DIGESTIVE EFFICIENCY IN BOX TURTLES MILLER, M. R., AND K. J. REINECKE. 1984. Proper expression of metabolizable energy in avian energetics. Condor 86:396–400. RAUBENHEIMER, D. 1995. Problems with ratio analyses in nutritional studies. Functional Ecology 9:21–29. RAUBENHEIMER, D., AND S. J. SIMPSON. 1992. Analysis of covariance: an alternative to nutritional indices. Entomologia Experimentalis et Applicata 62:221– 231. REAGAN, D. P. 1974. Habitat selection in the ThreeToed Box Turtle, Terrapene carolina triunguis. Copeia 1974:512–527. SAMMARTANO, D. V. 1994. Spatial, Dietary, and Temporal Niche Parameters of Two Species of Box Turtle (Terrapene) in Microsympatry. Unpubl.
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master’s thesis, Southwest Missouri State University, Springfield. STUART, M. D., AND G. C. MILLER. 1987. The Eastern Box Turtle, Terrapene c. carolina (Testudines: Emydidae), in North Carolina. Brimleyana 13:123–131. THOMAS, R. B., D. MOLL, AND J. STEIERT. 1994. Evidence of a symbiotic relationship between cellulolytic bacteria and a freshwater herbivorous turtle. Southwestern Naturalist 39:386–388. ZIMMERMAN, L. C., AND C. R. TRACY. 1989. Interactions between the environment and ectothermy and herbivory in reptiles. Physiological Zoology 62:374–409. Accepted: 8 June 2006.
Journal of Herpetology, Vol. 40, No. 3, pp. 371–377, 2006 Copyright 2006 Society for the Study of Amphibians and Reptiles
New Species of Colostethus (Anura, Dendrobatidae) from the Penı´nsula de Paria, Venezuela CE´SAR L. BARRIO-AMORO´S,1 GILSON RIVAS,2
AND
HINRICH KAISER3,4
1 Fundacio´n Andı´genA, Apartado Postal 210, 5101-A Me´rida, Venezuela Museo de Historia Natural La Salle, Apartado Postal 1930, Caracas 1010-A, Venezuela 3 Department of Biology, Victor Valley College, 18422 Bear Valley Road, Victorville, California 92395, USA; E-mail:
[email protected] 2
ABSTRACT.—We describe a new species of Colostethus from the Penı´nsula de Paria in northeastern Venezuela. It inhabits the cloud forest of Cerro El Humo at an altitude of around 1000 m. By its morphology, the new species most closely resembles Colostethus bromelicola from the central Venezuelan Coastal Range. It can be confidently distinguished from other Colostethus by a suite of characters, including its dorsal pattern, absence of fringes on fingers, absence of a lingual process, and absence of yellow or orange chest coloration. This description increases to 33 the number of Colostethus species known from Venezuela. RESUMEN.—Se describe un nuevo Colostethus de la Penı´nsula de Paria en el noreste de Venezuela. La nueva especie habita´ en las vertientes del Cerro El Humo a` altitudes cerca de 1000 m. Morfolo´gicamente, este estrechamente relacionada con Colostethus bromelicola del tramo central de la Cordillera de la Costa venezolana. Se distingue claramente de las dema´s especies conocidas para el ge´nero de coloracio´n dorsal, ausencia de plieges laterales en los dedos de la mano, ausencia de proceso lingual, y coloracio´n ventral blanquecina. Con esta se eleva a` 33 el nu´mero de especies pertenecientes al ge´nero Colostethus para Venezuela.
Venezuela has an extremely diverse fauna of dendrobatid frogs, including 50 species in six genera described to date (modified from BarrioAmoro´s and Fuentes, 1999; Barrio-Amoro´s, 1998, 2004, 2006; Barrio-Amoro´s et al., 2004). Among the six represented dendrobatid taxa, the genera Colostethus and Mannophryne are the most diverse with 33 and 11 described and recognized species, respectively. Recent systematic research on these genera using molecular 4
Corresponding Author.
and chromosomal data (e.g., La Marca et al., 2002; Kaiser et al., 2003) is confirming that this great diversity has been underestimated and that some species with purportedly wide ranges are actually composed of several distinct, more localized taxa (e.g., Barrio-Amoro´s et al., 2006; Mannophryne trinitatis). A case in point are populations north of the Orinoco River known as the Colostethus brunneus species complex (La Marca, 1996). La Marca (1996) stated that these cis-Orinoco populations should be considered a distinct species restricted to the central Coastal Range, and the other to the Penı´nsula
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de Paria. Henceforth, we refer to the Coastal Range species of C. brunneus as C. brunneus complex sp. A. The Penı´nsula de Paria in northeastern Venezuela is the easternmost extension of the Andes on the South American continent (sensu Duellman, 1979, 1999) and, therefore, the easternmost portion of the Venezuelan Coastal Range. Although its southern half is flat and harbors a mixture of amphibian elements from the Llanos and Eastern Amazonia, the northern half is a narrow (approximately 10 km), mountainous ridge that runs east–west for over 100 km. The ridge is covered with lush lowland forest up to an elevation of about 600 m, above which the vegetation shifts to cloud forest habitat. During our herpetological exploration of the Penı´nsula de Paria, we have discovered several new species of amphibians and reptiles, including the recently described snake Taeniophallus nebularis (Schargel et al., 2005), a lizard of the genus Riama (Rivas et al., 2005), a series of as yet undescribed dendrobatid frogs, and additional, unnamed species of rain frogs (genus Eleutherodactylus). Herein, we describe a new species of Colostethus from the Penı´nsula de Paria. MATERIALS AND METHODS Specimens and Locality.—Specimens were collected during September 2001 on the slopes of Cerro El Humo, Penı´nsula de Paria, Estado Sucre, Venezuela (Appendix 1). Collections occurred in the late afternoon and at dusk. Specimens examined are housed in the following collections: Coleccio´n de Vertebrados, Universidad de los Andes, Me´rida (CVULA); Museo de la Estacio´n Biolo´gica de Rancho Grande, Maracay (EBRG); Coleccio´n de Herpetologı´a del Museo de Biologı´a de la Universidad Central de Venezuela, Caracas (MBUCV); and the Museo de Historia Natural La Salle, Caracas (MHNLS). Species Description.—The diagnosis follows Grant and Rodrı´guez (2001), whereas the description of the type series follows BarrioAmoro´s et al. (2004). Comparative data were taken from La Marca (1996), La Marca et al. (2002), Myers and Donnelly (2001), Rivero (1961), and Test (1956). Sex was determined by dissection and the maturity of our specimens was indicated by the presence of swollen oviducts. Measurements were taken to the nearest 0.1 mm using calipers, and they are defined and abbreviated as follows: (1) SVL, straight length from tip of snout to vent; (2) SL, shank length from outer edge of flexed knee to heel; (3) HeL, head length from tip of snout to posterior end of skull (posterior edge of prootic); (4) HW, head width between angle of
jaws; (5) InD, internarial distance, taken between centers of nares; (6) IOD, interorbital distance, measured as the shortest distance between the medial edges of the orbits across the skull; (7) EN, distance of anterior edge of eye to posterior edge of naris on the same side of the head; (8) ED, horizontal eye diameter; (9) TD, horizontal tympanum diameter, based on an estimated circular tympanum (the posterior portion of the tympanum is obscured by adductor musculature); (10) ETS, distance between the anterior edge of the eye to the tip of snout; (11) FD, disc width of Finger (F) III; (12) T4D, disc width of Toe (T) IV; (13) 1FL, length of FI from proximal edge of thenar tubercle to tip of disc; (14) 2FL, length of FII from the junction of FI and FIII to the tip of finger disc. Colostethus caribe n. sp. Figure 1 Holotype.—MHNLS 17462, an adult female collected by J. D. Trujillo on 6 September 2001 on the southern slope of Cerro El Humo, Penı´nsula de Paria, Estado Sucre, Venezuela (10u4190940N, 62u3791470W; Fig. 2), elevation 1050 m. Paratopotypes.—MHNLS 17463 and 17498, two adult females with same collection data as the holotype. Diagnosis.—A small Colostethus (up to 20.8 mm SVL); length of FI 5 FII; swollen condition of FIII in males unknown; rudimentary toe webbing; venter uniformly colored; sexual dimorphism unknown; dorsolateral stripe present, but faint; oblique lateral stripe absent; ventrolateral stripe absent; median lingual process absent; anal sheath absent; black arm bands absent; dorsum brown, sides dark brown, venter light orange laterally, transitioning to whitish medially. Species Comparisons.—Colostethus caribe is here compared to the only morphologically similar small-bodied Venezuelan species (Colostethus bromelicola, C. brunneus complex sp. A, Colostethus humilis, Colostethus undulatus). These species all lack an oblique lateral stripe and possess only rudimentary toe webbing. Colostethus caribe (whose characteristics appear in parentheses after those of compared species) can be distinguished from C. bromelicola by the well-defined whitish dorsolateral stripe (faint), black tips of digits (pointed with white), the black coloration in life of the flanks in lateral view (brown with small whitish spots), and by its arboreal, bromeliad-bound lifestyle (terrestrial). Colostethus brunneus complex sp. A has a strongly shagreened dorsum covered with tubercles (smooth), is larger in SVL (up to 24 mm, with only a maximum of 20.8 mm in C. caribe), has a snout rounded in dorsal and
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Fig. 1. Female of Colostethus caribe n. sp. (MHNLS 17498) in life.
FIG. 2. Distribution of Colostethus caribe n. sp. (black circle) and four similar species. Symbols used represent Colostethus bromelicola (white square), Colostethus brunneus species complex A (black squares), Colostethus humilis (white circles), and Colostethus undulatus (black cross). Shaded areas indicate topography, with darker shading representing higher altitudes.
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TABLE 1. Measurements given are for the type series of Colostethus caribe. Abbreviations are as explained in the text. Specimens are (1) female holotype (MHNLS 17462), (2) female paratype (MHNLS 17463), (3) female paratype (MHNLS 17498). No.
SVL
SL
HeL
HW
IOD
ED
TD
FD
T4D
ETS
1FL
2FL
1 2 3
20.8 19.5 19.6
8.9 9.0 7.8
7.9 7.8 6.9
7.0 6.8 6.8
2.5 2.5 2.5
2.7 2.3 2.5
1.2 1.0 1.0
0.8 0.6 0.6
0.9 0.7 0.8
3.2 3.0 3.0
2.9 2.9 2.8
2.7 2.9 2.9
ventral view (truncated). Colostethus humilis is a small frog with a maximum known SVL of 21.8 mm (La Marca et al., 2002). It possesses an indistinct tympanum (inferior half distinct), and a diamond-shaped dorsal pattern (no pattern). Colostethus undulatus has an undulating dorsal pattern (no pattern), no trace of any dorsolateral stripes (faint), and black flanks (brown with small white spots). Three other species of Colostethus have similarities to C. caribe but are not known to occur in Venezuela. Colostethus marchesianus (from its type locality in northern Brazil; see Caldwell et al., 2002) is a smaller species, with females reaching a SVL of up to 17 mm (20.8 mm), and with FI . FII (FI 5 FII). It possesses webbing between TII and TIII and between TIII and TIV (only rudimentary webbing to which webbing formula can be applied), has a wide and distinct dorsolateral stripe (faint) as well as a diffuse oblique lateral stripe (absent). Colostethus degranvillei is known from French Guyana and Surinam. This species has FI , FII (FI 5 FII), possesses a white posttympanic bar (absent), and its ventral surfaces are brown with white spots (orange tending to whitish). Colostethus beebei is known from Guyana, French Guyana, and Surinam. Its tympanum is indistinct (conspicuous), FI . FII (FI 5 FII), and it possesses a ventrolateral stripe (absent). Colostethus caribe can easily be distinguished from other Venezuelan and northern South American non-aposematically colored dendrobatids because it does not have a black collar nor a yellow throat (a defining feature of species in the genus Mannophryne; La Marca, 1992). Colostethus caribe also clearly has only rudimentary toe webbing, which eliminates from comparison those species with moderate to extensive toe webbing (Colostethus alboguttatus, Colostethus capurinensis, Colostethus dunni, Colostethus duranti, Colostethus haydeeae, Colostethus leopardalis, Colostethus mandelorum, Colostethus mayorgai, Colostethus meridensis, Colostethus molinarii, Colostethus orostoma, Colostethus saltuensis, Colostethus sanmartini, Colostethus serranus), all of which occur in biogeographically distinct and geographically distant regions (most from the Andes). Finally, C. caribe does not have a median lingual process, which eliminates from closer
comparison the Guianan Shield species of Colostethus (Colostethus ayarzaguenai, Colostethus breweri, Colostethus guayanaensis, Colostethus murisipanensis, Colostethus parkerae, Colostethus praderioi, Colostethus roraima, Colostethus shrevei, Colostethus tamacuarensis, Colostethus tepuyensis, Colostethus triunfo, Colostethus wothuja). Description of Type Series.—Complete sets of measurements for the type series are provided in Table 1. Body slender and elongate, quadrangular in vertical section. Dorsal skin smooth to shagreened (in MHNLS 17463), ventral skin smooth. Head longer than wide, greatest head width between angles of jaws 34% SVL (Table 1). Snout sloping, subacuminate in profile, nearly truncate in dorsal and ventral view. Nares situated near tip of snout and directed slightly posterolaterally; nares visible from the front, barely or not visible dorsally or ventrally. Canthus rostralis rounded; loreal region feebly concave (nearly flat), sloping slightly outward to lip. Interorbital region much wider than upper eyelid; snout longer than eye length. Tympanum conspicuous, large (nearly 50% ED), half concealed posterodorsally by a bulging adductor musculature; tympanum positioned closely behind eye and low, nearly touching angle of jaws. Teeth present on maxillary arch. Tongue round, triangular or quadrangular (shape differences may be artifacts of preservation); free posteriorly; median lingual process absent. Hand moderate in size (25% SVL). Relative lengths of adpressed fingers III . IV . II 5 I; FI of holotype just longer than II, equal in MHNLS 17463 and slightly shorter in MHNLS 17498; discs of all fingers expanded, round; FIII disc 1.5 times wider than distal end of adjacent phalanx. Base of palm with large, rounded palmar tubercle; elliptical thenar tubercle on base of FI; one or two subarticular tubercles (one each on FI and FII, two each on FIII and FIV); all tubercles flat and rounded; no supernumerary tubercles. Fringes on fingers absent. Hind limbs of moderate length, shank 43– 46% SVL. Relative lengths of adpressed toes IV . III . V . II . I; tip of TI reaching base of subarticular tubercle of TII. Toe discs moderately expanded, about 1.5 times wider than distal end of adjacent phalanx. Feet only basally webbed, webbing formula not applicable. No
NEW VENEZUELAN COLOSTETHUS fringes on toes. One to three nonprotuberant, small subarticular tubercles present (one on TI and TII, two on TIII and TV, three on TIV). Two metatarsal tubercles present, including a small conical outer, a slightly larger elliptical inner, and sometimes (e.g., MHNLS 17463) a median metatarsal tubercle; median metatarsal tubercle round, less distinct than others. Tarsal keel present on distal half of tarsus but not prominent; tarsal keel straight, reaching inner tarsal tubercle; a small ridge present at proximal end of tarsal keel. Anal opening at upper level of thighs; a short skin flap present above vent in MHNLS 17463, absent in other specimens. Color, Pattern, and Variation.—In preservative, dorsal color pale brown, without consistent pattern. MHNLS 17463 has a faint, dark brown interorbital blotch extending onto the upper eyelids, a poorly defined elongate middorsal mark, and a centered smaller spot on the sacrum. The holotype has a faint X-shaped pattern centered on the dorsum. It also has the interorbital blotch and some lighter colored irregular markings that may indicate dorsal scarring from past wounds or fungal infections. Forearms pale brown, with no pattern visible from above, but with a conspicuous longitudinal dark brown bar along the ventral surfaces of the forearms. There are also horizontal dark brown stripes crossing the anterior surfaces of the forearms. Hind limbs pale brown, crossed with one (MHNLS 17463) or two (MHNLS 17462, 17498) narrow dark brown lines, and in MHNLS 17498 with two symmetric, short white stripes on each side of the vent opening. Flanks dark brown, darker close to the border with the dorsum, well delimited from the dorsum by a faint, ill-defined dorsolateral line, made of fine whitish spots, reaching the tip of the snout above the upper eyelids; sides of the head covered with a mask emanating directly from the dark brown flanks, reaching the tip of snout where the tympanum is easily visible by its paler color. Narrow dark brown stripe along the upper lip and above and below the face mask with a white (dirty white in holotype) stripe; the darker superior border of the flanks continues posteriorly and meets on the upper part of the anal opening. Flanks covered with small whitish spots. No trace of an inguinal stripe. Ventrolateral stripe absent, but even without a stripe the flank pattern changes abruptly from brown to the orange ventrolateral coloration. Chin and lower jaw bordered by a broken line of brown melanophores, some widespread on the throat and the anterior part of the chest (holotype). In MHNLS 17498, only the chin has some melanophores. Apart of a few melanophores on the chin, MHNLS 17463 has a small dot formed by melanophores anterior and
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lateral to the chest. Palms and soles are dirty brown. Otherwise, the ventral color is an immaculate creamy white. Every tip of the digits, when viewed from above, has two ivory white spots on each disc, and sometimes even a third spot on the distal phalanx, just anterior to the discs. In life (from several photographs taken at the type locality) the dorsum is brown, with dark brown spots and cross marks; small white spots present on the dorsolateral stripes; flanks dark brown anteriorly and adjacent to the dorsolateral stripe; venterolaterally and posteriorly towards the groin this color fades; bluish spots present on flanks, the upper surfaces of hind limbs look tubercular. Ventral parts immaculate and whitish. Natural History.—Only three females were seen and caught when the species was first encountered. One animal sat at the entrance of a hole (perhaps an armadillo burrow) at the edge of a path in the cloud forest. The other two animals were observed initially in a dry streambed, along with young Craugastor biporcatus. No calls attributable to this species were heard. All animals moved rapidly when approached and proved hard to catch. Despite its superficial similarity with C. bromelicola, our observations suggests that this species is terrestrial, but no bromeliads were checked close to the type locality. During three additional brief visits to the type locality while conducting general surveys (August 2002, July 2003, December 2003) the species was not seen, although no focused searches for C. caribe were conducted. Etymology.—The species name caribe is a nominative used in apposition. The Spanish word caribe is normally used to describe the Carib native people who used to inhabit the coasts surrounding what is now known as the Caribbean Sea. Because their culture and voice have nearly disappeared, we name this frog, which inhabits some of Venezuela’s ancient caribe lands, for one of the ancestral people of the Caribbean. DISCUSSION The four species most closely resembling C. caribe live in geographically and ecologically disparate areas of Venezuela (Fig. 2). Colostethus humilis is known from the eastern slopes of the Cordillera de Me´rida in the Andes (the closest locality in Trujillo state lies 850 km westsouthwest of the Penı´nsula de Paria). Colostethus bromelicola is known only from its type locality, Rancho Grande in Henri Pittier National Park (545 km west). Colostethus undulatus is known only from Yutaje´ tepui (700 km southwest) in the Guiana Shield south of the Orinoco River, and C. brunneus complex sp. A is only known
´ S ET AL. C. L. BARRIO-AMORO
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from scattered localities along the central Coastal Range, including some in Henri Pittier National Park and in the Sierra de San Luis, Falco´n state. Morphologically, C. caribe most closely resembles C. bromelicola. Thus, C. caribe may have its origins in an allopatric speciation event as mountain formation separated portions of the Coastal Range from the Andes and the Penı´nsula de Paria. Such a speciation event can explain the difference in microhabitat use, active in the forest litter for C. caribe and inhabiting bromeliads 2–5 m above-ground for C. bromelicola (Dixon and Rivero-Blanco, 1985). Although the Andes are not closely connected with the Penı´nsula de Paria by their Colostethus fauna, with no species known to occur in both mountain chains, the Coastal Range and, to some degree, the Venezuelan Guayana are indeed botanically connected (Steyermark, 1979). A connection between these mountains involving an ancestral Colostethus is, therefore, one scenario with biogeographic merit. Acknowledgments.—Fieldwork in the Penı´nsula de Paria was supported by a grant from the College of Arts and Sciences at La Sierra University to HK. We are indebted to J. D. Trujillo, W. E. Schargel, E. Bonaccorso, and J. M. Guyasamı´n for companionship in the field during our different expeditions to the Penı´nsula de Paria. We gratefully acknowledge the funding provided by the Small Grants Program of the Panorama Society, Natural History Museum and Biodiversity Research Center, University of Kansas to J. M. Guayasamı´n and by a grant from the Phi Sigma Biological Honor Society to W. E. Schargel. Collection permits (01-11-0510 and 01-11-0590) were issued to GR by the Ministerio del Ambiente y de los Recursos Naturales through the Fundacio´n La Salle de Ciencias Naturales. LITERATURE CITED BARRIO-AMORO´S, C. L. 1998. Sistema´tica y biogeografı´a de los anfibios (Amphibia) de Venezuela. Acta Biolo´gica Venezuelica 18:1–93. ———. 2004. Amphibians of Venezuela, systematic list, distribution and references; an update. Review of Ecology in Latin America 9:1–48. ———. 2006. A new dendrobatid frog (Anura: Dendrobatidae: Colostethus) from Aprada tepui, southern Venezuela. Zootaxa 1110:59–68. BARRIO-AMORO´S, C. L., AND O. FUENTES. 1999. Sinopsis de la familia Dendrobatidae (Amphibia: Anura) de Venezuela. Acta Biolo´gica Venezuelica 19:1–10. BARRIO-AMORO´S, C. L., O. FUENTES, AND G. RIVAS. 2004. Two new species of Colostethus (Anura: Dendrobatidae) from the Venezuelan Guayana. Salamandra 40:183–200.
BARRIO-AMORO´S, C. L., G. RIVAS, C. R. MOLINAS, AND H. KAISER. 2006. Mannophryne trinitatis (Anura: Dendrobatidae) is a Trinidadian single-island endemic. Herpetological Review in press. CALDWELL, J. P., A. P. LIMA, AND C. KELLER. 2002. Redescription of Colostethus marchesianus (Melin, 1941) from its type locality. Copeia 2002:157–165. DIXON, J. R., AND C. RIVERO-BLANCO. 1985. A new dendrobatid frog (Colostethus) from Venezuela, with notes on its natural history and that of related species. Journal of Herpetology 19:177–184. DUELLMAN, W. E. 1979. The South American herpetofauna: a panoramic view. In W. E. Duellman (ed.), The South American Herpetofauna: Its Origin, Evolution and Dispersal, pp. 1–28. Museum of Natural History, University of Kansas, Monograph 7:1–485. ———. 1999. Distribution patterns of amphibians in South America. In W. E. Duellman (ed.), Patterns of Distribution of Amphibians, pp. 255–328. Johns Hopkins University Press, Baltimore, MD. GRANT, T., AND L. O. RODRIGUEZ. 2001. Two new species of frogs of the genus Colostethus (Dendrobatidae) from Peru´ and a redescription of C. trilineatus (Boulenger, 1883). American Museum Novitates 3355:1–24. KAISER, H., C. STEINLEIN, W. FEICHTINGER, AND M. SCHMID. 2003. Chromosome banding of six dendrobatid frogs (Colostethus, Mannophryne). Herpetologica 59:203–218. LA MARCA, E. 1992. Cata´logo taxono´mico, biogeogra´fico y bibliogra´fico de las ranas de Venezuela. Cuadernos Geogra´ficos, Univesidad de Los Andes, Me´rida 9:1–197. ———. 1996. Ranas del ge´nero Colostethus (Amphibia: Anura: Dendrobatidae) de la Guayana Venezolana, con la descripcio´n de siete nuevas especies. Publicaciones de la Asociacion Amigos Don˜ana 9:1–64. LA MARCA, E., M. VENCES, AND S. LO¨TTERS. 2002. Rediscovery and mitochondrial relationships of the dendrobatid frog Colostethus humilis suggest parallel colonization of the Venezuelan Andes by poison frogs. Studies of the Neotropical Fauna and Environment 37:233–240. MYERS, C. W., AND M. DONNELLY. 2001. Herpetofauna of the Yutaje´-Corocoro Massif, Venezuela: second report from the Robert G. Goelet American Museum Terramar Expedition to the northwestern tepuis. Bulletin of the American Museum of Natural History 261:1–85. RIVAS, G., W. E. SCHARGEL, AND J. MEIK. 2005. A new species of Riama (Squamata: Gymnopthalmidae), endemic to the Penı´nsula de Paria, Venezuela. Herpetologica 61:461–468. RIVERO, J. A. 1961. Salientia of Venezuela. Bulletin of the Museum of Comparative Zoology 126:1– 267. SCHARGEL, W. E., G. RIVAS, AND C. W. MYERS. 2005. An enigmatic new snake from cloud forests of the Penı´nsula de Paria, Venezuela (Colubridae: genus Taeniophallus). American Museum Novitates 3484:1–22. STEYERMARK, J. A. 1979. Plant refuge and dispersal centres in Venezuela: their relict and endemic elements. In K. Larsen and L. B. Holm-Nielsen
NEW VENEZUELAN COLOSTETHUS (eds.), Tropical Botany, pp. 185–221. Academic Press, London. TEST, F. H. 1956. Two new dendrobatid frogs from Northern Venezuela. Occasional Papers of the Museum of Zoology, University of Michigan 577:1–9. Accepted: 10 June 2006.
APPENDIX 1 Material Examined Colostethus bromelicola.—MBUCV 5123, from Rancho Grande, Estado Aragua, elevation 1100 m. (On the
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specimen label the location is given as lying in Miranda state at an altitude of 200 m, both of which constitute bookkeeping errors.) Colostethus brunneus complex sp. A.—MBUCV 1996, from Rancho Grande, Estado Aragua, Venezuela (as determined by F. H. Test). EBRG 3879, Sierra de San Luis, Cerro Galicia, Estado Falco´n, elevation 1300 m. EBRG 3946–47, Cerro Paraguaniba, Sierra de San Luis, Estado Falco´n. Colostethus humilis.—CVULA 3790, from the Doradas-Uribante Rivers region (no more detailed locality given), Estado Tachira, Venezuela. Colostethus undulatus.—EBRG 3040–42 (paratypes), from Cerro Yutaje´, elevation 1750 m, Estado Amazonas, Venezuela.
