom. illeg., superfl. TYPE: Costa Rica. Cartago: Irasú, Oersted 14284 (holotype, C!). Local names. BM). Chichicaste (Mexico: E. Kerber 333, Habitat and distribution. Montane forest, shade coffee farms, Pinus–Quercus–Liquidambar forest, from 500–2800 m. Mexico (Chiapas, Veracruz), Guate- Comments. Urera verrucosa most closely resembles U. caracasana. The two species can be distinguished from each other based on leaf texture and inflorescence peduncle size as follows: (1) for U. verrucosa, leaves bullate; staminate peduncle unbranched at base for 40– 80 mm, densely pubescent; pistillate peduncle unbranched at base for 27–98 mm; (2) for U. caracasana, leaves chartaceous; staminate peduncle branched to base or unbranched at base for 2–13 mm; pistillate peduncle branched to base or unbranched at base for 2–20 mm. Selected specimens examined. BOLIVIA. La Paz: M. Lewis 882155 (K). COSTA RICA. Cartago: N of Cartago, Rı́o Reventada, R. Khan, M. Tebbs & A. R. Vickery 959 (BM). EL SALVADOR. Ahuachapán: Lago de Ninfas, Cordillera Grande de Apaneca, NW of Juayua, G. Davidse, K. Sidwell, A. Monro, M. A. Renderos & C. Cortez. 37383 (BM, ITIC, LAGU, MO). GUATEMALA. Chimaltenango: On hwy. CA1 betw. turnoff to Patzúm & Sololá, 14.8 mi. NNW of turnoff to Patzúm, 23.5 mi. SSE of turnoff to Sololá, T. B. Croat & D. P. Hannon 64732 (BM, MO). MEXICO. Veracruz: Orizaba, M. Botteri 288 (BM). PANAMA. Chiriquı́: 3 km NW of Cerro Punta, along dirt rd. on rte. to Las Nubes, B. Hammel 1363 (BM, MO, NY). PERU. San Martı́n: T. D. Pennington & A. Daza 16676 (K, MOL). Literature Cited Badilla, B., G. Mora, A. J. Lapa & J. A. Silva E. 1999. Antiinflammatory activity of Urera baccifera (Urticaceae) in Sprague-Dawley rats. Revista Biol. Trop. 47: 365–371. Britton, N. L. & P. Wilson. 1924. Botany of Porto Rico and the Virgin Islands. Sci. Surv. Porto Rico & Virgin Islands 5: 243. Burger, W. 1977. Urera. In Flora Costaricensis. Fieldiana, Bot. 40: 276–280. Davidse, G., M. S. Sousa & A. O. Chater. 1994. Introducción general. Pp. xiii–xiv in G. Davidse, M. S. Sousa & A. O. Chater (editors), Flora Mesoamericana, Vol. 6, Alismataceae a Cyperaceae. Universidad Nacional Autónoma de México, México D. F.; Missouri Botanical Garden, St. Louis; The Natural History Museum, London. de Rooij, M. J. M. 1975. Urera. Pp. 301–309 in J. Lanjouw & A. L. Stoffers (editors), Flora of Suriname, Vol. 5(1). E. J. Brill, Leiden. Field Museum of Natural History. 2006. The Botany Collections Database. Department of Botany, Field Museum of Natural History, Chicago. ,http://emuweb. fieldmuseum.org/botany/Query.php., accessed 3 September 2008. Friis, I. 1989. The Urticaceae: A systematic review. Pp. 285– 308 in P. R. Crane & S. Blackmore (editors), Evolution, Systematics, and Fossil History of the Hamamelidae, Vol. 2. Systematics Association Special Volume 40B. Oxford Science Publications, Oxford, United Kingdom. Gaudichaud-Beaupré, C. 1830. Urera. Pp. 496–497 in Voyage autour du Monde, entrepris par ordre du roi,. . . exécuté sur les corvettes de S.M. l’Uranie et la Physicienne. . . par M. Louis de Freycinet. Botanique. Pilletainé, Paris. Annals of the Missouri Botanical Garden mobt-96-02-05.3d 13/5/09 20:10:08 282 Volume 96, Number 2 2009 Monro & Rodrı́guez New Species and Synopsis of Urera Gonzáles, J. C. 1994. Botánica Medicinal Popular, Etnobotánica Medicinal de El Salvador. Jardı́n Botánico La Laguna, Cuscatlán, El Salvador. Google Earth. 2008. ,http://earth.google.com., accessed 12 April 2008. Guánchez, F. J. 1999. Plantas amazónicas de uso medicinal y mágico. Fundación Polar, Servicio Autónomo para el Desarrollo Ambiental del Amazonas, Caracas. House, P. R., S. Lagos-Witte, L. Ochoa, C. Torres, T. Mejı́a & M. Rivas. 1995. Plantas Medicinales Comúnes de Honduras. Litografı́a López, S. de R. L. Universidad Nacional Autónoma de Honduras/Comité Internacional de Medicina Natural en Honduras/Cooperación Internacional para el Desarrollo (Programa de Cooperación Técnica Británica en Honduras)/Deutsche Gessellshaft für Technische Zusammenaerbeit, Tegucigalpa. International Plant Names Index. 2008. ,http://www.ipni. org., accessed 19 June 2008. IUCN. 2001. IUCN Red List Categories and Criteria, Version 3.1. Prepared by the IUCN Species Survival Commission. IUCN, Gland, Switzerland and Cambridge, United Kingdom. Janzen, D. H. & W. Hallwachs. Caterpillar food plants of the Area de Conservación Guanacaste, northwestern Costa Rica. ,http://janzen.sas.upenn.edu/Wadults/searchfood. lasso., accessed 9 November 2005. Linneaus, C. 1763. Species Plantarum, ed. 2. Stockholm. Miquel, F. A. W. 1853. Urera. Pp. 188–193 in Flora Brasiliensis (Martius), Vol. 4(1). Fleischer, Leipzig. Monro, A. K. 2006. The revision of species-rich genera: A phylogenetic framework for the strategic revision of Pilea (Urticaceae) based on cpDNA, nrDNA, and morphology. Amer. J. Bot. 93: 426–441. Pool, A. 2001. Urera. In W. D. Stevens, C. Ulloa, A. Pool & O. M. Montiel (editors), Flora de Nicaragua. Monogr. Syst. Bot. Missouri Bot. Gard. 85: 2492–2495. Stafleu, F. A. & R. S. Cowan. 1985. Taxonomic Literature, 2nd ed., Vol. 5. Bohn, Scheltema & Holkena, Utrecht/Antwerpen, and Dr. W. Junk b.v. Publishers, The Hague/Boston. ———, ——— & H. A. Weddell. 1988. Taxonomic Literature, 2nd ed., Vol. 7. Bohn, Scheltema & Holkena, Utrecht/Antwerpen, and Dr. W. Junk b.v. Publishers, The Hague/Boston. Standley, P. C. & J. A. Steyermark. 1952. Urera. In Flora of Guatemala. Fieldiana, Bot. 3: 424–428. Steinmann, V. W. 2005. Four new neotropical species and a new combination of Urera (Urticaceae). Acta Bot. Mex. 71: 19–43. Vellozo, J. M. C. 1827 [1831]. Florae Fluminensis 10. Typographia Nationali, Rio de Janeiro. Weddell, H. A. 1856. Urera. Pp. 143–162 in G. Baudry & J. Baudry (editors), Monographie de la Famille des Urticacées. G. and J. Baudry, Paris. ———. 1869. Urera. Pp. 199–203 in A. de Candolle (editor), Prodromus (D.C.), Vol. 16. Paris. 4. U. glabriuscula V. W. Steinm. 5. U. guanacastensis A. K. Monro & Al. Rodr. 6. U. killipiana Standl. & Steyerm. 7. U. laciniata Goudot. ex Wedd. 8. U. lianoides A. K. Monro & Al. Rodr. 9. U. simplex Wedd. 10. U. verrucosa (Liebm.) V. W. Steinm. APPENDIX 1. Index to Exsiccatae. Collections are listed alphabetically by collector name and then in ascending numerical order. Species are numbered as in the list provided here. LIST OF SPECIES 1. Urera baccifera (L.) Gaudich. ex Wedd. 2. U. caracasana (Jacq.) Griseb. 3. U. fenestrata A. K. Monro & Al. Rodr. 283 Acosta, L. 480 (3), 486 (3); Aguilar, I. 387 (10); Aguilar, R. et al. 3591 (8); Alclos S., J. 92 (9); Carlson, M. C. 391 (10), 2056 (1); Alcrasquilla, L. 180 (10); Alfaro, E. 329 (8), 745 (3), 1260 (3), 1569 (3); Allen, P. H. 911 (cf. 2), 926 (1), 1471 (10), 2008 (8), 3617 (9), 6315 (1), 6325 (9), 6330 (2), 6946 (10), 855 (cf. 2); Anderson, R. & Mori, S. 147 (1); Angulo, L. 88 (5); Antonio, T. 611 (1), 1118 (9), 1738 (9), 3528 (2), 4988 (9), 5111 (3); Araquistain, M. 3182 (9); Araquistain, M. & Castro, D. 1905 (1); Araquistain, M. & Moreno, P. P. 1502 (2), 2009 (1), 2235 (1), 2285 (1), 2358 (9), 2473 (9), 2605 (9), 2790 (9), 2863 (1); Atwood, J. T. & Moore, A. D. 304 (2), 489a (1); Aviles, S. 32 (9), 106 (8). Baker, C. F. 2011 (2); Baker, R. R53 (2); Bangham, W. N. 211 (9); Barkley, F. A. & Barkley, E. D. 40130 (1); Barkley, F. A. & Hernandez R., J. 40263 (1); Barkley, F. A. & LeivaWelchez, L. 39579 (2); Barkley, F. A. & Smith, M. L. 40859 (9); Barrelier, M. 11 (9); Bello, E. 2237 (5), 3143 (2), 5181 (8); Bittner, J. 1882 (3), 1894 (3); Blackmore, S. & Heath, G. L. A. 1916 (9), Botteri, M. 288 ‘854’ (10); Boyle, B. 2450 (3); Breedlove, D. E. 9107 (1), 9994 (4), 10003 (5), 19906 (8), 20207 (9), 23388 (4), 24109 (9), 24249 (4), 24998 (9), 34570 (4), 51467 (5); Breedlove, D. E. & Raven, P. H. 13559 (4), 13583 (9); Breedlove, D. E. & Smith, A. R. 19905 (9), 21681 (9), 31520 (4); Breedlove, D. E. & Thorne, R. F. 30986 (10); Brenes, A. M. 136 (5544) (3), 220 (4208) (3), 339 (9), 3480 (9), 3596 (9), 5524 (2), 5793 (8), 6039 (2), 6308 (8), 6764 (9), 23063 (8); Brown, E. 6 (2); Bunting, G. S. & Licht, L. 1137 (1); Burch, D. 4603 (9); Burger, W. C. 4161 (1); Burger, W. C. & Antonio, T. 10956 (9), 10993 (1); Burger, W. C. & Baker, R. 10013 (1); Burger, W. C. & Burger, M. 7678 (9), 8496 (9); Burger, W. C. & Gentry, J. L. 8639 (3), 8752 (3), 8836 (9), 8984 (9), 9092 (7); Burger, W. C. & Liesner, R. L. 6605 (2), 6879 (1), 6952 (8); Burger, W. C. & Matta U., G. 4819 (9); Burger, W. C. & Stolze, R. G. 4912 (3), 5608 (9), 5904 (2); Burger, W. C. et al. 10502 (1), 10563 (1). Cabrera, E. et al. 2666 (4); Cafferty, S. & Monro, A. K. 16 (9); Calderón, S. 727 (5), 850 (2), 1510 (2), 1539 (1), 1687 (2), 1775 (2), 1809 (2); Calonico S., J. et al. 21101 (4), 21242 (4); Campos, P. (8); Chacón, I. A. 1881 (3); Chavarria, U. 303 (2); Chavelas, P. et al. ES-2280 (9); Chávez, C. 95 (5); Chávez, J. L. 874 (8); Chickering, A. M. 126 (9); Chorley, M. & R. Atkinson 88 (2); Christopherson, E. D. 178 (10); Chrysler, M. A. 5612 (2); Conrad, J. et al. 2796 (4), 2815 (9); Cooper, G. P. & Slater, G. M. 178 (9); Cordero, G. 95 (9); Correa, M. D. et al. 2368 (2); Cosentino, K. 83 (9); Cowan, C. P. 1721 (9), 3064 (9), 3109 (9), 3151 (9); Cowan, C. P. et al. 3951 (9); Cowell, J. F. 257 (1); Croat, T. B. 10593 (3), 10597 (10), 12333 (9), 12466 (8), 15895 (9), 15897 (9), 23728 (9), 24254 (9), 26599 (9), 35138 (9), 36151 (3), 36777 (9), 40192 (8), 40230 (8), 40599 (5), 40862 (9), 40956 (8 aff.), 43488 (3), 43886 (9), 47437 (10), 49785 (7), 66252 (9), 66612 (2), 66795 (2), 68307 (1), 68317 (9), 78493 (2), 78524 (9), 78545 (9); Croat, T. B. & Hannon, D. P. 63356 (cf. 2), 64732 (10), 65180 (8), 65332 (9), 65354 (4); Croat, T. B. & Porter, D. M. 15663 (9); Croat, T. B. & Zhu, G. 76527 (9); Cruz, R. 31 (1), s.n. ‘WB-1176’ (10). D’Arcy, W. G. 4239 (9), 10767 (9), 10998 (10); D’Arcy, W. G. & Hammel, B. 12235 (9); D’Arcy, W. G. & Sytsma, K. 14516 (cf. 2); D’Arcy, W. et al. 12658 (3); Darı́o, M. 461 (9); Darwin, S. et al. 2148 (1); Davidse, G. et al. 28277 (8), 37383 Annals of the Missouri Botanical Garden mobt-96-02-05.3d 13/5/09 20:10:09 283 284 Annals of the Missouri Botanical Garden (10), 37493 (1); Davidson, C. & Donahue, J. 8353 (9); Davidson, C. 7202 (9); Davidson, M. E. 349 (3), 484 (9), 487 (10); Deam, C. C. 85 (1); Delgado, R. 24 (5), 76 (9); Delprete, P. 5150 (9); Döbbeler, P. 5162 (3); Douglas, W. & Krukoff, B. A. 3522 (2), 4001 (2); Dryer, V. J. 179 (3), 233 (3), 234 (3), 1646 (10); Duke, J. A. 9020 (10), 9266 (7), 11844 (9), 12008 (7); Dunlap, V. C. 174 (2), 241 (9); Dunn, D. et al. 23234 (2); Dwyer, J. D. 2414 (2), 2895 (9), 15338 (5), 15365 (2); Dwyer, J. D. & Correa, M. 7923 (8), 14670 (1) (8); Dwyer, J. D. & Correa A., M. D. 7503 (10), 7964 (9); Dwyer, J. D. et al. 4837 (9). Ebinger, J. E. 812 (9), 967 (2); Echevarria C., J. A. 207 (10), C. 268 (cf. 2); Edwards, J. B. 107 (1), P-107 (1); Espinoza, R. 771 (5). Fernández, R. & Acosta-Zamudio, N. 2204 (4); Flores, K. & Flores, M. 130 (8); Folsom, J. P. 5932 (1); Folsom, J. P. et al. 2240 (3), 5571 (9); Fonseca Z., A. 79 (2), 113 (9); Fosberg, F. R. 27335 (7); Frankie, G. W. 79c (2), 138c (9). Gamboa, B. 45 (5), Garcı́a, A. R. & Martı́nez, E. 47 (1); Garwood, N. C. et al. 386 (3), 1134 (1), 2728 (9); Gaumer, G. F. 501 (1), 936 (1); Gentle, P. H. 2118 (9), 2781 (1), 2808 (9), 2819 (9); Gentry, A. 2043 (3), 6246 (9), 6648 (1), 7897 (9), 8427 (9); Gentry, A. et al. 43950 (8), 43976 (9), 79328 (9); Gilly, C. L. & Hernández X., E. 320 (5); Gómez L. 5535 (3), 8078 (3); Gómez P. 2207 (9), 21040 (9), 23129 (8), 23172 (8); González, J. & Aragón, C. 2327 (8); Grayum, M. H. & R. Evans 9866 (9); Grayum, M. H. 7237 (cf. 3); Grayum, M. H. et al. 5878 (8); Grijalva, A. & Burgos, F. 1547 (9); Grijalva, A. 1916 (2); Grijalva, A. et al. 2906 (2), 2924 (8); Guadalupe J., S. & Lessette F., F. 33 (2). Haber, W. ex Bello, E. 6403 (1); Haber, W. & E. Bello 7403 (8); Haber, W. & Zuchowski, W. 9293 (9), 9294 (2), 9377 (8); 10889 (10), 10907 (9); Haber, W. et al. 11279 (9), 11296 (cf. 2); Hall, J. S. & Bockus, S. M. 7564 (2); Hamilton, C. & Stockwell, H. 3537 (9); Hamilton, C. 558 (8); Hammel, B & Chavarrı́a, E. 17536 (5); Hammel, B. 1363 (10), 1578 (10), 2149 (2), 2676 (9), 3016 (10), 4769 (9); Hammel, B. et al. 6840 (9); Hampshire, R. J. & Whitefoord, C. 125 (9), 133 (1), 203 (3), 214 (3); Hampshire, R. J. et al. 632 (9); Hancock, W. s.n. (678139) (10); Harmon, W. E. & Dwyer, J. D. 3384 (9); Hatch, W. R. & Wilson, C. L. s.n. (864375) (9); Hatheway, W. H. 1354 (10); Hawkes, J. G. et al. 2150 (9); Hawkins, T. 983 (9), 1119 (1), 1132 (2); Hayes, S. 84 (1), 86 (1), 750 (9); Heath, M. & Long, A. MA44 (9), MA53 (9); Henrich, J. E. & Stevens, W. 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