Sibon perissostichon sp. nov. pan köhler et al. 2010
Herpetologica, 66(1), 2010, 80–85E 2010 by The Herpetologists’ League, Inc. A NEW SPECIES OF SIBON (SQUAMATA: COLUBRIDAE) FROM WESTERN PANAMA GUNTHER KOHLER1,3, SEBASTIAN LOTZKAT1,2, ¨ AND ANDREAS HERTZ1,2 1 Senckenberg Forschungsinstitut und Naturmuseum, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany 2 Johann Wolfgang Goethe-University, Institute for Ecology, Evolution and Diversity, BioCampus—Westend, Siesmayerstr. 70, 60323 Frankfurt am Main, Germany ´ ABSTRACT: We describe a new species of Sibon from the Reserva Forestal La Fortuna, Chiriquı Province, Panama. The new species is distinguished from all species presently assigned to Sibon by having 17 rows of dorsal scales at midbody and 15 rows one head length anterior to cloaca (vs. 13 or 15 rows of dorsal scales at midbody without reduction on posterior portion of body in all other species of Sibon), and dorsal scale rows that change orientation from the sides of body toward middorsum (vs. maintaining more or less the same orientation). A key to the Lower Central American species of Sibon is provided. RESUMEN: Describimos una nueva especie de Sibon procedente de la Reserva Forestal La Fortuna, ´ ´ Provincia de Chiriquı, Panama. La nueva especie se diferencia de todas las especies actualmente asignadas al ´ genero Sibon por tener 17 hileras de escamas dorsales en la mitad del cuerpo y 15 hileras una longitud de ´ ´ cabeza anterior a la cloaca (13 o 15 hileras de escamas dorsales en la mitad del cuerpo sin reduccion en la ´ ´ parte posterior del cuerpo en todas las demas especies de Sibon), ası como hileras de escamas dorsales que ´ cambian su orientacion desde los flancos del cuerpo hacia el centro del dorso (conservan su orientacion ´ ´ longitudinal en todas las demas especies de Sibon). Presentamos una clave para las especies de Sibon que ´ ocurren en el sur de Centroamerica. Key words: Colubridae; New species; Panama; Sibon; Squamata THE GENUS Sibon is a moderately diverse took additional data for the Lower Centralgroup of small to moderate-sized snakes that are American species of Sibon from McCraniedistributed from Nayarit and Veracruz, Mexico, (2006), Savage and McDiarmid (1992), andthroughout Central America, and well into ´ Solorzano (2002). Abbreviations for museumSouth America to Ecuador and Brazil. Recently, collections follow those of Leviton et al. (1985) ´McCranie (2006, 2007) and Solorzano (2002) ´ except for CRB (Coleccion de la Reservadescribed new species of Sibon, bringing the ´ Bartola, Rıo San Juan, Nicaragua). Specimensnumber of Central American species in this in the Appendix labeled with SL fieldgenus to 11 (Ko ¨hler, 2008). Three fieldwork numbers will be deposited in the Universidadseasons in Panama (January 2006, January and ´ ´ ´ Autonoma de Chiriquı, Davıd, Panama, andFebruary 2007, and May to August 2008) those labeled with JS field numbers will beproduced several specimens of Sibon annulatus deposited in the Museo de Ciencias Naturalesand S. nebulatus. Among the specimens collect- de la Universidad Centroamericana (UCA),ed was a single adult female of Sibon that differs Managua, Nicaragua. We follow Dowlingstrikingly from all its congeners in the number (1951) for the method of counting ventralsand arrangement of dorsal scale rows. Compar- and Savage (1973) for the terminology ofisons with the known species of Sibon from scales in the loreal region of the head. The sexCentral America demonstrated that this speci- of the holotype was determined by visualmen represents an undescribed species. examination of relative tail length and basal girth. Head and scale measurements were MATERIALS AND METHODS made to the nearest 0.1 mm with dial calipers The format for the description of the new held under a dissecting microscope. Snout–species generally follows that of McCranie vent length and tail length measurements(2007). We provide a list of the specimens were taken to the nearest millimeter alongsideexamined for comparison in the Appendix. We a meter stick. Terminology for measurements is abbreviated as: snout–vent length, SVL; tail 3 CORRESPONDENCE: e-mail, email@example.com length, TL; head length, HL; snout length, 80
March 2010] HERPETOLOGICA 81 FIG. 1.—Holotype of Sibon perissostichon (SMF 88716)in life, SVL 458.0 mm.SL; and head width, HW. Eye length equalsthe horizontal distance across eye. Scaledimensions were measured at the longest orwidest points along the longitudinal or per-pendicular axis of the body, respectively. Thecapitalized colors and color codes (the latter inparentheses) are those of Smithe (1975–1981).Drawings of the head were made using acamera lucida attachment for a dissecting FIG. 2.—Head of holotype of Sibon perissostichonmicroscope. A small tissue sample of For- (SMF 88716). Scale bars 5 2.0 mm.schungsinstitut Senckenberg (SMF) 88716was taken from the edge of a ventral scale rows one head length anterior to cloaca (vs. 13about midbody and deposited in the tissue or 15 rows of dorsal scales at midbody withoutsample collection of the Senckenberg For- reduction on posterior portion of body in allschungsinstitut und Naturmuseum, Frankfurt, other species of Sibon), and dorsal scale rowsGermany. that change orientation from the sides of body toward middorsum (vs. maintaining more or SPECIES ACCOUNT less the same orientation). Additionally, S. perissostichon differs from its congeners by Sibon perissostichon sp. nov. the following characteristics: subcaudals 108 (Figs. 1–4) in S. perissostichon vs. 113–126 in female S. Holotype.—SMF 88716, adult female as annulatus. Light interspaces between darkjudged by the shape of the base of the tail, bands on body Sayal Brown (223C) in S.collected near Lost and Found Ecohostel, perissostichon vs. interspaces orange to red-Reserva Forestal La Fortuna, 8u 40.479 N, 82u orange middorsally, grading to cream laterally12.979 W, 1434 m elevation, by Sebastian in S. anthracops. Enlarged penultimate su-Lotzkat on 14 May 2008. Original field pralabial not in contact with eye in S.number SL 145. perissostichon vs. this scale in contact with Diagnosis.—Sibon perissostichon can be eye in S. argus and S. longifrenis. Also, darkdistinguished from its congeners by having dorsal bands or spots reach well onto venter or17 rows of dorsal scales at midbody and 15 even across venter in S. perissostichon vs. dark
82 HERPETOLOGICA [Vol. 66, No. 1 FIG. 4.—Scalation (middorsal portion of body) of (a) Sibon perissostichon (SMF 88716) and (b) S. annulatus FIG. 3.—Holotype of Sibon perissostichon (SMF 88716) (SMF 88715). Scale bars 5 2.0 mm.at its capture site. no dark dorsal bands and venter immaculatedorsal bands or spots reach at most to outer in S. manzanaresi and S. sanniolus. Ventralsedges of ventrals, although dark shading may 186, subcaudals 108, bands solid dark brown,be present on venter in S. argus and S. darker than interspaces in S. perissostichon vs.longifrenis. One anterior temporal present in ventrals 168–171 in females, subcaudals 95–S. perissostichon vs. no such scale in S. carri. 100 in females, bands dark brown ventrally,Therefore, penultimate supralabial not in their centers paler brown, paler than inter-contact with parietal in S. perissostichon vs. spaces in S. miskitus. Postmental present;these scales in contact in S. carri. Bands solid therefore, first infralabials not in contact withdark brown, darker than interspaces in S. each other, vertebral row not enlarged in S.perissostichon vs. bands dark brown ventrally, perissostichon vs. first infralabials in contacttheir centers paler brown, paler than inter- with each other, no postmental, vertebral rowspaces in S. dimidiatus. Dorsal surface of head moderately enlarged in S. nebulatus.with black markings, ground color of body Description of the holotype.—Adult femalebrown, postmental single in S. perissostichon (Figs. 1, 2); SVL 458 mm; TL 210 mm; HLvs. dorsal surface of head with green or yellow 15.0 mm; HW 9.0 mm; SL 4.2 mm. Head veryas well as red and black markings, ground distinct from neck; snout short, blunt in dorsalcolor of body green, and postmental divided in and lateral outline; rostral wider (2.6 mm) thanS. lamari. Ventrals 186, loreal in contact with high (2.0 mm), not extending posteriorlyeye in S. perissostichon vs. ventrals 155, loreal between internasals, its length visible fromseparated from eye in only known female above one-quarter length of median internasalspecimen of S. linearis. Dark dorsal bands suture (1.2 mm); internasals shorter (1.5 mm)present, reaching well onto venter or even than wide (2.0 mm), two-thirds length ofacross venter in S. perissostichon vs. usually prefrontal suture (2.6 mm); prefrontals longer
March 2010] HERPETOLOGICA 83(2.6 mm) than wide (2.2 mm), their median Brown (223B) with Jet Black (89) blotches;suture two-thirds length of frontal; prefrontals ventral surface of head dirty white with Rawbordering eye above loreal; frontal longer Umber (223) mottling and Jet Black (89) chin(4.1 mm) than wide (3.5 mm), widest anteri- shields and first ventral scale; iris Mars Brownorly, length two-thirds that of parietal, in (223A). Color in preservative (70% ethanol)contact with prefrontals, supraoculars, and similar to color in life.parietals; supraoculars longer (3.6 mm) than Etymology.—The name perissostichon is awide (1.6 mm), nearly as long as frontal; noun in apposition and is derived from theparietals longer (5.9 mm) than wide (3.8 mm), Greek words perissos (beyond the regularwidest anteriorly, their median suture (4.4 mm) number or size) and stichos (row, line),slightly longer than frontal; parietals in contact referring to the high number of dorsal scalewith frontal, supraoculars, upper postoculars, rows in this species.anterior and posterior temporals, and eight Natural history notes.—The holotype wasnuchal scales; nasal divided, in contact with encountered at 2053 h in secondary Premon-first two supralabials, loreal, prefrontal, inter- tane Forest while it was active within thenasal, and rostral; loreal single, longer (1.6 mm) dense, fog-moistened undergrowth about 0.5 mthan high (0.7 mm), its length less than one- above the forest floor (Fig. 3). At the time ofhalf length of snout (tip of snout to anterior encounter, the snake was moving about on theedge of eye) and one-half length of eye inclined trunk of a young tree covered with(3.3 mm); preoculars absent; postoculars three foliose hepatics. The undergrowth surround-(right) and two (left); temporals one plus three, ing this trunk consisted of herbaceous as wellanterior one above fifth supralabial, lowest as young woody plants, among which we couldposterior one above sixth supralabial; suprala- distinguish different ferns as well as membersbials six, with third and fourth (right) or fourth of the families Rubiaceae and Melastomata-(left) bordering eye (enlarged penultimate ceae. The type locality is situated about 20supralabial separated from eye by lower altitudinal meters below a narrow ridge on apostocular); pupil vertically elliptical; eye very steep slope (approximately 45u inclina-length two times length of loreal; infralabials tion) facing the southern Pacific lowlands.seven, first pair separated posterior to mental Most trees’ diameters at breast height do notby a single, triangular postmental, first infra- exceed 50 cm. The dense canopy layer extendslabial in contact with postmental, infralabials from about 10 m aboveground upward.2–5 in contact with anterior chin shields; Detached rocks are scattered among theanterior chin shields paired, longer (5.6 mm) scenery, and there are no streams nearby.than wide (1.6 mm), longer than posterior chin Air parameter measurements conducted aftershields; posterior chin shields longer (3.4 mm) the capture showed a temperature of 17.7 uCthan wide (2.5 mm), paired, in broad contact and 87% relative humidity. In the vicinity ofmedially; mental groove present; ventrals 186; the type locality of S. perissostichon, wecloacal scute single; subcaudals 108, divided recorded the following species of amphibiansexcept for third and fourth pairs that are fused; and reptiles: Craugastor podiciferus, Dia-ventrals plus subcaudals 294; dorsal scales sporus diastema, Pristimantis cruentus, P.smooth, in 15–17–15 rows (anterior–mid- ridens, Anolis capito, A. fortunensis, A.body–hindbody), without apical pits or supra- humilis, Ptychoglossus plicatus, Dendrophi-cloacal ridges; dorsal scale rows changing dion paucicarinatum, Geophis sp., Imantodesorientation from the sides of body toward cenchoa, Liophis epinephelus, S. annulatus,middorsum; vertebral row not enlarged. Trimetopon slevini, and Urotheca sp. Color in life.—Dorsal ground color SayalBrown (223C) with Sepia (119) blotches DISCUSSIONreaching onto venter, mostly continuous Aside from the new species describedacross venter, some ventrally offset; ventral herein, four species of Sibon are known tosurface between dark blotches dirty white, ¨ occur in western Panama (Kohler, 2008): S.suffused with Buff (124) and mottled with annulatus, S. argus, S. longifrenis, and S.Sepia (119); dorsal surface of head Verona nebulatus. During our fieldwork, we have
84 HERPETOLOGICA [Vol. 66, No. 1collected eight specimens of S. annulatus 3(b). Enlarged penultimate supralabial not in(including a specimen collected the same contact with eye; dark dorsal bands or spots reach well onto venter or evennight as the holotype of S. perissostichon just across venter 5 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _260 m WSW of the type locality; see SMF 4(a). 181–201 ventrals; 112–121 subcau-numbers in Appendix). All eight specimens of dals Sibon argus _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _S. annulatus have a single postmental scale, 4(b). 151–173 ventrals; 82–106 subcau- dals Sibon longifreniswhereas this species is said to have usually a _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 5(a). First infralabials in contact with eachpair of postmentals; the presence of a single other behind the mental Sibon nebulatus _ _ _ _ _ _ _ _postmental is considered typical for S. dimi- 5(b). First infralabials separated from each ¨diatus (Kohler, 2008; Savage, 2002). However, other by one or two postmentals behindon the basis of characteristics of dorsal the mental 6 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _coloration, we refer these specimens to S. 6(a). Bands dark brown ventrally, their cen- ters paler brown, paler than interspaces;annulatus as opposed to S. dimidiatus. They usually one postmental present _ _ _ _ _ _ _ _ _ _ _ _all have more or less solid dark brown bands Sibon dimidiatus _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _without paler centers, reaching well onto 6(b). Bands dark brown ventrally and lateral-venter. Our Panamanian specimens of S. ly, their centers not paler brown to cream laterally; lateral portion of bandsannulatus have 184–193 ventrals and 103– darker than interspaces; one or two118 subcaudals in males (n 5 2), and 170–185 postmentals present 7 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ventrals and 118–130 subcaudals in females (n 7(a). Dorsal surface of head with green or5 4). Two juveniles (SMF 88713–14) have yellow as well as red and black markings;177–184 ventrals and 108–114 subcaudals. ground color of body green Sibon lamari _ _ _ _ _ _ _ _ 7(b). Dorsal surface of head with pale andThus, our specimens show slightly greater dark brown markings; ground color ofvariation in the number of ventral scales than body usually olive to pale brown _ _ _ _ _ _ _ _ _ _has been documented for this species so far Sibon annulatus _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _(McCranie, 2007). Within the Dipsadini(sensu Cadle, 2007), S. perissostichon is Acknowledgments.—Collecting and exportation permitsunique in having 17 dorsal scale rows at were provided by A. Salazar, Y. Hidalgo, and J. Garcıa, ´midbody and a reduction to 15 on the Autoridad Nacional del Ambiente, Panama City, Panama.posterior portion of body. Reduction of the Q. D. Fuenmayora and V. Martinez, Panama City, Panama, provided valuable assistance with acquisition ofnumber of dorsal rows on the posterior body these permits. For field assistance, we thank L. Stadleris unusual in the genus Sibon. Only S. and N. Hamad, as well as A. Bennett, who also providednebulatus is known to occasionally have logistical support. For the loan of or access to specimens,‘‘reductions from 15 to 13’’ (Peters, 1960:174). we thank L. Ford, C. J. Raxworthy, D. R. Frost, American Museum of Natural History (AMNH), New York; C. J. McCarthy, The Natural History Museum (BMNH), KEY TO THE SPECIES OF SIBON FROM LOWER London; W. E. Duellman and J. E. Simmons, University CENTRAL AMERICA of Kansas, Natural History Museum (KU), Lawrence; R. W. McDiarmid and W. R. Heyer, National Museum of Natural History (USNM), Washington, DC. This paper is 1(a). Seventeen rows of dorsal scales at mid- body and 15 rows one head length based on work funded to AH by the FAZIT-STIFTUNG, anterior to cloaca; dorsal scale rows and to SL by the Studienstiftung des deutschen Volkes. changing orientation from the sides of An earlier version of this manuscript was read by J. R. body toward middorsum (Fig. 4a) _ _ _ _ _ _ _ _ McCranie, L. D. Wilson, and an anonymous reviewer who Sibon perissostichon _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ all helped to improve the paper with their comments and corrections. 1(b). Thirteen or 15 rows of dorsal scales at midbody without reduction on posterior portion of body; dorsal scale rows oriented longitudinally (Fig. 4b) 2 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ LITERATURE CITED 2(a). Thirteen dorsal rows at midbody _ _ _ _ _ _ _ _ _ CADLE, J. E. 2007. The snake genus Sibynomorphus Sibon anthracops _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ (Colubridae: Dipsadinae: Dipsadini) in Peru and 2(b). Fifteen dorsal rows at midbody 3 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ Ecuador, with comments on the systematics of the 3(a). Enlarged penultimate supralabial in Dipsadini. Bulletin of the Museum of Comparative contact with eye; dark dorsal bands or Zoology 158:183–283. spots reach at most to outer edges of DOWLING, H. G. 1951. A proposed standard system of ventrals, although dark shading may be counting ventrals in snakes. British Journal of Herpe- present on venter 4 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ tology 1:97–99.
March 2010] HERPETOLOGICA 85KOHLER, G. 2008. Reptiles of Central America. Herpeton, ¨ 13.129 W, 1320 m: SMF 88715; Pianista trail, 8u 529 17.10 Offenbach, Germany. N, 82u 249 57.40 W, 1005 m: SMF 86411; Pianista trail, CasaLEVITON, A. E., R. H. GIBBS, JR., E. HEAL, AND C. E. ´ de Ancon: SMF 88713–14; Veraguas: Cerro Mariposa, DAWSON. 1985. Standards in herpetology and ichthyol- 680 m: SMF 89596; Cerro Mariposa, 1000 m: SMF 89597. ogy: Part I. Standard symbolic codes for institutional Sibon anthracops.—COSTA RICA: Guanacaste: Volcan resource collections in herpetology and ichthyology. Rincon de la Vieja: SMF 77133; EL SALVADOR: No Copeia 1985:802–832. additional locality data: SMF 68197; La Libertad: SantaMCCRANIE, J. R. 2006. New species of Sibon (Squamta: Tecla, Pinares de Suiza: SMF 81286; San Salvador: San Colubridae) from northeastern Honduras. Journal of Salvador: SMF 79239; GUATEMALA: Zacapa: San Vi- Herpetology 40:16–21. cente, Aldea Santa Elena: SMF 82594; NICARAGUA:MCCRANIE, J. R. 2007. A second new species of Sibon ´ Carazo: Diriamba: KU 174287; Estelı: 15 km E Estelı, ´ (Squamta: Colubridae) from La Mosquitia, northeast- 800 m: KU 116975; Granada: Volcan Mombacho: SMF ern Honduras. Herpetologica 63:213–218. 81024; Managua: 1 mi N Sabana Grande: KU 42299; 6 miPETERS, J. A. 1960. The snakes of the subfamily Dipsadinae. WSW Sabana Grande: KU 42300; 8 km on South road: KU Miscellaneous Publications Museum of Zoology Univer- 174286; Casa Colorada, Las Nubes: KU 174282–85; Rivas: sity of Michigan 114:1–224. Morgan’s Rock, 11u 189 28.90 N, 85u 549 59.10 W, 35–265 m:SAVAGE, J. M. 1973. A revised terminology for plates in the JS field number 806, 838, SMF 88971–72. Sibon loreal region of snakes. British Journal of Herpetology dimidiatus.—HONDURAS: Copa ´n: Near Quebrada 5:360–362. Grande, 1300 m: SMF 79119; NICARAGUA: Matagalpa:SAVAGE, J. M. 2002. The Amphibians and Reptiles of Costa Selva Negra, 12u 59.969 N, 85u 54.559 W: SMF 77800, Rica. A Herpetofauna between Two Continents, 77860, 79027–28; Matagalpa: BMNH 1822.214.171.124. Sibon between Two Seas. University of Chicago Press, longifrenis.—COSTA RICA: Heredia: Puerto Viejo de Chicago, Illinois, USA. ´ Sarapiquı, 50 m: SMF 79966; HONDURAS: Gracias aSAVAGE, J. M., AND R. W. MCDIARMID. 1992. Rediscovery ´ ´ ´ Dios: Reserva de Biosfera Rıo Platano: SMF: 86012; of the Central American colubrid snake, Sibon argus, NICARAGUA: Jinotega: Bosawas, Kulum Kitang, 14u with comments on related species from the region. 19.89 N, 84u 56.29 W, 180 m: SMF 88182. Sibon Copeia 1992:421–432. nebulatus.—COSTA RICA: Guanacaste: Volcan Arenal:SMITHE, F. B. 1975–1981. Naturalist’s Color Guide. SMF 77134; EL SALVADOR: Ahuachapa ´n: Parque Part I. Color Guide. 182 Color Swatches. American Nacional El Imposible, La Fincona, 720 m: SMF 81283; Museum of Natural History, New York, New York, USA. ´ HONDURAS: Atlantida: La Ceiba: SMF 79879; NICARA-SOLORZANO, A. 2002 (2001). Una nueva especie de ´ ´ GUA: Atla ´ntico Norte: Wanı, Siuna: UCA 214; Big Falls, ´ serpiente del genero Sibon (Serpentes: Colubridae) ´ Rıo Pis Pis: AMNH 12682; Rancho Alegre, 13u 399 47.50 N, de la vertiente del Caribe de Costa Rica. Revista de ´ Biologıa Tropical 49:1111–1120. 85u 019 38.90 W, 315 m: SMF 85510; Bosawas, Muru Lak, 14u 21.219 N, 84u 56.529 W, 185 m: SMF 87225; Parque Nacional Saslaya, Cerro El Toro, 830 m: SMF 83122; .Accepted: 18 November 2009 ´ Atlantico Sur: Kukra: AMNH 12680–81; betw. Kukra and .Associate Editor: Christopher Raxworthy ´ Kanawa: AMNH 12679; Isla del Maız Grande: AMNH ´ 97068–70; Isla del Maız Grande, Mt. Pleasant: KU 174303; Boaco: near Boaco: KU 174302; Granada: Volcan Mom- ´ bacho, near lower antenna, 11u 49.999 N, 85u 58.779 W, APPENDIX I 1100 m: SMF 79561, 79835; Managua: Casa Colorada, El Crucero: KU 174295–99; Casa Colorada, Las Nubes: KU Comparative Material Examined 174288–994, 174300; Nueva Segovia: 5 km N, 2.5 km E ´ Sibon annulatus.—HONDURAS: Colon: Reserva de ´ Jalapa, 680 m: KU 112982; Rıo San Juan: Los Guatuzos, ´ ´ ´Biosfera Rıo Platano, El Ocotillal, Cabeceras de Rıo ´ ´ ´ Rıo Frıo, Fundeverde, 11u 049 37.00 N, 84u 449 55.70 W, ´Platano, 370 m: SMF 86014; Gracias a Dios: Reserva de ´ 45 m: SMF 87266; Los Guatuzos, along Rıo Papaturro, ´ ´ ´Biosfera Rıo Platano, Pomokir, 150 m: SMF 86013; 11.0227u N, 85.0513u W, 40 m: JS field number 995; nearNICARAGUA: Atla ´ntico Norte: Parque Nacional Saslaya, ´ El Castillo, finca of JOHN KINLACH: KU 174301; Rıo SanCampamento Los Monos, 13u 45.19 N; 85u 02.29 W, 850 m: Juan, Boca de San Carlos, 10u 47.269 N, 84u 11.709 W, 20 m:SMF 79655; Jinotega: Reserva Biosfera Bosawas, ca. 0.5 km ´ SMF 79836; Rıo San Juan, Bartola, 10u 58.379 N, 84u 20.359SE Pueblo Wiso, 13u 59.609 N, 85u 19.609 W, 200 m: SMF W, 30 m: CRB 011–13, SMF 79837; Camp Machado: ´ ´ ´78369; Matagalpa: Cerro Musun, Palan, Bilampı, 13u 009 USNM 19558; San Juan del Norte: USNM 5571, 62997;41.00 N, 85u 149 11.60 W, 620 m: SMF 88180; PANAMA: ´ PANAMA: Chiriquı: Pianista trail, Casa de Calisto: SMFBocas del Toro: Palo Seco, 8u 469 39.50 N, 82u 129 330 W, 88717; Los Algarrobos: Casa Culebra, 134 m: SL field ´1030 m: SMF 85077–78; Chiriquı: near Lost and Found number 165; Volante, 480 m: SMF 89599; Veraguas: CerroEcohostel, Reserva Forestal La Fortuna, 8u 40.439 N, 82u Mariposa, 933 m: SMF 89598.