Sunday, April 3, 2016

House Gecko, Hemidactylus mabouia (Family Gekkonidae)



Gekko mabouia Moreau De Jonnès 1818: 138
Hemidactylus mabouia - Duméril & Bibron 1836: 362

Type locality: Restricted to St Vincent in the Lesser Antilles (Stejneger, 1904), but the type specimen apparently does not belong to the taxon to which this name is usually applied. Instead it is a H. angulatus, which is similar to those from the Greater Antilles and Colombia, particularly the latter (Kluge, 1969) [from Carranza & Arnold 2006].

Adults have a SVL up to 63 mm; hatchlings are about 20 mm SVL; tails are about 1.2x the body length. Diagnosis. Dorsum covered with small granular scales studded with trihedral; tubercles in 6–14 longitudinal rows. Ventral scales overlap and are large than dorsals. Upper labials 8–12, lower labials 7–9. Lamellae on fourth finger 7–9, lamellae on fourth toe 7–8. Total pores in males 29–33. Color change in this lizard can be dramatic. Similar species. It is most easily confused with Hemidactylus palaichthus which has a more spiny appearance and more lamellae on the base of toes four and five that extend to the base of the digit. Distribution. Probably occurs island wide and associated with human modified habitats, most often buildings and frequently near lights that attract insects. Other common names. House gecko, wall mabuia, African woodslave, twenty-four hours.

Duméril, A.M. C. and G. Bibron. 1836. Erpetologie Générale ou Histoire Naturelle Complete des Reptiles. Vol.3. Libr. Encyclopédique Roret, Paris, 528 pp.

Kluge, A. G. 1969. The evolution and geographical origin of the New World Hemidactylus mabouia-brooki complex (Gekkonidae, Sauria). Misc. PubI. Mus. Zool. Univ. Michigan 138:1-78.

Moreau de Jonnès, A. 1818. Monographie du mabouja des murailles, on Gecko Mabouja des Antilles. Bull. Scient. Soc. Philomath. Paris, Ser. 3,5:138-139.


Stejneger, L. 1904. The herpetology of Porto Rico. Rept. United States Natl. Mus. 1902: 549-724.

Trinidad Anole, Anolis trinitatis (Family Dactyloidae)

Anolis trinitatis Reinhardt & Lütken 1862: 269.
Anolis vincenti Garman 1887: 46.
Anolis trinitatis trinitatis — Underwood In Williams et al. 1959: 212.
Anolis trinitatis vincentii — Underwood In Williams et al. 1959: 213.
Anolis trinitatis — Schwartz & Henderson 1991: 349.


Holotype: ZMUC (UZM) R.37145. Type locality: “Trinidad”. Revised to “Kingstown, St. Vincent” by Lazell 1972.

Distribution: Anolis trinitatis inhabits the Lesser Antilles (St. Vincent, St. Lucia, Chateaubelair Island) and it has been introduced to Trinidad (Lazell 1972). May be present in Venezuela’s Peninsula de Paria (Donoso-Barros 1968).

Anolis trinitatis hybridizes with Anolis aeneus on Trinidad (Lazell 1972). However, hybrids show greatly reduced reproductive fitness (Gorman et al. 1971). Males, which reach 74 mm snout-to-vent, are green to green-blue, with blue stippling on the head and anterior trunk. They have yellow coloring on the throat and ventral surface, and the area around the eye is dark. Males have a large dewlap that extends into the abdominal region. Females are duller and have a smaller dewlap.

Synonymy: Boulenger (1885) listed Anolis trinitatis (and Anolis aeneus) as synonyms of Anolis alligator, which is now A. roquet.


Karyotype: Gorman et al. (1968) reported Anolis trinitatis and Anolis aeneus have 36 and 34 chromosomes, respectively. Hybrids have 35 chromosomes.

Literature
Donoso-Barros, R. 1968. The lizards of Venezuela (checklist and key). Carib. J. Sci. 8 (3-4): 105-122

Gorman GC. 1969. The zoogeography of Lesser Antillean Anolis lizards; an analysis based upon chromosomes and lactic dehydrogenases. Bull. Mus. Comp. Zool. Harvard 138 (3): 53-80.

Gorman GC, Atkins L. 1968. Natural hybridization between two sibling species of Anolis lizards: Chromosome cytology. Science 159 (March): 1358-1360

Gorman G, Licht P, Desshauser HC, Boos JO 1971. Reproductive failure among the hybridizing Anolis lizards of Trinidad. Syst. Zool. 20: 1–18

Lazell JD. 1972. The anoles (Sauria: Iguanidae) of the lesser Antilles. Bull. Mus. comp. Zool. Harvard 143 (1): 1-115

Poe S. 2004. Phylogeny of anoles. Herpetological Monographs 18: 37-89

Poe S. 2013. 1986 Redux: New genera of anoles (Squamata: Dactyloidae) are unwarranted. Zootaxa 3626 (2): 295–299

Reinhardt J, Lütken CH 1862. Bidrag tii det vestindiske Öriges og navnligen tii de dansk-vestindiske Oers Herpetologie. Vidensk. Meddel. Naturhist. For. Kjöbenhavn (10-18): 153-291 [1862]

Schwartz A, Henderson RW. 1991. Amphibians and Reptiles of the West Indies. University of Florida Press, Gainesville, 720 pp.

Williams EE, Quesnel VC, Kenny JS, Underwood G. 1959. The anoles of the Eastern Caribbean (Sauria, Iguanidae) Part. I. Preface; Part II. Two sibling species of anoles in Trinidad; Part. III. Revisionary notes. Bull. Mus. Comp. Zool. Harvard 121 (5): 185-226 + 1 plate

Anolis wattsi (Dactyloidae)

Anolis wattsi Boulenger 1894: 375. 
Type locality: Antigua. Restricted to St. John’s, St. John Parish, Antigua, by Lazell (1972). Syntypes: BMNH 1946.8.29.12-13. White and Hailey (2006) give a detailed account of this lizard from Trinidad. It is known from Anguilla, Antigua, Barbuda, St. Martin, St. Eustatius, St. Christopher, Nevis, St. Lucia, and was discovered in Trinidad in 1992. See this web site: http://www.ahailey.f9.co.uk/anolis.htm.

Giant Crown Anole, Anolis richardi Duméril & Bibron

Anolis Richardi Duméril & Bibron 1837: 141
Ptychonotus (Ctenodeira) richardii — Fitzinger 1843: 66
Anolis richardii — Boulenger 1885: 37
Anolis trossulus Garman 1887: 38
Anolis trossulus — Barbour 1916
Anolis richardii richardii — Underwood In Williams et al. 1959



Holotype: MNHN Ig. 53 (fide Underwood 1959); MCZ Type locality: “Tortola”. Revised to “Crown Point, Tobago” by Lazell 1972.

Distribution Lesser Antilles: Grenadines, Grenada, Tobago, St. Vincent; Surinam (introduced according to Hoogmoed 1981: 281). The Tobago population seems to have been introduced from the Grenadines or Grenada.

This is large, morphologically variable anole, with males reaching 140 mm snout-to-vent. It is often seen sitting on tree trunks with its head down. It has a dark green or brown dorsal surface, with a green-gray to yellow ventral surface. Its dewlap is orange, yellow, or gray-green. Females and juveniles often have a yellow or cream-colored lateral stripe. This species reaches relatively high population densities along streams.

Barbour T. 1916. Amphibians and reptiles of Tobago. Proc. Biol. Soc. Washington 29: 221-224

Boulenger G.A. 1885. Catalogue of the lizards in the British Museum (Natural History). Vol. 2, Second edition. London.

Creer DA, Kevin de Queiroz, Todd R. Jackman, Jonathan B. Losos and Allan Larson 2001. Systematics of the Anolis roquet series of the southern Lesser Antillies. Journal of Herpetology 35 (3): 428-441.

Duméril AMC, Bibron G. 1837. Erpétologie Générale ou Histoire Naturelle Complete des Reptiles. Vol. 4. Libr. Encyclopédique Roret, Paris, 570 pp.

Fitzinger L. 1843. Systema Reptilium, fasciculus primus, Amblyglossae. Braumüller et Seidel, Wien: 106 pp.

Garmam S. 1887. On West Indian reptiles. Iguanidae. Bull. Essex Inst. 19: 25-50.

Gorman GC, Atkins, L. 1968. The relationships of the Anolis of the roquet species group (Sauria: Iguanidae). III. Comparative study of display behavior. Breviora (284): 1-31

Hoogmoed MS. 1981. Introduced species of reptiles in Surinam. Notes on the herpetofauna of Surinam VIII. Amphibia-Reptilia 1: 277-285

Köhler G. 2014. Characters of external morphology used in Anolis taxonomy—Definition of terms, advice on usage, and illustrated examples. Zootaxa 3774 (2): 201–257

Lazell JD. 1972. The anoles (Sauria: Iguanidae) of the lesser Antilles. Bull. Mus. comp. Zool. Harvard 143 (1): 1-115

Malhotra A, Thorpe RS. 1999. Reptiles & Amphibians of the Eastern Caribbean. MacMillan, London & Oxford

Mertens R. 1972. Herpetofauna Tobagana. Stuttgarter Beitr. zur Naturkunde nr. 252 22 pp.

Poe S. 2004. Phylogeny of anoles. Herpetological Monographs 18: 37-89

Roughgarden J. 1995. Anolis Lizards of the Caribbean: Ecology, Evolution, and Plate Tectonics. Oxford University Press, 200 pp.

Schwartz A., Henderson RW. 1991. Amphibians and Reptiles of the West Indies. University of Florida Press, Gainesville, 720 pp.

Williams EE. Quesnel VC. Kenny JS, Underwood G. 1959. The anoles of the Eastern Caribbean (Sauria, Iguanidae) Part. I. Preface; Part II. Two sibling species of anoles in Trinidad; Part. III. Revisionary notes. Bull. Mus. Comp. Zool. Harvard 121 (5): 185-226 + 1 plate

Leaf Anole, Anolis planiceps (Family Dactyloidae)

Anolis planiceps Troschel 1848: 649

Anolis chrysolepis planiceps - Vanzolini & Williams 1970: 85

Anolis nitens nitens - Avila-Pires 1995


Type locality: Caracas, Venezuela. Holotype: ZMB 529.

Distribution. E Venezuela, Guyana, Brazil (Roraima, Amazonas), Trinidad.

The leaf anole, Anolis planiceps, occurs in Venezuela, Guyana, and Trinidad . Trinidad localities include Tucker Valley; Port of Spain, Maracas Valley, Nariva Swamp, Chacachacare and Huevos, Aripo savannas, Arena Forest; southwestern peninsula near Bonasse. St. Augustine; Mt. St. Benedict; La Seita; Toco; Morgua; Guayaguayare; Mayaro; Chacachacare Island; and Huevos Island. This lizard is found on tree trunks, on the ground, and sleeping on leaves at night. It is a forest and forest edge species and
until now the only species of Anolis on Trinidad that is native to the island. All other species have been introduced. However we have recently found Anolis tigrinus on Tobago and seen photographic evidence that it is also present on Trinidad.








Based on specimens labeled Anolis planiceps in Vertnet

Zandolie, Jungle Runner, Giant Ameiva, Ameiva atrigularis (Family Teiidae)


Ameiva surinamensis var. atrigularis Garman 1887.
Ameiva ameiva ameiva — Marcuzzi 1950: 101 (part.)
Ameiva ameiva tobagana — Peters & Donoso-Barros 1970: 20.
Ameiva ameiva atrigularis — Tuck & Hardy 1973: 241
Ameiva ameiva — Murphy 1997: 152
Ameiva atrigularis — Ugueto & Harvey 2011

Taxonomy & Systematics. This account follows
 Ugueto and Harvey (2011).

Distribution. Northern Venezuela including the Distrito Capital, Vargas, Miranda, Sucre, Peninsula de Paria, Isla de Margarita), Trinidad, Tobago, Little Tobago and satellites.

Natural History. This is a giant species of Ameiva, with adults reaching a body size that may exceed 0.5 m. The Zandolie is omnivorous, feeding on insects, small vertebrates, and fruits. It is visible on bright sunny days when it is
searching for food and mates, at other times they
are
concealed in burrows. Females are oviparous and probably lay several clutches of eggs annually. Habitats used are deciduous tropical and premontane humid forest, forest-edge, as well as areas of forest with open undergrowth where patches of light reach the forest floor, scrub lands, secondary growth, suburban yards and gardens, roadsides, and beaches.



Barbour, T. and Noble G.K.. 1915. A revision of the lizards of the genus Ameiva. Bull. Mus. Comp. Zool. Harvard 59: 417-479. - get paper here

Burt, C.E. & Burt, M.D. 1931. South American lizards in the collection of the American Museum of Natural History. Bull. Amer. Mus. nat. Hist. 61 (7): 227-395

Garman, S. 1887. On the West Indian Teiids in the Museum of Comparative Zoology. Bull. Essex Inst. 19: 1-12.

Gorzula, Stefan & Senaris, J. C. 1999. In: Contribution to the herpetofauna of the Venezuelan Guayana. I: a data base. Scientia Guaianae, Caracas, No. 8 [1998], 269+ pp.

Murphy, J.C. 1997. Amphibians & Reptiles of Trinidad & Tobago. Krieger Publishing Company, Malabar, Florida, 304 pp.

Peters, James A. & Donoso-Barros, R. 1970. Catalogue of the Neotropical Squamata: Part II. Lizards and Amphisbaenians. Bull. US Natl. Mus. 297: 293 pp.

Rivas Fuenmayor, G. and Barrio Amorós C.L. 2005. New Amphibian and Reptile records from Cojedes State, Venezuela. Herpetological Review 36 (2):205-209.

Roze, J.A. 1964. La herpetologia de la Isla de Margarita, Venezuela. Soc. de Cienc. Natur. 'La Salle', Caracas, Mem. 24(69): 209-241.

Tuck, R. G., Hardy, J.D. 1973. Status of the Ober Tabago collection, Smithsonian Institution, and the proper allocation of Ameiva surinamensis tobaganus Cope (Sauria: Teiidae). Proc. Biol. Soc. Washington 86 (19): 231-242

Ugueto, G. N. and Harvey M.B. 2011. Revision of Ameiva ameiva Linnaeus (Squamata: Teiidae) in Venezuela: Recognition of Four Species and Status of Introduced Populations in Southern Florida, USA. Herpetological Monographs 25 (1): 112-170.

Friday, April 1, 2016

Paradox Frog, Pseudis paradoxus caribensis Galllardo

Rana paradoxa Linnaeus, 1758: 212. Type locality: Suriname.

Pseudes paradoxa — Wiegmann, 1832: 148.

Pseudis paradoxus caribensis Gallardo, 1961. Holotype: MCZ 19890, by original designation. Type locality: Mayaro Bay, Trinidad.


Adults are 45-75 mm in total length. Body is short and stout; eyes dorsolateral; dorsal and ventral skin; fingers free of webbing, toes heavily webbed. Dorsum green anteriorly, brown to green posteriorly; flash marks on posterior surface of femur. Ventral immaculate white. has the largest, or near largest tadpoles known when compared to adult size. Tadpoles are 220 mm (3 to 4 times longer than adult frogs). As currently defined it is known from the following countries: Argentina, Bolivia, Brazil, Colombia, French Guiana, Guyana, Paraguay, Peru, Suriname, Trinidad (unknown from Tobago), and Venezuela. Usually found in open marshy areas with floating vegetation, in permanent and temporary ponds. Calls can be heard day and night and consists of a sequence of 8-11 pulses, but frogs are very wary during the day. At night they can be seen floating on the surface and grasping vegetation with their hands. Mating may be stimulated by rainfall. Eggs are green and laid along the shore among aquatic plants.

 Abstract from Aguiar et al. (2007)
The previous uncertain placement of Lysapsus and Pseudis within the neobatrachians was recently resolved by molecular and morphological studies, which supported them as members of the Hylinae subfamily. Their inter- and intrageneric relationships, however, have long been under debate and no studies shed light on these questions. Aiming to elucidate such questions, this paper used 3.2 kb comprising the mitochondrial genes 12S, tRNA valine, 16S and cytochrome b, and the nuclear exon 1 coding for rhodopsin, to all representatives of both genera (except to two subspecies of Pseudis paradoxa). The results identified three major clades: the clade 1 was composed by Lysapsus species and subspecies; clade 2 included the subspecies of the Pseudis paradoxa (Pseudis paradoxa paradoxa, P. paradoxa platensis and P. paradoxa occidentalis), P. fusca, P. bolbodactyla and P. tocantins, and clade 3 was composed by Pseudis southern Brazil species (Pseudis cardosoi and P. minuta). As closely related taxa we found Pseudis minuta + P. cardosoi; P. tocantins + P. fusca, and the subspecies within each genus. Evidence that Pseudis is not monophyletic with respect to Lysapsus was found and we suggest Lysapsus to be a junior synonym of Pseudis. Based on pair-wise comparison among gene sequences, we also suggest that the subspecies of Pseudis paradoxa and Lysapsus limellum must be considered as full species.

Aguiar O, Bacci M, Lima AP, Rossa‐Feres D, Haddad CF, Recco‐Pimentel SM. Phylogenetic relationships of Pseudis and Lysapsus (Anura, Hylidae, Hylinae) inferred from mitochondrial and nuclear gene sequences. Cladistics. 2007 Oct 1;23(5):455-63.

Pseudis paradoxa habitat on Trinidad
Range of Pseudis paradoxa from AmphibiaWeb

Range of Pseudis paradoxa on Trinidad

Suriname Toad, Pipa pipa (Family Pipidae)

Rana Pipa Linnaeus, 1758: 210.
Pipa americana Laurenti, 1768: 25.

Ranges throughout South America from Ecuador to Guianas and southward to Peru, Bolivia, and Brazil. It is also present on Trinidad (absent from Tobago). Pipa pipa is aquatic inhabiting the muddy bottoms of tropical rivers. The fingers end in star-shaped, filamentous tactile organs to detect food. Pipa pipa uses vacuum feeding similar to that of the mata mata turtle. It feeds on aquatic invertebrates and fish. During courtship the male fertilizes the eggs and then positions them so they attach to the female's back. The skin of the female grows around the fertilized eggs and the embryos develop with the females skin. They hatch after 90-120 days of incubation as miniature adults, about 20 mm in length. On Trinidad all of the records are from the southern portion of the island.

Bromeliad-dwelling Treefrog, Phytotriades auratus (Family Hylidae)

Amphodus auratus Boulenger, 1917:185. Syntypes: BMNH 1917.9.27.39-41. Type locality: Mount Tucuche, a little above 3000 feet altitude, Trinidad.

Phyllodytes auratus — Bokermann, 1968, Rev. Brasil. Biol., 28: 157-160.

Phytotriades auratus — Jowers, Downie, and Cohen, 2009, Stud. Neotrop. Fauna Environ., 43: 186.

Previously known from the type locality at the summit of Mt. Tucuche and Mt Aripo. it has been recently discovered on Cerro de Humo, Península de Paria, Sucre, Venezuela , in Venezuela. It does not occur on Tobago. The total range of the species is estimated to be 10km² or less on Trinidad, and about 2 hectares in Venezuela. It is known from 940-1250 m asl. It is found in montane rainforest and elfin woodland. It has a very specific microhabitat and is associated with the giant epiphytic bromeliad, Glomeropitcairnia erectiflora. It breeds by larval development in bromeliads. Probably the greatest threat to this species is climate change. As the climate becomes warmer and drier, it could be threatened by fires. Rivas and Freitas. 2015. Herp Review 46(2):153-156. 

Flashy Narrow-mouth Frog, Elachistocleis surinamensis (Family Microhylidae)




See comment under E. ovalis. This species ranges widely in northern Venezuela, east to northern Suriname, and it is also present on Trinidad (unknown from Tobago). It occurring up to at least 500 m asl, and all records are from close to sea level. It is terrestrial in tropical forest, including closed-canopy secondary forest, and possibly also coastal savannahs. In Trinidad 
animals occur in the forest leaf-litter; males may be found calling from small water-filled stump holes. It is not found in open country on Trinidad. but is associated with wooded areas at Aripo Savanna. Spawning takes place in temporary pools, the eggs float and are non-adhesive, and the larvae develop in the same water bodies. It has been suggested this species is possibly the same species as E. ovalis. It is possible that Relictivomer pearsei is a synonym of this species (and it was treated as such by Kenny (1969).  Like other members of the genus it is a leaf litter inhabitant that feeds on ants.

Leaf-nesting Frog, Phyllomedusa trinitatis (Family Hylidae)

Phyllomedusa trinitatis Mertens, 1926: 145. Holotype: SMF 2633. Type locality: Port of Spain, Trinidad.

Phyllomedusa burmeisteri trinitatis — Rivero, 1961, Bull. Mus. Comp. Zool., 126: 148.

Phyllomedusa trinitatis — Rivero, 1961, Bull. Mus. Comp. Zool., 126: 148.


Pithecopus trinitatis — Lutz, 1966, Copeia, 1966: 236.

This species is widespread on Trinidad, it is unknown from Tobago. It has also been reported from the Paria Peninsula, but the exact identity of the Venezuelan populations remains to be determined. In elevation it ranges from about sea level to more than 950 m. It is a forest and savanna species that uses the understory trees and bushes often near ponds. It is also found by ditches in secondary forests and urbanized habitats with standing water and adjacent vegetation. The female folds a leaf or leaves into a nest which are held together by the adhesive properties of the gelatinous covering of the eggs. When the eggs hatch the tadpoles drop into the water below. We have found the cat-eyed snake (Leptodeira annulata ashmeadi) very close to the nests of this frog.

Trinidad Stream Frog, Mannophryne trinitatis (Family Aromobatidae)

A male transporting tadpoles.
Phyllobates trinitatis Garman, 1888 (1887), Bull. Essex Inst., 19: 13. Type locality: "Trinidad".

Prostherapis trinitatis — Mole and Urich, 1894, J. Trinidad Field Nat. Club, 2: 87.

Prostherapis trinitatis trinitatis — Rivero, 1961, Bull. Mus. Comp. Zool., 126: 155.

Colostethus trinitatus — Edwards, 1971, Proc. Biol. Soc. Washington, 84: 148.

Mannophryne trinitatus — La Marca, 1992, Cuad. Geograf., Mérida, 9: 32.

Mannophryne trinitatis — La Marca, 1994, Publ. Asoc. Amigos Doñana, 4: 52.

Barrio-Amorós et al., (2006) provided morphological and chromosomal evidence that Mannophryne trinitatis is restricted to the island of Trinidad. The Peninsula de Paria, Venezuelan population was found to be a different species, Mannophryne venezuelensis (Manzanilla et al., 2007).

Distribution. Forested areas of the Northern and Central Ranges of the island of Trinidad, occurring from the upper elevations of the mountainous areas to sea level.

Natural History. Associated with gravel, and sandy substrate streams with shaded. Females deposit eggs at terrestrial locations guarded by the male. Males transport tadpoles to predator free pools that may be some distance from the stream.



Barrio Amorós, C.L. 2004. Amphibians of Venezuela Systematic List, Distribution and References, An Update. Review of Ecology in Latin America 9(3): 1-48.

Barrio-Amorós, C.L. Rivas, G., Molina, C., and Kaiser, H. 2006. Mannophryne trinitatis (Anura: Dendrobatidae) a Trinidadian single-island endemic. Herpetological Review 37: 298-299.

Edwards, S.R. 1974. Taxonomic notes on South American Dendrobatid frogs of the genus Colostethus. Occasional Papers of the Museum of Natural History of the University of Kansas: 1-14.

IUCN. 2010. IUCN Red List of Threatened Species (ver. 2010.2). Available at:http://www.iucnredlist.org.

Jowers, M.J. and Downie, J.R. 2004. Distribution of the frog Mannophryne trinitatis (Anura: Dendrobatidae) in Trinidad, West Indies. Living World. Journal of the Trinidad and Tobago Field Naturalists´ Club: 17-19.

Kenny, J.S. 1969. The Amphibia of Trinidad. Studies on the Fauna of Curacao and Other Caribbean Islands.

La Marca, E. 1992. Catálogo taxonómico, biogeográfico y bibliográfico de las ranas de Venezuela. Cuadernos Geográficos, Universidad de Los Andes 1: 1-197.

La Marca, E. 1994. Taxonomy of the frogs of the genus Mannophryne (Amphibia: Anura: Dendrobatidae). Publicaciones de la Asociacion Amigos de Doñana: 1-75.

La Marca, E. 1995. Biological and systematic synopsis of a genus of frogs from Northern mountains of South America (Anura: Dendrobatidae: Mannophryne). Bulletin of the Maryland Herpetological Society: 40-77.

La Marca, E. 1997. Lista actualizada de los anfibios de Venezuela. In: La Marca, E. (ed.), Vertebrados Actuales y Fosiles de Venezuela, pp. 103-120. Museo de Ciencias y Tecnología de Mérida, Mérida.

Manzanilla, J., García-París, M. and La Marca, E. 2002. Filogenia molecular del Género Mannophryne. VII Congreso Luso-Español de Herpetología, Evora, pp. 61. Portugal.

Manzanilla, J., Jowers, M.J., La Marca, E., and García-París, M. 2007. Taxonomic reassessment of Mannophryne trinitatis (Anura: Dendrobatidae) with a description of a new species from Venezuela. The Herpetological Journal 17: 31-42.

Manzanilla, J., La Marca, E. and García-París, M. 2003. Evidencias moleculares de la monofilia del género Mannophryne (Anura: Dendrobatidae). VI Congreso Latinoamericano de Herpetología, Enero 2003, Lima, Perú.

Mole, R.R. and Urich, F.W. 1894. A preliminary list of the reptiles and batrachians of the island of Tobago. Journal of the Trinidad Field Naturalists’ Club: 77-90.

Murphy, J.C. 1997. Amphibians and Reptiles of Trinidad and Tobago. Krieger Publishing Company, Malabar, Florida.

Parker, H.W. 1933. A list of the frogs and toads of Trinidad. Tropical Agriculture 10(1): 8-12.