DEDICATION

We are happy to dedicate this paper to our friend and

colleague, Josiah H. Townsend, Associate Professor of

Biology at Indiana University of Pennsylvania, in Indiana,

Pennsylvania. Over the last two decades, since he was

a student in one of Larry Wilson’s classes, Joe has built

an imposing reputation as the principal authority on the

herpetofauna of the biogeographically significant Chortis

Highlands of northern Central America. During this

time he amassed important collections, and their study

is demonstrating that the herpetofaunal diversity of this

region of Mesoamerica has been seriously underestimated,

especially among anurans, salamanders, and squamates.

Since 2006, Joe has authored or co-authored the descriptions

of 21 new taxa from northern Central America, including

one anuran, 10 salamanders, four lizards, and six snakes.

He also has produced important summary papers on the

Mesoamerican herpetofauna, including several coauthored

chapters in the 2010 book Conservation of Mesoamerican

Amphibians and Reptiles (Wilson et al. 2010; Johnson et al.

2010; Townsend and Wilson 201 0a, b), a2014 paper entitled

“Characterizing the Chortis Block Biogeographic Province:

geological, physiographic, and ecological associations

and herpetofaunal diversity,’ and a 2016 paper entitled

“Amphibians of the Cordillera Nombre de Dios, Honduras:

COI barcoding suggests underestimated taxonomic

diversity in a threatened endemic fauna.” Additionally, Joe

co-authored the 2006 book The Amphibians and Reptiles

of the Honduran Mosquitia (McCranie et al. 2006) and the

2008 book Amphibians and Reptiles of Cusuco National

Park, Honduras (Townsend and Wilson 2008). During

his career Joe has collaborated with a sizable number

of colleagues and students to underscore the significant

biodiversity of northern Central America, as an important

component of the overall Mesoamerican herpetofauna. The

collections he assembled in remote regions will continue

to provide insights into the phylogenetic relationships and

phylogeography of this herpetofauna, and we believe his

trajectory will position him as one of the most influential

herpetologists of his era.

The frog in this photograph is a partially metamorphosed

individual of Rana (Lithobates) lenca, recently described

as a new species from Honduras in the following paper:

Luque-Montes, Ileana, James D. Austin, Kayla D.

Weinfurther, Larry David Wilson, Erich P. Hofmann, and

Amphib. Reptile Conserv.

Josiah H. Townsend. 2018. An integrative assessment

of the taxonomic status of putative hybrid leopard frogs

(Anura: Ranidae) from the Chortis Highlands of Central

America, with description of a new species. Systematics

and Biodiversity 2018: 1-17. This paper is an example

of the seminal work being conducted by Joe Townsend

and his colleagues, which is exposing the underestimated

herpetofaunal diversity of the biogeographically significant

Chortis Highlands. The frog was photographed in a shallow

pond above the Thomas Cabot Biological Station at an

elevation of 1,640 m, within Reserva Biologica Cerro

Uyuca in the department of Francisco Morazan. The pond

is located in pine forest, along a trail that leads from the

biological station to the summit of Cerro Uyuca. This frog

is an inhabitant of the “weeping woods” or cloud forest, so

beautifully described by Archie Carr in the first chapter of

his 1953 book High Jungles and Low.

“ae

glee

Josiah H. Townsend photographed tn 2008 along the Rio

Arcaqual tn Parque Nacional Celaque, on a trail between

the visitors’ center and the summit of Cerro Celaque. At that

time Joe was undergoing a marathon period of fieldwork

in an effort to assess the composition, distribution, and

conservation status of the amphibian herpetofauna of

Honduras. Cerro Celaque is the highest mountain in

Honduras, and a site that contains substantial herpetofaunal

diversity and endemicity. Ongoing research in Joe’s lab

likely will add to the diversity of salamander species found

on this mountain. Joe is considered the principal authority

on the herpetofauna of the Chortis Highlands of Central

America and is one of the leading investigators employing

next-generation techniques in molecular systematics to

recover underestimated phylogenetic diversity, especially in

threatened endemic herpetofaunas. Photo by Ileana Luque-

Montes.

January 2019 | Volume 13 | Number 1 | e168

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

13(1) [General Section]: 1-64 (e168).

The endemic herpetofauna of Central America:

a casualty of anthropocentrism

‘Vicente Mata-Silva, Dominic L. DeSantis, *Eli Garcia-Padilla, *Jerry D. Johnson,

and ®Larry David Wilson

'24Department of Biological Sciences, The University of Texas at El Paso, El Paso, Texas 79968-0500, USA *Oaxaca de Juarez, Oaxaca 68023,

MEXICO *Centro Zamorano de Biodiversidad, Escuela Agricola Panamericana Zamorano, Departamento de Francisco Morazan, HONDURAS

Abstract.—The endemic herpetofauna of Central America is of global significance, and currently consists of

623 species, 56.9% of a total herpetofauna of 1,095 species. During the last two years 43 endemic species have

been added to this total, and one species has been deleted. The endemic herpetofauna of Central America is

distributed unevenly among 10 physiographic regions, ranging from six species in the Yucatan Platform to

254 in the Isthmian Central American highlands. The distributions of close to three quarters of the 623 species

are limited to a single physiographic region, and our assessment of their conservation status indicates that

about nine-tenths of these species lie within the high vulnerability range of the Environmental Vulnerability

Score (EVS). We prioritized the conservation significance of the Central American species by combining

the data on physiographic distribution with those of the EVS and recognize 14 priority levels. About eight of

every 10 endemic species occupy the first two priority levels, i.e., high vulnerability species limited to one or

two physiographic regions. Protecting the endemic component of the Central American herpetofauna is the

greatest challenge currently facing conservation professionals working in this region. We conclude that this

goal will not be reached until humanity, in general, addresses the issues generated by the widespread adoption

of the anthropocentric worldview.

Keywords. Anthropocentric worldview, anurans, caudates, caecilians, conservation significance, endemism, extinc-

tion risk, squamates, turtles

Resumen.—La herpetofauna endemica de Centroamerica es de importancia global y actualmente consiste de

623 especies, 56.9% de una herpetofauna total de 1,095 especies. Durante los dos ultimos anos, 43 especies

endemicas han sido agregadas a la lista, y una especie ha sido eliminada. La herpetofauna endémica de

Centroameérica esta distribuida de forma desigual entre 10 regiones fisiograficas, que va de seis especies

en la Plataforma de Yucatan, a 254 en las Tierras Altas del Istmo Centroamericano. Las distribuciones de

aproximadamente tres cuartos de las 623 especies estan limitadas a una sola region fisiografica, y nuestra

evaluacion sobre su estatus de conservacion indica que alrededor de nueve décimas de estas especies se

localizan dentro de la categoria de vulnerabilidad alta del sistema de puntaje de vulnerabilidad ambiental

(EVS). Ordenamos la importancia de conservacion de las especies centroamericanas combinando los datos

sobre su distribucion fisiografica con los de EVS, y reconocemos 14 niveles de prioridad. Aproximadamente

ocho de cada 10 especies endémicas ocupan los dos primeros niveles de prioridad, por ejemplo, especies

con vulnerabilidad alta limitadas a una o dos regiones fisiograficas. La proteccion del componente endémico

de la herpetofauna de Centroamérica es el mayor reto para los profesionales de la conservacion que trabajan

en esta region. Concluimos que esta meta no sera lograda hasta que la humanidad en general confronte los

problemas generados por una vision antropoceéntrica del mundo.

Palabras Claves. Vision antropocéntrica del mundo, anuros, caudados, cecilios, importancia de conservacion, ende-

mismo, riesgo de extincion, escamosos, tortugas

Citation: Mata-Silva, V, DeSantis DL, Garcia-Padilla E, Johnson JD, Wilson LD. 2019. The endemic herpetofauna of Central America: a casualty of

anthropocentrism. Amphibian & Reptile Conservation 13(1) [General Section]: 1-64 (e168).

NoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any medium, provided

the original author and the official and authorized publication sources are recognized and properly credited. The official and authorized publication

credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation; official journal website <amphibian-

reptile-conservation.org>.

Received: 30 April 2018; Accepted: 10 September 2018; Published: 20 January 2019

Correspondence. ! vmata@utep.edu * didesantis@miners.utep.edu * eligarcia_18@hotmail.com *jjohnson@utep.edu * bufodoc@aol.com

Amphib. Reptile Conserv. 3 January 2019 | Volume 13 | Number 1 | e168

Mata-Silva et al.

“.. losses, whether of species, landscapes, or seascape,

-a world without-will become simply the new default. But

whether we notice them or not, these losses matter be-

cause each diminishes our experiences, pleasures, and

possibilities...”

David W. Orr (2016)

Introduction

Johnson et al. (2017) examined the biodiversity and

conservation status of the endemic herpetofauna of Mexico,

which then consisted of 789 species and presently amounts

to 792 species. Johnson et al. (2017) concluded that this

endemic herpetofauna is of global significance and has

become severely imperiled as a consequence of actions

by humans. These authors calculated that the endemic

Mexican herpetofauna constituted 61.0% of the total of

1,292 herpetofaunal species in the country.

This paper is a companion piece to Johnson et al. (2017),

and places the 623 endemic species in Central America into

14 conservation priority levels. Our approach is the same as

that stipulated by Johnson et al. (2017) in their Introduction

and section on Biodiversity Decline.

Given that the foundation and conclusions of this paper

are based on the results of Johnson et al. (2017), we briefly

quote the first four conclusions in that paper (see p. 614), as

follows:

“A. The complex interplay among the atmosphere,

hydrosphere, and lithosphere allows for the existence of life

on planet Earth.”

“B. Humans are faced with the consequences of an

interrelated amalgam of global problems of their own

making, which impact the atmosphere, hydrosphere,

lithosphere, and biosphere. These problems are sufficiently

grave to threaten the continued existence of life.”

“C. Biodiversity decline is a problem of global dimensions.

This decline impacts life at all levels: from the ecosystem,

through the species comprising these ecosystems, to the

genes prescribing the traits of these species.”

“—D. Throughout its history, life has been subjected to a series

of mass extinction episodes that have preceded the current

sixth episode of humanity’s design.”

In this paper we underscore the significance of the endemic

herpetofauna of Central America and assess its conservation

status. The continued existence of this endemic herpetofauna

hinges on sustaining the region’s life-support systems,

i.e., the group of interacting elements in the atmosphere,

hydrosphere, and lithosphere that allow for the maintenance

of life on the planet. As a rational species, we believe that

humans are responsible for protecting and preserving the

diversity of life on the planet, as well as improving the

Amphib. Reptile Conserv.

quality of life for humans and other life forms. For this

reason, we are diametrically opposed to the ideas recently

promulgated by R. Alexander Pyron in a perspective

published in The Washington Post (2017), that “we don’t

need to save endangered species” because “extinction 1s

part of evolution,” and further that, “the only creatures we

should go out of our way to protect are Homo sapiens.”

Recent Changes to the Central American

Herpetofauna

Even while facing an increasing tempo for global biodiversity

decline (Ceballos et al. 2017), herpetologists continue to

increase the number of known species of amphibians and

reptiles from around the world (see the AmphibiaWeb and

Reptile Database websites). Johnson et al. (2015) reported

92 additional taxa to the Central American herpetofaunal

list, based on the cutoff date established for the list in Wilson

and Johnson (2010). For this paper, which considers the

endemic members of the Central American herpetofauna,

we added 19 taxa of amphibians and 24 of reptiles to the

list presented in Johnson et al. (2015). We list these taxa and

their supportive publications below.

Incilius mayordomus—Savage et al. 2013. Copeia 2013:

8—12. New species.

Hyalinobatrachium dianae—Kubicki et al. 2015. Zootaxa

3920: 69-84. New species.

Craugastor gabbi—Arias et al. 2016. Zootaxa 4132: 347-

363. New species.

Diasporus darienensis—Batista et al. 2016a. Zoological

Journal of the Linnean Society 178: 267-311. New

Species.

Diasporus majeensis—Batista et al. 2016a. Zoological

Journal of the Linnean Society 178: 267-311. New

Species.

Diasporus pequeno—Batista et al. 2016a. Zoological

Journal of the Linnean Society 178: 267-311. New

Species.

Diasporus sapo—Batista et al. 2016a. Zoological Journal

of the Linnean Society 178: 267-311. New species.

Plectrohyla_ calvata—McCranie. 2017a. Mesoamerican

Herpetology 4: 389-401. New species.

Smilisca manisorum—McCranie. 2017c. Mesoamerican

Herpetology 4: 512-526. Resurrected from synonymy

of Smilisca baudinit.

Lithobates lenca—Luque-Montes et al. 2018. Systematics

and Biodiversity 2018: 1-17. New species.

Bolitoglossa aurae—Kubicki and Arias. 2016. Zootaxa

4184: 329-346. New species.

Bolitoglossa chucutaniensis—Batista et al. 2014.

Mesoamerican Herpetology 1: 96-121. New species.

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The endemic herpetofauna of Central America

Cryptotriton xucaneborum—Rovito et al. 2015. Zoological

Journal of the Linnean Society 175: 150-166. New

Species.

Nototriton costaricense—Arias and Kubicki. 2018. Zootaxa

4369: 487-500. New species.

Nototriton nelsoni—Townsend. 2016. Zootaxa 4196: 511—

528. New species.

Nototriton oreadorum—Townsend. 2016. Zootaxa 4196:

511-528. New species.

Oedipina _ berlini—Kubicki. —_2016.

Herpetology 3: 819-840. New species.

Mesoamerican

Oedipina capitalina—Solis et al. 2016. Salamandra 42:

125-133. New species.

Oedipina salvadorensis—Brodie et al. 2012. Journal of

Herpetology 46: 233—240. Resurrected from synonymy

of Oedipina taylori.

Celestus laf—Lotzkat et al. 2016. Mesoamerican

Herpetology 3: 962-975. New species.

Mesaspis cuchumatanus—Solano-Zavaleta et al. 2016.

Journal of Herpetology 50: 327-335. New species

Mesaspis salvadorensis—Solano-Zavaleta and _ Nieto-

Montes de Oca. 2018. Molecular Phylogenetics and

Evolution 120: 16-27. Elevated from subspecies to

species level.

Dactyloa brooksi—Poe and Ryan. 2017. Amphibian &

Reptile Conservation 11: 1-16. Resurrected from the

synonymy of Dactyloa insignis.

Dactyloa kathydayae—Poe and Ryan. 2017. Amphibian &

Reptile Conservation 11: 1-16. New species.

Dactyloa maia—Batista et al. 2015b. Zootaxa 4039: 57-84.

New species.

Dactyloa savagei—Poe and Ryan. 2017. Amphibian &

Reptile Conservation 11: 1-16. New species.

Norops elcopeensis—Poe et al. 2015. Amphibian & Reptile

Conservation 9: 1-13. New species.

Norops mccraniei—Kohler et al. 2016. Mesoamerican

Herpetology 3: 8-41. New species.

Norops oxylophus—McCranie and Kohler. 2015. Bulletin

of the Museum of Comparative Zoology SPS(1): 1-292.

Recognition as distinct from Norops lionotus.

Norops_ wilsoni—Kohler et al. 2016. Mesoamerican

Herpetology 3: 8-41. New species.

Lepidoblepharis emberawoundule—Batista et al. 2015a.

Zootaxa 3994: 187-221. New species.

Lepidoblepharis rufigularis—Batista et al. 2015a. Zootaxa

3994: 187-221. New species.

Lepidoblepharis_ victormartinezi—Batista et al. 2015a.

Zootaxa 3994: 187-221. New species.

Ameiva fuliginosa—McCranie and Gotte. 2014.

Proceedings of the Biological Society of Washington

Amphib. Reptile Conserv.

127: 543-556. Elevation from subspecies to species

level.

Holcusus miadis—Meza-Lazaro et al. 2015. Zoological

Journal of the Linnean Society 2015: 1-22. Elevation

from subspecies to species level.

Tantilla berguidoi—Batista et al. 2016b. Mesoamerican

Herpetology 3: 949-960. New species

Tantilla excelsa—McCranie and Smith. 2017. Herpetologica

73: 338-348. New species.

Tantilla gottei—McCranie and Smith. 2017. Herpetologica

73: 338-348. New species.

Tantilla stenigrammi—McCranie and Smith. 2017.

Herpetologica 73: 338-348. New species.

Rhadinella_ lisyae—McCranie. 2017b. Mesoamerican

Herpetology 4. 243-253. New species.

Epictia martinezi—Wallach. 2016. Mesoamerican

Herpetology 3: 216-374. New species.

Epictia pauldwyeri—Wallach. 2016. Mesoamerican

Herpetology 3: 216-374. New species.

Bothriechis nubestris—Doan et al. 2016. Zootaxa 4138:

271-290. New species.

The Distributional Status of Dipsas viguieri

Peters (1960) reviewed the taxonomic status of members

of the subfamily Dipsadinae, and placed nine species in

the Dipsas articulata group, collectively distributed from

western and southeastern Mexico to northwestern Ecuador.

Peters’ view of the species-level relationships among the

members of the articulata group was impacted by the

paucity of specimens of each taxon known at that time,

and he noted that only minimal scale and color differences

separated certain species. Peters (1960) indicated the range

of D. viguieri as the Pacific coast of Panama.

Pérez Santos and Moreno (1988) reported on a specimen

of D. gracilis from the Pacific coast of Colombia (see

discussion below), and Pérez Santos (1999) noted the

occurrence of D. viguieri from both versants of Panama,

including a specimen from the province of Bocas del Toro

in the western part of the country. Subsequently, Kohler

(2001; 2003; 2008) noted the range of Dipsas viguieri as

eastern Panama and western Colombia but did not provide

additional information. Cadle (2005), in a paper on the

systematics of the Dipsas oreas complex, tentatively referred

to a specimen (FMNH 74376) from northwestern Colombia

near the Panama border as D. viguieri, which previously

had been identified as D. gracilis. Nonetheless, Cadle

(2005) stated that these two taxa were not distinguishable

by any reported characteristics, and on p. 128 noted that,

“Without additional study, I am unable to adequately

differentiate Dipsas viguieri (eastern Panama and northern

Choco, Colombia) and D. gracilis (western Ecuador and

extreme northern Peru).” Further, based on an examination

of morphological characters, Cadle indicated geography as

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Mata-Silva et al.

Plate 1. Ate/opus varius (Lichtenstein and Martens, 1856). The Harlequin Frog is a priority ten species with an EVS of 11, distributed

on both versants of the cordilleras of Costa Rica and western Panama (Frost 2018). This individual is from one of three known

surviving populations of this species, and in Costa Rica it is being surveyed near Uvita, in the province of Puntarenas. Photo by

César Barrio-Amoros.

distribution to Volcan Barva, Costa Rica (Frost 2018). This individual was located in Alto del Roble, in the province of Heredia.

Photo by Victor Acosta-Chaves.

Plate 3. [ncilius melanochlorus (Cope, 1877). The Wet Forest Toad is a priority eight species with an EVS of 12, with a distribution

on the Atlantic versant of Costa Rica and adjacent Panama, and likely in adjacent Nicaragua (Frost 2018). This individual was found

in Centro Soltis, San Isidro de Pefias Blancas, in the province of Alajuela, Costa Rica. Photo by Victor Acosta-Chaves.

Amphib. Reptile Conserv. 6 January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

the only currently reliable means of assigning names to these

species. Similarly, in discussing a number of poorly-known

Dipsas from South America, Harvey (2008) commented that

he was unable to distinguish D. viguieri from D. gracilis, and

thus did not include D. viguieri in his key but noted that the

D. articulata complex requires further study.

Jaramillo et al. (2010) and Johnson et al. (2015) regarded

Dipsas viguieri as endemic to Panama. Wallach et al. (2014:

235), however, considered D. viguieri as occupying “Eastern

Panama (Canal Zone, Darién, Panama) and Colombia (?

Choco, Piura), NSL-60 m.” In their D. gracilis account,

however, these authors noted, “Colombian record doubtful

fide Cadle (2005: 123): possibly D. viguieri. Dipsas gracilis

and D. viguieri possibly conspecific fide Harvey (2008:

429).” Wallach et al. (2014: 232), however, apparently

confused the information provided by Cadle (2005), as the

FMNH specimen tentatively was referred to D. viguieri and

not D. gracilis. Finally, Ray (2017) indicated the range of

D. viguieri as eastern Panama to northwestern Colombia.

The historical timeline for information on the distribution

and taxonomic status of Dipsas viguieri has been unclear,

as different workers have maintained that this species

is endemic to Panama or occurs in both Panama and

Colombia. In the absence of a definitive analysis involving

morphological and molecular approaches, for the purpose

of this paper we are considering D. viguieri as not endemic

to Panama.

Global Status of the Central

Herpetofauna

American

As with the Mexican herpetofauna (Johnson et al. 2017),

the Central American herpetofauna also is highly diverse,

consisting of 60 families, 214 genera, and 1,095 species

(Table 1), organized into six orders (Anura, Caudata,

Gymnophiona, Crocodylia, Squamata, and Testudines).

The level of herpetofaunal diversity in Central America

is intermediate between that found in Mexico and North

America (United States—Canada). The number of species

in the United States-Canada is the same as Johnson et

al. (2017) reported, i.e., 650 (Center for North American

Herpetology website; accessed 9 December 2017). Johnson

et al. (2017) reported the number of species in Mexico as

1,292.

Even though the number of herpetofaunal species

occurring in Central America is intermediate between that

found in the United States-Canada and Mexico, Central

America contains about 8.5 times the number of taxa by

area as found in Mexico, and 155.6 times the number found

in the United States—Canada. Thus, the relative degree of

biodiversity 1s significantly higher in Central America when

compared to that in Mexico and the United States—Canada.

If we consider Central America as a single region tn our

analysis (i.e., not one divided into seven countries), then its

herpetofauna also is significant when compared to that of

other areas in Latin America. With respect to amphibians,

the 509 species occurring in Central America is the fifth

Amphib. Reptile Conserv.

largest in Latin America (amphibiaweb.com; 15 April

2018), and is closest to that for the country of Ecuador, at

562. The area/species ratio for Ecuador however, is 504.6,

compared to 998.0 for Central America.

Considering the numbers of crocodylian, squamate,

and turtle species, the 586 species in Central America is

comparable to that recorded for the neighboring country

of Colombia, which is 611 (reptile-database.org; accessed

29 December 2017). Colombia, however, with an area of

1,141,748 km/?, is 2.25 times the size of Central America,

which contains an area of 507,966 km? (www. Oéi.es/

historico/cultura2/Colombia/03.htm; accessed 29 December

2017). Thus, the area/species ratio for Colombia is 1,868.7,

compared to 868.0 for that of Central America. Only Brazil

(799) and Colombia in South America contain more species

than Central America (reptile-database.org; accessed 29

December 2017).

Endemism within the Central American

Herpetofauna

The proportion of herpetofaunal endemism in Central

America is slightly less than in Mexico, the other major

segment of Mesoamerica. The percentage in Central

America is 56.9 (Table 2) compared to 61.1 in Mexico

(Johnson et al. 2017). This percentage in Central America

is based on an endemic herpetofauna of 623 species and

a total herpetofauna of 1,095 species (Table 2). Both of

the comparable figures for the Mexican herpetofauna are

higher, 1.e., 789 and 1,292 (Johnson et al. 2017). As noted

by Johnson et al. (2015: 26), “Mesoamerica is one of the

world’s most important biodiversity reservoirs, and Central

America contains a substantial component of that region’s

herpetofauna.” We illustrate the breakdown of the total and

endemic components of the Central American herpetofauna

in Fig. 1. This graph shows the close correspondence

between the endemic and total number of salamander

species, the relatively distant correspondence between the

endemic and total number of squamate species, and the

intermediate correspondence between the two figures for

anurans (Fig. 1).

Of the 60 families represented in Central America, 38

(63.3%) contain endemic species (Table 2). This leaves

22 families with no endemic representation, including

the anuran families Aromobatidae, Hemiphractidae, and

Rhinophrynidae, the crocodylian families Alligatoridae

and Crocodylidae, the squamate families Amphisbaenidae,

Hoplocercidae, Polychrotidae, Xenosauridae, Boidae,

Charinidae, Loxocemidae, Natricidae, Sibynophiidae,

and Tropidophiidae, and the turtle families Cheloniidae,

Chelydridae, Dermatemydidae, Dermochelyidae,

Emydidae, Staurotypidae, and Testudinidae. In Central

America these are small-content families, with species

numbers ranging from one to five (Table 2). The families

with endemic representation have total numbers ranging

from one to 166; the endemic numbers vary from one to

143 (Table 2).

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Mata-Silva et al.

Endemic and Total Components of the

Central American Herpetofauna

600

560

500

328

Number of Species

200

100

15 ; A 23 ,

6 =... = _

Anura Caudata Gymnophiona Crocodylia Squamata Testudines

Native Endemic

Fig. 1. Graph comparing the endemic and total number of species for the Central American herpetofauna, arranged by order.

Table 1. Diversity of the Central American herpetofauna at the familial, generic, and specific levels.

Orders Families Genera Species

Anura 14 58 328

Caudata 1 7 166

Gymnophiona 7 4 15

Crocodylia 2 2 3

Squamata 32 130 560

Testudines 9 13 D8

Totals 60 214 1,095

Table 2. Degree of endemism of the Central American herpetofauna at the species level, arranged by family.

Total Number of Species Number of Endemic Species | Percentage of Endemism

l = =

Aromobatidae

Bufonidae

Centrolenidae

Craugastoridae

Dendrobatidae

Eleutherodactylidae

Hemiphractidae

Hylidae

Leptodactylidae

Microhylidae

Phyllomedusidae

Pipidae

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The endemic herpetofauna of Central America

Table 2 (continued). Degree of endemism of the Central American herpetofauna at the species level, arranged by family.

Total Number of Species Number of Endemic Species | Percentage of Endemism

11 6

Ranidae

Rhinophrynidae

Subtotals

Plethodontidae

Subtotals

Caeciltidae

Dermophiidae

Subtotals

Totals

Alligatoridae

Crocodylidae

Subtotals

Amphisbaenidae

Anguidae

Corytophanidae

Dactyloidae

Eublepharidae

Gymnophthalmidae

Helodermatidae

Hoplocercidae

Iguanidae

Mabuyidae

Phrynosomatidae

Phyllodactylidae

Polychrotidae

Scincidae

Sphaerodactylidae

Sphenomorphidae

Telidae

Xantusiidae

Xenosauridae

Anomalepididae

Boidae

Charinidae

Colubridae

Dipsadidae

Elapidae

Leptotyphlopidae

Loxocemidae

Natricidae

Sibynophiidae

Tropidophiidae

Typhlopidae

Viperidae

Subtotals

Cheloniidae

Amphib. Reptile Conserv. 9 January 2019 | Volume 13 | Number 1 | e168

Mata-Silva et al.

Table 2 (continued). Degree of endemism of the Central American herpetofauna at the species level, arranged by family.

Chelydridae

Dermatemydidae

Dermochelyidae

Emydidae

Geoemydidae

Kinosternidae

Staurotypidae

Testudinidae

Subtotals

Totals

Sum Totals

Of the 14 anuran families with representatives in Central

America, 11 contain endemic species, which include 192

(58.5%) of the total of 328 species (Table 2). Of these 11

families, the largest numbers of endemics are 77 in the

Craugastoridae and 52 in the Hylidae. Other than tn the

Pipidae, with one total and one endemic species (100%),

as might be expected, the percentage of endemism is next

highest in the Craugastoridae (75.5%), but the third highest is

in the Dendrobatidae (66.7%), and not the Hylidae (60.9%).

The value for the Eleutherodactylidae (62.5%) also is higher

than that for the Hylidae. The remaining families contain

from one to 24 endemic species (Table 2).

A single family of salamanders, the Plethodontidae,

occurs in Central America. The percentage of endemism

(86.1%) is amazingly high and is the highest in all the 38

families represented (Table 2).

The endemic species of caecilians (seven) make up less

than one-half (46.7%) of the total number of 15 in Central

America. Three of the endemics are caeciliids and four are

dermophiids.

None of the three species of crocodylians in Central

America is endemic. Crocodylus acutus and Caiman

crocodilus rather are among the naturally most broadly

distributed herpetofaunal species in the Western Hemisphere.

The squamates are the most speciose group of

herpetofaunal organisms in Central America, with 560

species distributed among 32 families (Table 2). Only the

endemic proportions of turtles (8.7%) and crocodylians

(0.0%) are lower than those of squamates (49.8%). The

endemic squamates are more or less evenly divided between

the lizards (143) and snakes (136). Of the 19 families of

lizards with representatives in Central America, 15 contain

endemic species (78.9%). The largest numbers of endemic

squamate species are found within the families Dactyloidae

(74) and Dipsadidae (77). The next largest number of

endemic lizards (25) 1s allocated to the family Anguidae.

The remaining 12 lizard families contain only one to 13

endemic species. The percentage of endemism among the

lizard families ranges from 11.1% in the Corytophanidae

to 80.0% in the Mabuyidae (Table 2). Thirteen families of

snakes are represented in Central America, of which seven

Amphib. Reptile Conserv.

er of Species Number of Endemic Species | Percentage of Endemism

p) = =

contain endemic species (53.8%). The greatest numbers of

endemic species are in the families Dipsadidae (77) and

Colubridae (30). The next largest number (16) lies within the

family Viperidae; the remaining four snake families contain

from one to six endemic species (Table 2). The percentage

of endemism among the snake families ranges from 31.6%

in the Elapidae to 60.0% in the Typhlopidae (Table 2).

The percentage of endemism among turtles is very low,

with only two such species among a total of 23 (8.7%), one

each in the families Geoemydidae and Kinosternidae (Table

2).

At the ordinal level, the percentage of endemism 1s

highest among the salamanders at 86.1%, the same figure

as for the family Plethodontidae, since it is the only family

of salamanders found in Central America (Table 2). The

percentage is lowest among the crocodylians at 0.0%.

Intermediate values are evident for the anurans (58.5%),

squamates (49.8%), and caecilians (46.7%).

The level of herpetofaunal endemism tn Central America

is comparable to that found in the other portions of the North

American continent, 1.e., Mexico, as well as the United

States—Canada (Table 3). The overall level for Central

America, however, is slightly lower (56.9%) than for either

Mexico (61.5%) or the United States—Canada (61.2%).

Although, as expected, the total number of herpetofaunal

species is lower in the United States—Canada than to the

south in Mesoamerica (Table 3); even with the greater area

of the two northern nations, the level of endemism still is

impressive. Of the total of 650 species in the United States—

Canada, 398 are endemic, for a percentage of endemism of

61.2%. Notably, this level of endemism is based heavily on

the amphibians, especially the salamanders. The proportion

of amphibian endemism is more than 10 points higher

in the United States-Canada (78.6%) than in Mexico

(68.3%) or Central America (67.2%). The percentage of

endemism is higher for both anurans and salamanders in

the United States—Canada (64.4% and 86.4%, respectively)

than for these two groups in either Mexico (60.0% and

82.8%, respectively) or Central America (58.5% and

86.1%, respectively). Significantly, the level of amphibian

endemism in all three regions heavily depends on the

January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

Plate 4. Craugastor sandersoni (Schmidt, 1941). Sanderson’s

Rainfrog is a priority two species with an EVS of 19, which is

distributed on the “Caribbean slopes of the Maya Mountains in

east-central Belize southward to the Montafias del Mico in eastern

Guatemala and westward into the Sierra de Santa Cruz, the eastern

portion of the Sierra de las Minas, and the foothills of the northern

Alta Verapaz” (Frost 2018). This individual was found in Montafias

del Mico, Guatemala. Photo by Sean Michael Rovito.

Plate 6. Oophaga pumilio (Schmidt, 1857). The Strawberry

Poison Frog is a priority one species with an EVS of 16, and

its distribution extends along the Atlantic versant from “eastern

Nicaragua...south through the lowlands of Costa Rica and

northwestern Panama (Savage 2002: 388). This individual came

from the mainland in the province of Bocas del Toro, Panama.

Photo by Abel Batista.

Plate 5. Oophaga granulifera (Taylor, 1958). The Granular Poison

Frog 1s a priority two species with an EVS of 17, distributed in the

Golfo Dulce region of Pacific coastal Costa Rica and presumably

in adjacent Panama (Frost 2018). This individual was encountered

in Ciudad Cortéz de Osa, in the province of Puntarenas, Costa

Rica. Photo by Victor Acosta-Chaves.

=’ -

Plate 7. Oophaga vicentei wee Weyoldt, and Juraska,

1996). This dendrobatid frog is a priority two species with an

EVS of 16, distributed on the Atlantic versant “of the provinces

of Veraguas and Coclé and the upper reaches of Pacific versant

in Coclé, central Panama” (Frost 2018). This individual is from

Santa Fé National Park, in the province of Veraguas, Panama.

Photo by Abel Batista.

Plate 8. Phyllobates lugubris (Schmidt, 1857). The Lovely Poison

Frog is a priority one species with an EVS of 17, which ranges

along the Atlantic versant from “extreme southeastern Nicaragua to

northwestern Panama; a single record from just west of the Panama

Canal” (Savage 2002: 390). This individual was found in the Donoso

region, in the province of Colon, Panama. Photo by Abel Batista.

Amphib. Reptile Conserv.

Plate 9. Diasporus hylaeformis (Cope, 1876). The Pico Blanco

Robber Frog is a priority one species with an EVS of 17, with

a distribution in the highlands of the cordilleras of Costa Rica

and Panama (Frost 2018). This individual came from Alto del

Roble, in the province of Heredia, Costa Rica. Photo by Victor

Acosta-Chaves.

January 2019 | Volume 13 | Number 1 | e168

Mata-Silva et al.

Table 3. Total number of species, endemic species, and relative endemism within the herpetofaunal groups in Central America,

North America (United States—Canada), and Mexico. Data for Central America from this paper, for North America from CNAH

(www.cnah.org; accessed 9 January 2018), and for Mexico from updated figures in Johnson et al. (2017).

Herpetofaunal Total Endemic Relative Total

Groups Speciesin Speciesin Endemism Species

Central Central in Central in North

America America America America

(“%)

Anurans 328 192 58.5 104

Salamanders 166 143 86.1 191

Caecilians LS 7 46.7 —

Subtotals 509 342 67.2 295

Crocodylians 3 — — 2

Squamates 560 279 49.8 287

Turtles 23 2 8.3 66

Subtotals 586 281 48.0 355

Totals 1,095 623 56.9 650

salamanders, 1.e., over 80%. Curiously enough, such high

incidences of salamander species-level endemism are not

correlated with the incidence of family-level endemism,

which decreases markedly from the United States—Canada

through Mexico to Central America. The 191 salamander

species in the United States—Canada are organized within

eight families, but with the majority allocated to the family

Plethodontidae (Center for North American Herpetology

website; accessed 2 January 2018). Four of these eight

families, the Amphiumidae, Cryptobranchidae, Proteidae,

and Rhyacotritonidae occur no farther south than the United

States. Four families, the Ambystomatidae, Plethodontidae,

Salamandridae, and Sirenidae are represented in Mexico, but

with the greatest number of species in the Plethodontidae,

as in the United States—Canada. Only a single family, the

Plethodontidae, is found in Central America. In the United

States—Canada, 145 of the 191 species of salamanders are

in the family Plethodontidae (75.9%). Of the 151 species

found in Mexico 130 (86.1%) are in the Plethodontidae

(Wilson et al., 2017); the remaining 21 species are allocated

to three families (See above). Finally, of the 166 species

of salamanders found in Central America, all are in the

Plethodontidae; obviously the percentage of occupancy is

100%.

Unlike the situation among the amphibians, the level of

endemism among the crocodylians, squamates, and turtles

is significantly lower in all three regions dealt with in Table

3. The level of endemism among these taxa is lowest in

the United States—Canada (46.8%), next lowest in Central

America (48.0%), and highest tn Mexico (58.6%). Given

that squamates constitute the largest group, compared

to the other two, the same pattern would be expected for

them as for the entire group. Thus, the level of squamate

endemism is lowest in the United States—Canada (40.8%),

intermediate in Central America (49.8%), and highest in

Mexico (60.0%).

The differential between the percentages of endemism

for amphibians versus the remainder of the herpetofauna

Amphib. Reptile Conserv.

Endemic Relative Total Endemic Relative

Species Endemism - Speciesin Speciesin Endemism

in North in North Mexico Mexico in Mexico

America America (%)

(%)

67 64.4 247 148 60.0

165 86.4 151 125 82.8

— — 3 1 33.3

232 78.6 401 274 68.3

1 50.0 3 — —

117 40.8 863 517 60.0

48 72.7 51 20 39.2

166 46.8 917 537 58.6

398 61.2 1,318 811 61.5

(Table 3) increases from that seen in Mexico (9.7%), through

Central America (19.2%), to the United States—Canada

(31.8%). Thus, in all three regions amphibians contribute

more to the degree of endemicity than the remainder of the

herpetofauna (Table 3).

Physiographic Distribution of the Endemic

Central American Herpetofauna

Given the considerable global significance of the diversity

and endemicity of the Central American herpetofauna, as

documented above, it is of paramount importance to protect

its elements. As an initial step to determine the distributional

patterns of these organisms in Central America, we collated

the available information on the occurrence of the members

of the herpetofauna among the 10 physiographic regions

traditionally recognized in this portion of Mesoamerica

(Campbell, 1999; Wilson and Johnson, 2010; Fig. 2). Six

of these regions occupy the lowlands of Central America,

including the Yucatan Platform, the Caribbean lowlands of

eastern Guatemala and northern Honduras, the Caribbean

lowlands from Nicaragua to Panama, the Pacific lowlands

from eastern Chiapas to south-central Guatemala,

the Pacific lowlands from southeastern Guatemala to

northwestern Costa Rica, and the Pacific lowlands from

central Costa Rica through Panama (Table 4). Four regions

are found in the highlands of Central America, including the

western nuclear Central American highlands, the eastern

nuclear Central American highlands, the Isthmian Central

American highlands, and the highlands of eastern Panama

(Table 4). We document the distribution of the 623 endemic

members of the Central American herpetofauna among the

10 physiographic regions in Table 4, and summarize these

data in Table 5 and Fig. 3.

Thetotal number ofthe endemic species distributed within

the 10 physiographic regions ranges from a low of six in the

Yucatan Platform to 254 in the Isthmian Central American

highlands (Table 5). The mean regional occupancy figure

January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

»=Jamaica-

Guatemala

> XSEl'Salvadors

Nicaragua

Fig. 2. Physiographic regions of Central America, after Campbell (1999). Abbreviations are as follows: CG = western nuclear

Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from

central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP =

highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean

lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central

American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP =

Yucatan Platform.

Table 4. Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions. Abbreviations

for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to

south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands);

CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern

Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes

associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to

Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. ? = species known from indeterminate locality

(see text).

Physiographic Regions of Central America Totals

Anura (192 species)

Bufonidae (24 species)

Atelopus certus

Atelopus chiriquiensis

Atelopus chirripoensis

Atelopus limosus

Atelopus senex

Atelopus varius

Atelopus zeteki

Amphib. Reptile Conserv. 13 January 2019 | Volume 13 | Number 1 | e168

Mata-Silva et al.

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic

regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands

from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes

associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific

lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern

Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands

from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. ? = species known from

indeterminate locality (see text).

Incilius aucoinae

Incilius chompipe

Incilius epioticus

Incilius fastidiosus

Incilius guanacaste

Incilius holdridgei

Incilius ibarrai

Incilius karenlipsae

Incilius leucomyos

Incilius majordomus

Incilius melanochlorus

Incilius periglenes

Incilius peripatetes

Incilius porteri

Incilius signifer

Rhinella centralis

Rhinella chrysophora

Centrolenidae (4 species)

Cochranella granulosa

Hyalinobatrachium dianae

Hyalinobatrachium talamancae

Hyalinobatrachium vireovittatum

Craugastoridae (77 species)

Craugastor adamastus

Craugastor anciano

Craugastor andi

Craugastor angelicus

Craugastor aphanus

Craugastor aurilegulus

Craugastor azueroensis

Craugastor bocourti

Craugastor bransfordii

Craugastor campbelli

Craugastor catalinae

Craugastor chac

Craugastor charadra

Craugastor chingopetaca

Craugastor chrysozetetes

Craugastor coffeus

Craugastor cruzi

Craugastor cuaquero

Amphib. Reptile Conserv. 14 January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic

regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands

from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes

associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific

lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern

Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands

from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. ? = species known from

indeterminate locality (see text).

Craugastor cyanochthebius

Craugastor daryi

Craugastor emcelae

Craugastor emleni

Craugastor epochthidius

Craugastor escoces

Craugastor evanesco

Craugastor fecundus

Craugastor fleischmanni

Craugastor gabbi

Craugastor gollmeri

Craugastor gulosus

Craugastor inachus

Craugastor jota

Craugastor laevissimus

Craugastor lauraster

Craugastor megacephalus

Craugastor melanostictus

Craugastor merendonensis

Craugastor milesi

Craugastor mimus

Craugastor monnichorum

Craugastor myllomyllon

Craugastor nefrens

Craugastor noblei

Craugastor obesus

Craugastor olanchano

Craugastor omoaensis

Craugastor pechorum

Craugastor persimilis

Craugastor phasma

Craugastor podiciferus

Craugastor polyptychus

Craugastor punctariolus

Craugastor ranoides

Craugastor rayo

Craugastor rhyacobatrachus

Craugastor rivulus

Craugastor rostralis

Craugastor rugosus

Craugastor sabrinus

Amphib. Reptile Conserv.

Physiographic Regions of Central America Totals

| Physiographie Regions of CentralAmerica |

Int NS (SRE eos (Tn [eee Nn cn eae

January 2019 | Volume 13 | Number 1 | e168

Mata-Silva et al.

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic

regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands

from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes

associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific

lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern

Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands

from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. ? = species known from

indeterminate locality (see text).

Physiographic Regions of Central America Totals

[______Phesigraphie Resins of entat Amerie

Craugastor saltuarius + 1

Craugastor sandersoni + F 2

Craugastor stadelmani + 1

Craugastor stejnegerianus of 3

Craugastor tabasarae 1

Craugastor talamancae 1

Craugastor taurus 1

Craugastor trachydermus + 1

Craugastor underwoodi 1

Craugastor xucanebi F 1

Pristimantis adnus + 1

Pristimantis altae z

Pristimantis caryophyllaceus 2

Pristimantis cerasinus + 3

Pristimantis museosus 1

Pristimantis pardalis 3

Pristimantis pirrensis + 1

Strabomantis laticorpus + 1

Dendrobatidae (14 species)

Ameerega maculata? ?

Andinobates claudiae 1

Andinobates geminisae 1

Colostethus latinasus = 1

Ectopoglossus astralogaster ste 1

Ectopoglossus isthminus ae 1

Oophaga arborea 1

Oophaga granulifera 2

Oophaga pumilio 1

Oophaga speciosa 1

Oophaga vicentei 4p 2

Phyllobates lugubris 1

Phyllobates vittatus 1

Silverstoneia flotator 3

Eleutherodactylidae (10 species)

Diasporus citrinobapheus 1

Diasporus darienensis =; 1

Diasporus diastema F +

Diasporus hylaeformis 1

Diasporus igneus 1

Diasporus majeensis aD 1

Diasporus pequeno ae 1

Diasporus sapo ts 1

Amphib. Reptile Conserv. 16 January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic

regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands

from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes

associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific

lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern

Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands

from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. ? = species known from

indeterminate locality (see text).

eee “ae of Central America Totals

Amphib. Reptile Conserv. 17 January 2019 | Volume 13 | Number 1 | e168

Diasporus tigrillo

Diasporus ventrimaculatus

Hylidae (52 species)

Atlantihyla panchoi

NY WN

Atlantihyla spinipollex

—

Bromeliohyla melacaena

—

Dryophytes bocourti

—

Duellmanohyla legleri

Duellmanohyla lythrodes

—

Duellmanohyla rufioculis

Duellmanohyla salvadorensis

Duellmanohyla salvavida

Duellmanohyla soralia

Duellmanohyla uranochroa

Ecnomiohyla bailarina

Ecnomiohyla fimbrimembra

e NY NY NY NY NY WN

Ecnomiohyla minera

—

Ecnomiohyla rabborum

—

Ecnomiohyla salvaje

Ecnomiohyla sukia

—

Ecnomiohyla thysanota

—

Ecnomiohyla veraguensis

—

Exerodonta catracha

—

Exerodonta perkinsi

Hyloscirtus colymba

—

Isthmohyla angustilineata

—

Isthmohyla calypso

Isthmohyla debilis

— —

Isthmohyla graceae

—

Isthmohyla infucata

—

Isthmohyla insolita

Isthmohyla lancasteri

—

Isthmohyla picadoi

—

Isthmohyla pictipes

—

Isthmohyla pseudopuma

—

Isthmohyla rivularis

—

Isthmohyla tica

—

Isthmohyla xanthosticta

—

Isthmohyla zeteki

—

Plectrohyla calvata

—

Plectrohyla chrysopleura

Mata-Silva et al.

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic

regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands

from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes

associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific

lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern

Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands

from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. ? = species known from

indeterminate locality (see text).

es “ee of Central America Totals

Plectrohyla dasypus

Plectrohyla exquisite

Plectrohyla glandulosa +

Plectrohyla pokomchi +

Plectrohyla psiloderma

Plectrohyla quecchi

Plectrohyla tecunumani

Plectrohyla teuchestes

Ptychohyla dendrophasma

Ptychohyla hypomykter

+ + + + + +

Quilticohyla sanctaecrucis

Scinax altae

Smilisca manisorum

Smilisca puma

Leptodactylidae (1 species)

Leptodactylus silvanimbus

Microhylidae (1 species)

Hypopachus pictiventris

Phyllomedusidae (2 species)

Agalychnis annae

Agalychnis saltator

Pipidae (1 species)

Pipa myersi

Ranidae (6 species)

Lithobates juliani +

Lithobates lenca

Lithobates miadis

Lithobates taylori

Lithobates vibicarius

Lithobates warszewitschii

Caudata (143 species)

Plethodontidae (143 species)

Bolitoglossa alvaradoi

Bolitoglossa anthracina

Bolitoglossa aurae

Bolitoglossa aureogularis

Bolitoglossa bramei

Bolitoglossa carri

Bolitoglossa cataguana

Bolitoglossa celaque

Bolitoglossa centenorum of

Amphib. Reptile Conserv.

+ + + + +

a a \

+ 1

+ 2

ee ee ee ee ee)

—

January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic

regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands

from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes

associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific

lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern

Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands

from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. ? = species known from

indeterminate locality (see text).

Bolitoglossa cerroensis

Bolitoglossa chucantiensis

Bolitoglossa colonnea

Bolitoglossa compacta

Bolitoglossa conanti

Bolitoglossa copia

Bolitoglossa cuchumatana

Bolitoglossa cuna

Bolitoglossa daryorum

Bolitoglossa decora

Bolitoglossa diaphora

Bolitoglossa diminuta

Bolitoglossa dofleini

Bolitoglossa dunni

Bolitoglossa epimela

Bolitoglossa eremia

Bolitoglossa gomezi

Bolitoglossa gracilis

Bolitoglossa heiroreias

Bolitoglossa helmrichi

Bolitoglossa huehuetenanguensis

Bolitoglossa indio

Bolitoglossa insularis

Bolitoglossa jacksoni

Bolitoglossa jugivagans

Bolitoglossa kamuk

Bolitoglossa kaqchikelorum

Bolitoglossa la

Bolitoglossa lignicolor

Bolitoglossa longissima

Bolitoglossa magnifica

Bolitoglossa marmorea

Bolitoglossa meliana

Bolitoglossa minutula

Bolitoglossa mombachoensis

Bolitoglossa morio

Bolitoglossa nigrescens

Bolitoglossa ninadormida

Bolitoglossa nussbaumi

Bolitoglossa nympha

Bolitoglossa obscura

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| Physiographic Regions of Central America

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Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic

regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands

from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes

associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific

lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern

Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands

from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. ? = species known from

indeterminate locality (see text).

Physiographic Regions of Central America Totals

ee PS See via ee | GER

Amphib. Reptile Conserv. 20 January 2019 | Volume 13 | Number 1 | e168

Bolitoglossa odonnelli

—

Bolitoglossa omniumsanctorum

—

Bolitoglossa oresbia

—

Bolitoglossa pacaya

—

Bolitoglossa pesrubra

—

Bolitoglossa porrasorum

—

Bolitoglossa psephena

—

Bolitoglossa pygmaea

—

Bolitoglossa robinsoni

—

Bolitoglossa robusta

Bolitoglossa salvinii

Ww N

Bolitoglossa schizodactyla

—

Bolitoglossa sombra

—

Bolitoglossa sooyorum

—

Bolitoglossa splendida

Bolitoglossa striatula

—

Bolitoglossa subpalmata

—

Bolitoglossa suchitanensis

—

Bolitoglossa synoria

—

Bolitoglossa taylori

—

Bolitoglossa tenebrosa

—

Bolitoglossa tica

—

Bolitoglossa tzultacaj

—

Bolitoglossa xibalba

—

Bolitoglossa zacapensis

—"

Cryptotriton monzoni

Cryptotriton nasalis

—

Cryptotriton necopinus

—

Cryptotriton sierraminensis

—

Cryptotriton veraepacis

—

Cryptotriton xucaneborum

—

Dendrotriton bromeliacius

—

Dendrotriton chujorum

—

Dendrotriton cuchumatanus

—

Dendrotriton kekchiorum

-¢

—

Dendrotriton rabbi

—

Dendrotriton sanctibarbarus

—

Nototriton abscondens

—

Nototriton barbouri

—

Nototriton brodiei

—

Nototriton costaricense

The endemic herpetofauna of Central America

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic

regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands

from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes

associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific

lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern

Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands

from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. ? = species known from

indeterminate locality (see text).

Nototriton gamezi

Nototriton guanacaste

Nototriton lignicola

Nototriton limnospectator

Nototriton major

Nototriton matama

Nototriton mime

Nototriton nelsoni

Nototriton oreadorum

Nototriton picadoi

Nototriton picucha

Nototriton richardi

Nototriton saslaya

Nototriton stuarti

Nototriton tapanti

Nototriton tomamorum

Oedipina alfaroi

Oedipina alleni

Oedipina altura

Oedipina berlini

Oedipina capitalina

Oedipina carablanca

Oedipina chortiorum

Oedipina collaris

Oedipina cyclocauda

Oedipina fortunensis

Oedipina gephyra

Oedipina gracilis

Oedipina grandis

Oedipina ignea

Oedipina kasios

Oedipina koehleri

Oedipina leptopoda

Oedipina maritima

Oedipina motaguae

Oedipina nica

Oedipina nimaso

Oedipina pacificensis

Oedipina paucidentata

Oedipina petiola

Oedipina poelzi

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Mata-Silva et al.

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic

regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands

from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes

associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific

lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern

Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands

from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. ? = species known from

indeterminate locality (see text).

Oedipina pseudouniformis

Oedipina quadra

Oedipina salvadorensis

Oedipina savagei

Oedipina stenopodia

Oedipina stuarti

Oedipina taylori

Oedipina tomasi

Oedipina tzutujilorum

Oedipina uniformis

Pseudoeurycea exspectata

Gymnophiona (7 species)

Caecilidae (3 species)

Caecilia volcani

Oscaecilia elongata

Oscaecilia osae

Dermophiidae (4 species)

Dermophis costaricensis

Dermophis gracilior

Dermophis occidentalis

Gymnopis multiplicata

Squamata (279 Species)

Anguidae (25 species)

Abronia anzuetoi

Abronia aurita

Abronia campbelli

Abronia fimbriata

Abronia frosti

Abronia gaiophantasma

Abronia meledona

Abronia montecristoi

Abronia salvadorensis

Abronia vasconcelosii

Celestus adercus

Celestus atitlanensis

Celestus bivittatus

Celestus cyanochloris

Celestus hylaius

Celestus laf

Celestus montanus

Celestus orobius

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The endemic herpetofauna of Central America

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic

regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands

from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes

associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific

lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern

Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands

from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. ? = species known from

indeterminate locality (see text).

Physiographic Regions of Central America Totals

| Physiographie Regions of CentralAmerica |

Rise ES Ree as eee ee

Amphib. Reptile Conserv. 23 January 2019 | Volume 13 | Number 1 | e168

Celestus scansorius

Coloptychon rhombifer

Diploglossus bilobatus

Diploglossus montisilvestris

Mesaspis cuchumatanus

Mesaspis monticola

Mesaspis salvadorensis

Corytophanidae (1 species)

Basiliscus plumifrons

Dactyloidae (74 species)

Dactyloa brooksi

Dactyloa casildae

Dactyloa ginaelisae

Dactyloa ibanezi

Dactyloa insignis

Dactyloa kathydayae

Dactyloa kunayalae

Dactyloa maia

Dactyloa microtus

Dactyloa savagei

Norops alocomyos

Norops altae

Norops amplisquamosus

Norops apletophallus

Norops aquaticus

Norops benedikti

Norops bicaorum

Norops campbelli

Norops carpenteri

Norops charlesmyersi

Norops cobanensis

Norops cryptolimifrons

Norops cupreus

Norops cusuco

Norops datzorum

Norops elcopeensis

Norops fortunensis

Norops fungosus

Norops gruuo

Norops haguei

Norops heteropholidotus

Mata-Silva et al.

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic

regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands

from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes

associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific

lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern

Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands

from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. ? = species known from

indeterminate locality (see text).

Norops humilis

Norops intermedius

Norops johnmeyeri

Norops kemptoni

Norops kreutzi

Norops leditzigorum

Norops limifrons

Norops lionotus

Norops loveridgei

Norops macrophallus

Norops magnaphallus

Norops marsupialis

Norops mccraniei

Norops monteverde

Norops morazani

Norops muralla

Norops ocelloscapularis

Norops osa

Norops oxylophus

Norops pachypus

Norops pijolensis

Norops polylepis

Norops pseudokemptoni

Norops pseudopachypus

Norops purpurgularis

Norops quaggulus

Norops roatanensis

Norops rubribarbaris

Norops salvini

Norops sminthus

Norops tenorioensis

Norops townsendi

Norops triumphalis

Norops tropidolepis

Norops utilensis

Norops villai

Norops wampuensis

Norops wellbornae

Norops wermuthi

Norops wilsoni

Norops woodi

Amphib. Reptile Conserv.

Physiographic Regions of Central America Totals

| Physiographie Regions of CentralAmerica |

iS ec Da ae Res eee

= = WYO —>— —

— —

+ +

eels mares Om, Wee 2

_ Re

+ 6

+ 3

+ 1

+ 4

January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic

regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands

from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes

associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific

lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern

Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands

from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. ? = species known from

indeterminate locality (see text).

Norops yoroensis

Norops zeus

Eublepharidae (1 species)

Coleonyx mitratus

Gymnophthalmidae (3 species)

Bachia blairi

Echinosaura panamensis

Echinosaura apodema

Helodermatidae (1 species)

Heloderma charlesbogerti

Iguanidae (7 species)

Ctenosaura bakeri

Ctenosaura flavidorsalis

Ctenosaura melanosterna

Ctenosaura oedirhina

Ctenosaura palearis

Ctenosura praeocularis

Ctenosaura quinquecarinata

Mabuyidae (4 species)

Marisora alliacea

Marisora magnacornae

Marisora roatanae

Marisora unimarginata

Phrynosomatidae (2 species)

Sceloporus lunaei

Sceloporus malachiticus

Phyllodactylidae (3 species)

Phyllodactylus insularis

Phyllodactylus palmeus

Phyllodactylus paralepis

Scincidae (1 species)

Mesoscincus managuae

Sphaerodactylidae (13 species)

Lepidoblepharis emberawoundule

Lepidoblepharis rufigularis

Lepidoblepharis victormartinezi

Sphaerodactylus alphus

Sphaerodactylus dunni

Sphaerodactylus graptolaemus

Sphaerodactylus guanaje

Sphaerodactylus homolepis

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Mata-Silva et al.

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic

regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands

from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes

associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific

lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern

Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands

from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. ? = species known from

indeterminate locality (see text).

Sphaerodactylus leonardovaldesi

Sphaerodactylus millepunctatus

Sphaerodactylus pacificus

Sphaerodactylus poindexteri

Sphaerodactylus rosaurae

Sphenomorphidae (1 species)

Scincella rara

Teiidae (5 species)

Cnemidophorus duellmani

Cnemidophorus ruatanus

Holcosus leptophrys

Holcosus miadis

Holcosus quadrilineatus

Xantusiidae (2 species)

Lepidophyma mayae

Lepidophyma reticulatum

Anomalepididae (1 species)

Helminthophis frontalis

Colubridae (30 species)

Dendrophidion apharocybe

Dendrophidion crybelum

Dendrophidion paucicarinatum

Dendrophidion rufiterminorum

Drymobius melanotropis

Leptodrymus pulcherrimus

Leptophis nebulosus

Mastigodryas alternatus

Mastigodryas dorsalis

Oxybelis wilsoni

Scolecophis atrocinctus

Tantilla albiceps

Tantilla armillata

Tantilla bairdi

Tantilla berguidoi

Tantilla brevicauda

Tantilla excelsa

Tantilla gottei

Tantilla hendersoni

Tantilla jani

Tantilla lempira

Tantilla olympia

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| Physiographie Regions of CentralAmerica |

HESS EN | CRE En | cea hee Een PSEA

January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic

regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands

from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes

associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific

lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern

Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands

from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. ? = species known from

indeterminate locality (see text).

Physiographic Regions of Central America Totals

[___Phnsigraphic Regions of entat Amerie

Tantilla psittaca

Tantilla ruficeps

Tantilla stenigrammi

Tantilla taeniata

Tantilla tecta

Tantilla tritaeniata

Tantilla vermiformis

Trimorphodon quadruplex

Dipsadidae (77 species)

Adelphicos daryi

Adelphicos ibarrorum

Adelphicos veraepacis

Atractus darienensis

Atractus depressiocellus

Atractus hostilitractus

Atractus imperfectus

Chapinophis xanthocheilus

Coniophanes joanae

Crisantophis nevermanni

Cubophis brooksi

Dipsas articulata

Dipsas bicolor

Dipsas nicholsi

Dipsas tenuissima

Enulius bifoveatus

Enulius roatanensis

Geophis bellus

Geophis brachycephalus

Geophis championi

Geophis damiani

Geophis downsi

Geophis dunni

Geophis fulvoguttatus

Geophis godmani

Geophis hoffmanni

Geophis nephodrymus

Geophis ruthveni

Geophis talamancae

Geophis tectus

Geophis zeledoni

Hydromorphus concolor

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Mata-Silva et al.

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic

regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands

from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes

associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific

lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern

Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands

from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. ? = species known from

indeterminate locality (see text).

Physiographic Regions of Central America Totals

| Physiographie Regions of CentralAmerica |

Se [EN CREE ee Pave cE Roa Re

Amphib. Reptile Conserv. 28 January 2019 | Volume 13 | Number 1 | e168

Hydromorphus dunni

Imantodes phantasma

Leptodeira rhombifera

Leptodeira rubricata

Ninia celata

Ninia espinali

Ninia maculata

Ninia pavimentata

Ninia psephota

Omoadiphas aurula

Omoadiphas cannula

Omoadiphas texiguatensis

Rhadinaea calligaster

Rhadinaea pulveriventris

Rhadinaea sargenti

Rhadinaea stadelmani

Rhadinaea vermiculaticeps

Rhadinella anachoreta

Rhadinella hempsteadae

Rhadinella lisyae

Rhadinella montecristi

Rhadinella pegosalyta

Rhadinella pilonaorum

Rhadinella rogerromani

Rhadinella serperaster

Rhadinella tolpanorum

Sibon anthracops

Sibon argus

Sibon carri

Sibon lamari

Sibon longifrenis

Sibon manzanaresi

Sibon merendonensis

Sibon miskitus

Sibon noalamina

Sibon perissostichon

Trimetopon barbouri

Trimetopon gracile

Trimetopon pliolepis

NON N

Trimetopon simile

Trimetopon slevini

The endemic herpetofauna of Central America

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic

regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands

from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes

associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific

lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern

Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands

from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. ? = species known from

indeterminate locality (see text).

Physiographic Regions of Central America Totals

| Physiographie Regions of CentralAmerica |

DEES cE an Te |e Na een ce eae

Amphib. Reptile Conserv. 29 January 2019 | Volume 13 | Number 1 | e168

Trimetopon viquezi

Urotheca guentheri

Urotheca myersi

Urotheca pachyura

Elapidae (6 species)

Micrurus alleni

Micrurus hippocrepis

Micrurus mosquitensis

Micrurus ruatanus

Micrurus stewarti

Micrurus stuarti

Leptotyphlopidae (3 species)

Epictia ater

Epictia martinezi

Epictia pauldwyeri

Typhlopidae (3 species)

Amerotyphlops costaricensis

Amerotyphlops stadelmani

Typhlops tycherus

Viperidae (16 species)

Agkistrodon howardgloydi

Atropoides indomitus

Atropoides picadoi

Bothriechis guifarroi

Bothriechis lateralis

Bothriechis marchi

Bothriechis nigroviridis

Bothriechis nubestris

Bothriechis supraciliaris

Bothriechis thalassinus

Cerrophidion sasai

Cerrophidion wilsoni

Lachesis melanocephala

Lachesis stenophrys

Porthidium porrasi

Porthidium volcanicum

Testudines (2 species)

Geoemydidae (1 species)

Rhinoclemmys funerea

Kinosternidae (1 species)

Kinosternon angustipons

Mata-Silva et al.

Herpetofaunal Composition by Physiographic Regions

300

254 (41%)

250

200

175 (28%)

150

123 (20%)

117 (19%)

Number of Species

100 94 (1596)

86 (14%)

50 z 41 (7%)

; a _ i

0 = =

CG HN CRP EP YP GH NP CGU GCR cP

Physiographic Region

Fig. 3. Graph indicating the number and percentage of Central American endemic species in each of the 10 physiographic regions

recognized.

Table 5. Distributional summary of herpetofaunal families containing priority level one species in Central America, among the 10

physiographic regions. See Table 4 for explanation of abbreviations.

Famili Number Physiographic Regions

Amphib. Reptile Conserv. 30 January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

Table 5 (continued). Distributional summary of herpetofaunal families containing priority level one species in Central America,

among the 10 physiographic regions. See Table 4 for explanation of abbreviations.

of Species

Ile Ss sles

[Mabuyidae ‘| 3

PPhytlodactyidae | 3

[sphacrodaciyidae [11

[Sphenomorphidae [1

Tcowwbriaae «| is

[Dipsadidae ‘| a

[Leptryphiopidae | 2

Piyphiopidae | 1

iperidae «|

[Subtotals iY

[Kinostemidae [1

[Subtotals fd

Protas =i)

[Sum Totals | 426

is 94.3. Four of the regional values lie above or close to the

mean figure, as follows: western nuclear Central American

highlands (CG; 117), eastern nuclear Central American

highlands (HN; 178), Isthmian Central American highlands

(CRP; 254), Caribbean lowlands of eastern Guatemala

and northern Honduras (GH; 94), and Caribbean lowlands

from Nicaragua to Panama (NP; 123). Given these species

numbers, the five regions with values above or close to the

mean are the most significant for conservation remediation.

The other five of the regional values lie below the mean

figure, as follows: highlands of eastern Panama (EP; 37),

Yucatan Platform (YP; 6), Pacific lowlands from eastern

Chiapas to south-central Guatemala (CGU; 9), Pacific

lowlands from southeastern Guatemala to northwestern

Costa Rica (GCR; 39), and Pacific lowlands from Central

Costa Rica through Panama (CP; 86). Even though these

values are relatively low, collectively they amount to 177

species, 28.4% of the total of 623 endemic species; thus,

they also are of considerable importance.

The five regions containing the highest numbers of

endemic species include three in highland and two in

lowland areas. The five regions with the lowest numbers

include one in highland and four in lowland areas. The

numbers in the four highland regions range from 37 to 254,

and in the six lowland regions from six to 86.

Obviously, the 623 Central American endemic species

are distributed unevenly throughout the 10 physiographic

regions we recognize. In order to examine their distribution,

we constructed a table indicating the total number of regions

inhabited by the component species (Table 6). The regions,

listed in order of their total number of constituent species,

range from six in the Yucatan Platform to 254 in the Isthmian

Amphib. Reptile Conserv.

Physiographic Regions

Central American highlands. The number of physiographic

regions occupied by these species ranges from one to eight,

and their corresponding number of species also decreases

markedly (Table 6). Thus, 450 species occupy a single

region, with the numbers ranging from one in the Yucatan

Platform to 154 in the Isthmian Central American highlands;

no single-region species are present in the Pacific lowlands

from eastern Chiapas to south-central Guatemala. At the

opposite extreme, a single species (Leptodeira rhombifera)

occupies eight regions, and one species (Hydromorphus

concolor) inhabits seven regions. The single-region species

comprise the most speciose categories for seven of the 10

physiographic regions (Table 6). The three exceptions are

subhumid regions on the Atlantic (Yucatan Platform) and

Pacific versants (Pacific lowlands from eastern Chiapas

to south-central Guatemala and Pacific lowlands from

southeastern Guatemala to northwestern Costa Rica).

The 450 single-region species comprise 72.2% of the

623 Central American endemic species. The 95 two-region

species contribute 15.2% of the total number. Together, the

single-region and two-region species constitute 545 taxa,

87.5% of the total. Thus, only 78 of the remaining species

occupy from three to eight regions. This feature is of

tremendous conservation significance for Central America,

and we review this matter in greater detail below.

Conservation Status of the Endemic Central

American Herpetofauna

In a previous paper on the Mexican endemic herpetofauna

(Johnson et al. 2017), we utilized the Environmental

Vulnerability Score (EVS) system of conservation

January 2019 | Volume 13 | Number 1 | e168

Mata-Silva et al.

Plate 10. Diasporus ventrimaculatus Chaves, Garcia-

Rodriguez, Mora, and Leal, 2009. This frog is a priority one

species “known only from the Valle del Silencio on the Caribbean

versant of the Cordillera de Talamanca, Limon Province, Costa

Rica” (Frost 2018). This individual was observed in Valle del

Silencio, Parque Internacional La Amistad, in the province of

Limon, Costa Rica. Photo by Victor Acosta-Chaves.

a -

Plate 12. /sthmohyla lancasteri (Barbour, 1928). Lancaster’s

Treefrog is a priority two species with an EVS of 14, which

occurs in “the Cordillera de Talamanca of Costa Rica and

western Panama” (Frost 2018). This individual was found in

Guayacan, in the province of Limon, Costa Rica. Photo by

Victor Acosta-Chaves.

Plate 14. Ptychohyla legleri (Taylor, 1958). Legler’s Stream

Frog is a priority one species with an EVS of 14, which is found

on the “Pacific slopes of the Sierra de Talamanca [of] eastern

Costa Rica and western Panama” (Frost 2018). This individual

was located in Alfombra de Pérez Zeledon, in the province of

San José, Costa Rica. Photo by Victor Acosta-Chaves.

Amphib. Reptile Conserv.

toe F ali

Plate 11. Duel/manohyla rufioculis (Taylor, 1952). This treefrog

is a priority one species with an EVS of 14, which ranges on

both “the Caribbean and Pacific slopes of the mountains of

Costa Rica” (Frost 2018). This individual came from Centro

Soltis, San Isidro de Pefias Blancas, in the province of Alajuela,

Costa Rica. Photo by Victor Acosta-Chaves.

Plate 13. Plectrohyla pokomchi Duellman and Campbell, 1984.

The Rio Sanaja Spikethumb Frog 1s a priority eight species with an

EVS of 13, which is distributed in “the Sierra de las Minas and the

contiguous Sierra de Xucaneb in central and eastern Guatemala”

(Frost 2018). This individual came from Purulha, in the department

of Baja Verapaz, Guatemala. Photo by Andres Novales.

a i Wee,

“ > . oe ill 7 i eo

Plate 15. Smilisca puma (Cope, 1885). The Tawny Smilisca is

a priority one species with an EVS of 14, distributed on the

“Caribbean lowlands of Costa Rica and adjacent Nicaragua”

(Frost 2018). This individual was encountered in La Selva

Biological Station, in the province of Heredia, Costa Rica.

Photo by Victor Acosta-Chaves.

January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

Table 6. Number of endemic species in each of the 10 physiographic regions inhabited in Central America. See Table 4 for

explanation of abbreviations.

Physiographic

Regions

Totals

assessment. Along with various other authors, we have

been involved with a series of papers published on the

Mesoamerican herpetofauna since 2013 (see Johnson

et al. 2017, for a listing) including a recent paper on the

herpetofauna of the Mexican state of Puebla (Woolrich-Pifia

et al. 2017). Herein, we use the same system to evaluate

the conservation status of the 623 species comprising the

Central American herpetofauna. In calculating the EVS for

these species, we used the scores included in Johnson et al.

(2015), supplemented by the scores we determined for the

43 species described since this paper was published. We

placed these scores in Table 7, incorporated them into those

for the entire Central American endemic herpetofauna in

Table 8, and provide a graph of the data in Fig. 4.

To illustrate the pattern of distribution of the EVS, we

organized these scores by family in Table 9. The data in this

table indicate that the scores range from 10 to 20, out of a

total theoretical range of 3 to 20. Thus, the scores occupy

the entire range of medium vulnerability (10-13) and high

vulnerability (14—20) in the EVS scale. None of the scores

for these endemic species extend into the low vulnerability

range (3-9).

The highest score of 20 is found only among the anurans

and, in particular, within the family Hylidae. This score

is shared by six hylid species, including four species of

Ecnomiohyla, one of Bromeliohyla, and one of Ptychohyla

(Table 8). The lowest score of 10 is seen in a broader

range of herpetofaunal families (Table 9), including the

Hylidae (one species), Ranidae (one), Dactyloidae (one),

Phrynosomatidae (one), and Leptotyphlopidae (one). The

greatest number of species, 1e., 146, were assessed an

EVS of 16, with species numbers decreasing more or less

gradually on either side of this apex to both extremes, L.e.,

10 and 20.

Of the 623 total scores, 63 (10.1%) lie within the medium

range and the remaining 560 (89.9%) in the high range

(Table 9). This large representation of high vulnerability

Species among the endemic species is of tremendous

conservation significance, and figures prominently tn the

system of prioritization we present below.

Amphib. Reptile Conserv.

Number of endemic species in each physiographic region

Total

iil: ic

= 6

—

(oo

—

Priority Listing for Central American Endemic

Herpetofaunal Species

In prioritizing the conservation significance of the endemic

herpetofaunal species in Central America, we used the

same simple system developed by Johnson et al. (2017).

This system involves combining the data on physiographic

distribution (Table 4) and the Environmental Vulnerability

Scores (Table 8) for the 623 endemic species. This procedure

resulted in the recognition of 14 priority levels, of which six

are high vulnerability and eight are medium vulnerability

groupings (Table 10).

We organized the high vulnerability species into six

groups based on the number of physiographic regions they

occupy, ranging from one to six (Table 10, Fig. 5). The

numbers of species in these seven groups decrease markedly

and consistently, as follows: Priority Level One (429

species); Priority Level Two (73); Priority Level Three (27);

Priority Level Four (21); Priority Level Five (nine); and

Priority Level Six (three). The most significant conclusion

of this study 1s that 562 (90.2%) of the endemic species in

Central America are allocated to the six high vulnerability

groups. This proportion is 10 percentage points higher than

the comparable figure (80.2%) for the Mexican endemic

species (Johnson et al., 2017). Furthermore, we believe

that the difficulty of protecting these high vulnerability

species increases with the fewer physiographic regions they

occupy. Thus, the most critically vulnerable species are in

the Priority One grouping, the 429 species that constitute

68.9% of the total number of Central American endemics.

The challenge of protecting the high vulnerability species

increases commensurately with the decrease in the priority

level number.

We arranged the medium vulnerability species into eight

groups, also on the basis of the number of physiographic

regions inhabited (Table 10, Fig. 5). Fewer species are

included in these eight groups compared to the high

vulnerability ones, as follows: Priority Level Seven (23);

Priority Level Eight (21 species); Priority Level Nine (5);

Priority Level Ten (four); Priority Level Eleven (four);

January 2019 | Volume 13 | Number 1 | e168

Mata-Silva et al.

Table 7. Environmental Vulnerability Scores (EVS) for 43 endemic members of the Central American herpetofauna not included

in Johnson et al. (2015) or requiring recalculation. Question marks indicate decisions made about reproductive mode based on

phylogenetic relationships.

Rivonia Veiner Score VS)

Incilius majordomus

Hyalinobatrachium dianae

Craugastor gabbi

Diasporus dariensis

Diasporus majeensis

Diasporus pequeno

Diasporus sapo

Plectrohyla calvata

Smilisca manisorum

Lithobates lenca

Bolitoglossa aurae

Bolitoglossa chucutaniensis

Cryptotriton xucaneborum

Nototriton costaricense

Nototriton nelson

Nototriton oreadorum

Oedipina berlini

Oedipina capitalina

Oedipina salvadorensis

Celestus laf

Mesaspis cuchumatanus

Mesaspis salvadorensis

Dactyloa brooksi

Dactyloa kathydayae

Dactyloa maia

Dactyloa savage

Norops elcopeensis

Norops mccraniei

Norops oxylophus

Norops wilsoni

Lepidoblepharis emberawoundule

Lepidoblepharis rufigularis

Lepidoblepharis victormartinezi

Ameiva fuliginosa

Holcosus miadis

Tantilla berguidoi

Tantilla excelsa

Tantilla gottei

Tantilla stenigrammi

Rhadinella lisyae

Epictia martinezi

Epictia pauldwyeri

Ana nrnnuna»aunrin WN aan annaa»anaaanananaannrnanrnnndndnDDHDD HD UNA ADHDNHD WDA aN

YN wWowonwoaonavrvriwnwaonwaonnanenNinNnNrNr waNNN WOT WAWAAHAN DAHA DAHADHDOHOAHAH WH NY

NA == re NN NY NY NY WW WW WW WW WW WwW WwW WwW WwW WwW Ww W W

Bothriechis nubestris

Amphib. Reptile Conserv. 34 January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

Plate 16. Agalychnis annae (Duellman, 1963). The Orange-eyed Treefrog is a priority two species with an EVS of 15, with a distribution

in the “Northern Cordillera de Talamanca, Cordillera de Tilaran and Cordillera Central of Costa Rica” (Frost 2018). This individual was

found in Heredia, Costa Rica. Photo by Victor Acosta-Chaves.

Plate 17. Agalychnis saltator Taylor, 1955. This leaf frog is a priority three species with an EVS of 14, which ranges along the “Caribbean

lowlands of northeastern Honduras, Nicaragua, to east-central Costa Rica” (Frost 2018). This individual was located in Centro Soltis, San

Isidro de Pefias Blancas, in the province of Alajuela, Costa Rica. Photo by Victor Acosta-Chaves.

Plate 18. Lithobates ee (Smith, 1959), The Peralta Tees isa aan, ra species with an EVS of 12, distributed “at scattered localities

on the humid Atlantic lowlands from eastern Nicaragua to southeastern Costa Rica and in the humid premontane and lower montane areas

of upland Costa Rica, including the Meseta Oriental and Meseta Occidental and probably the Cordillera Central” (Savage 2002: 402). This

individual was found in a pond at Llano Tugri, in the Serrania de Tabasara. Photo by Abel Batista.

Amphib. Reptile Conserv. 35 January 2019 | Volume 13 | Number 1 | e168

Mata-Silva et al.

Table 8. Environmental Vulnerability Scores (EVS) for the endemic members of the herpetofauna of Central America.

Atelopus certus

Atelopus chiriquiensis

Atelopus chirripoensis

Atelopus limosus

Atelopus senex

Atelopus varius

Atelopus zeteki

Incilius aucoinae

Incilius chompipe

Incilius epioticus

Incilius fastidiosus

Incilius guanacaste

Incilius holdridgei

Incilius ibarrai

Incilius karenlipsae

Incilius leucomyos

Incilius majordomus

Incilius melanochlorus

Incilius periglenes

Incilius peripatetes

Incilius porter

Incilius signifier

Rhinella centralis

Rhinella chrysophora

Cochranella granulosa

Hyalinobatrachium dianae

Hyalinobatrachium talamancae

Hyalinobatrachium vireovittatum

Craugastor adamastus

Craugastor anciano

Craugastor andi

Craugastor angelicus

Craugastor aphanus

Craugastor aurilegulus

Craugastor azueroensis

Craugastor bocourti

Craugastor bransfordii

Craugastor campbelli

Craugastor catalinae

Craugastor chac

Craugastor charadra

Craugastor chingopetaca

Craugastor chrysozetetes

Craugastor coffeus

Craugastor cruzi

Amphib. Reptile Conserv.

Oedipina ignea

Oedipina kasios

Oedipina koehleri

Oedipina leptopoda

Oedipina maritima

Oedipina motaguae

Oedipina nica

Oedipina nimaso

Oedipina pacificensis

Oedipina paucidentata

Oedipina petiola

Oedipina poelzi

Oedipina pseudouniformis

Oedipina quadra

Oedipina salvadorensis

Oedipina savagei

Oedipina stenopodia

Oedipina stuarti

Oedipina taylori

Oedipina tomasi

Oedipina tzutujilorum

Oedipina uniformis

Pseudoeurycea exspectata

Caecilia volcani

Oscaecilia elongata

Oscaecilia osae

Dermophis costaricensis

Dermophis gracilior

Dermophis occidentalis

Gymnopis multiplicata

Abronia anzuetoi

Abronia aurita

Abronia campbelli

Abronia fimbriata

Abronia frosti

Abronia gaiophantasma

Abronia meledona

Abronia montecristoi

Abronia salvadorensis

Abronia vasconcelosii

Celestus adercus

Celestus atitlanensis

Celestus bivittatus

Celestus cyanochloris

Celestus hylaius

36 January 2019 | Volume 13 | Number 1 | e168

Table 8 (continued). Environmental Vulnerability Scores (EVS) for the endemic members of the herpetofauna of Central America.

Craugastor cuaquero

Craugastor cyanochthebius

Craugastor daryi

Craugastor emcelae

Craugastor emleni

Craugastor epochthidius

Craugastor escoces

Craugastor evanesce

Craugastor fecundus

Craugastor fleischmanni

Craugastor gabbi

Craugastor gollmeri

Craugastor gulosus

Craugastor inachus

Craugastor jota

Craugastor laevissimus

Craugastor lauraster

Craugastor megacephalus

Craugastor melanostictus

Craugastor merendonensis

Craugastor milesi

Craugastor mimus

Craugastor monnichorum

Craugastor myllomyllon

Craugastor nefrens

Craugastor noblei

Craugastor obesus

Craugastor olanchano

Craugastor omoaensis

Craugastor pechorum

Craugastor persimilis

Craugastor phasma

Craugastor podiciferus

Craugastor polyptychus

Craugastor punctariolus

Craugastor ranoides

Craugastor rayo

Craugastor rhyacobatrachus

Craugastor rivulus

Craugastor rostralis

Craugastor rugosus

Craugastor sabrinus

Craugastor saltuarius

Craugastor sandersoni

Craugastor stadelmani

Amphib. Reptile Conserv.

The endemic herpetofauna of Central America

Celestus laf

Celestus montanus

Celestus orobius

Celestus scansorius

Coloptychon rhombifer

Diploglossus bilobatus

Diploglossus montisilvestris

Mesaspis cuchumatanus

Mesaspis monticola

Mesaspis salvadorensis

Basiliscus plumifrons

Dactyloa brooksi

Dactyloa casildae

Dactyloa ginaelisae

Dactyloa ibanezi

Dactyloa insignis

Dactyloa kathydayae

Dactyloa kunayalae

Dactyloa maia

Dactyloa microtus

Dactyloa savagei

Norops alocomyos

Norops altae

Norops amplisquamosus

Norops apletophallus

Norops aquaticus

Norops benedikti

Norops bicaorum

Norops campbelli

Norops carpenteri

Norops charlesmyersi

Norops cobanensis

Norops cryptolimifrons

Norops cupreus

Norops cusuco

Norops datzorum

Norops elcopeensis

Norops fortunensis

Norops fungosus

Norops gruuo

Norops haguei

Norops heteropholidotus

Norops humilis

Norops intermedius

Norops johnmeyeri

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Table 8 (continued). Environmental —_— Scores (EVS) for the endemic members of the herpetofauna of Central America.

Craugastor stejnegerianus

Craugastor tabasarae

Craugastor talamancae

Craugastor taurus

Craugastor trachydermus

Craugastor underwoodi

Craugastor xucanebi

Pristimantis adnus

Pristimantis altae

Pristimantis caryophyllaceus

Pristimantis cerasinus

Pristimantis museosus

Pristimantis pardalis

Pristimantis pirrensis

Strabomantis laticorpus

Ameerega maculate

Andinobates claudiae

Andinobates geminisae

Colostethus latinasus

Ectopoglossus astralogaster

Ectopoglossus isthminus

Oophaga arborea

Oophaga granulifera

Oophaga pumilio

Oophaga speciosa

Oophaga vicentei

Phyllobates lugubris

Phyllobates vittatus

Silverstoneia flotator

Diasporus citrinobapheus

Diasporus darienensis

Diasporus diastema

Diasporus hylaeformis

Diasporus igneus

Diasporus majeensis

Diasporus pequeno

Diasporus sapo

Diasporus tigrillo

Diasporus ventrimaculatus

Atlantihyla panchoi

Atlantihyla spinipollex

Bromeliohyla melacaena

Dryophytes bocourti

Duellmanohyla legleri

Duellmanohyla lythrodes

Duellmanohyla rufioculis

Amphib. Reptile Conserv.

Mata-Silva et al.

Norops kemptoni

Norops kreutzi

Norops leditzigorum

Norops limifrons

Norops lionotus

Norops loveridgei

Norops macrophallus

Norops magnaphallus

Norops marsupialis

Norops mcecraniei

Norops monteverde

Norops morazani

Norops muralla

Norops ocelloscapularis

Norops osa

Norops oxylophus

Norops pachypus

Norops pijolensis

Norops polylepis

Norops pseudokemptoni

Norops pseudopachypus

Norops purpurgularis

Norops quaggulus

Norops roatanensis

Norops rubribarbaris

Norops salvini

Norops sminthus

Norops tenorioensis

Norops townsendi

Norops triumphalis

Norops tropidolepis

Norops utilensis

Norops villai

Norops wampuensis

Norops wellbornae

Norops wermuthi

Norops wilsoni

Norops woodi

Norops yoroensis

Norops zeus

Coleonyx mitratus

Bachia blairi

Echinosaura panamensis

Echinosaura apodema

Heloderma charlesbogerti

Ctenosaura bakeri

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The endemic herpetofauna of Central America

Table 8 (continued). Environmental Vulnerability Scores (EVS) for the endemic members of the herpetofauna of Central America.

Duellmanohyla salvadorensis Ctenosaura flavidorsalis

Duellmanohyla salvavida

Duellmanohyla soralia

Duellmanohyla uranochroa

Ecnomiohyla bailarina

Ecnomiohyla fimbrimembra

Ecnomiohyla minera

Ecnomiohyla rabborum

Ecnomiohyla salvaje

Ecnomiohyla sukia

Ecnomiohyla thysanota

Ecnomiohyla veraguensis

Exerodonta catracha

Exerodonta perkinsi

HAyloscirtus colymba

Isthmohyla angustilineata

Isthmohyla calypso

Isthmohyla debilis

Isthmohyla graceae

Isthmohyla infucata

Isthmohyla insolita

Isthmohyla lancasteri

Isthmohyla picadoi

Isthmohyla pictipes

Isthmohyla pseudopuma

Isthmohyla rivularis

Isthmohyla tica

Isthmohyla xanthosticta

Isthmohyla zeteki

Plectrohyla calvata

Plectrohyla chrysopleura

Plectrohyla dasypus

Plectrohyla exquisita

Plectrohyla glandulosa

Plectrohyla pokomchi

Plectrohyla psiloderma

Plectrohyla quecchi

Plectrohyla tecunumani

Plectrohyla teuchestes

Ptychohyla dendrophasma

Ptychohyla hypomykter

QOuilticohyla sanctaecrucis

Scinax altae

Smilisca manisorum

Smilisca puma

Leptodactylus silvanimbus

Amphib. Reptile Conserv.

Ctenosaura melanosterna

Ctenosaura oedirhina

Ctenosaura palearis

Ctenosura praeocularis

Ctenosaura quinquecarinata

Marisora alliacea

Marisora magnacornae

Marisora roatanae

Marisora unimarginata

Sceloporus lunaei

Sceloporus malachiticus

Phyllodactylus insularis

Phyllodactylus palmeus

Phyllodactylus paralepis

Mesoscincus managuae

Lepidoblepharis emberawoundule

Lepidoblepharis rufigularis

Lepidoblepharis victormartinezi

Sphaerodactylus alphus

Sphaerodactylus dunni

Sphaerodactylus graptolaemus

Sphaerodactylus guanaje

Sphaerodactylus homolepis

Sphaerodactylus leonardovaldesi

Sphaerodactylus millepunctatus

Sphaerodactylus pacificus

Sphaerodactylus poindexteri

Sphaerodactylus rosaurae

Scincella rara

Cnemidophorus duellmani

Cnemidophorus ruatanus

Holcosus leptophrys

Holcosus miadis

Holcosus quadrilineatus

Lepidophyma mayae

Lepidophyma reticulatum

Helminthophis frontalis

Dendrophidion apharocybe

Dendrophidion crybelum

Dendrophidion paucicarinatum

Dendrophidion rufiterminorum

Drymobius melanotropis

Leptodrymus pulcherrimus

Leptophis nebulosus

Mastigodryas alternatus

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Mata-Silva et al.

Table 8 (continued). Environmental —— Scores (EVS) for the endemic members of the herpetofauna of Central America.

Hypopachus pictiventris

Agalychnis annae

Agalychnis saltatory

Pipa myersi

Lithobates juliani

Lithobates lenca

Lithobates miadis

Lithobates taylori

Lithobates vibicarius

Lithobates warszewitschii

Bolitoglossa alvaradoi

Bolitoglossa anthracina

Bolitoglossa aurae

Bolitoglossa aureogularis

Bolitoglossa bramei

Bolitoglossa carri

Bolitoglossa cataguana

Bolitoglossa celaque

Bolitoglossa centenorum

Bolitoglossa cerroensis

Bolitoglossa chucantiensis

Bolitoglossa colonnea

Bolitoglossa compacta

Bolitoglossa conanti

Bolitoglossa copia

Bolitoglossa cuchumatana

Bolitoglossa cuna

Bolitoglossa daryorum

Bolitoglossa decora

Bolitoglossa diaphora

Bolitoglossa diminuta

Bolitoglossa dofleini

Bolitoglossa dunni

Bolitoglossa epimela

Bolitoglossa eremia

Bolitoglossa gomezi

Bolitoglossa gracilis

Bolitoglossa heiroreias

Bolitoglossa helmrichi

Bolitoglossa huehuetenanguensis

Bolitoglossa indio

Bolitoglossa insularis

Bolitoglossa jacksoni

Bolitoglossa jugivagans

Bolitoglossa kamuk

Amphib. Reptile Conserv.

Mastigodryas dorsalis

Oxybelis wilsoni

Scolecophis atrocinctus

Tantilla albiceps

Tantilla armillata

Tantilla bairdi

Tantilla berguidoi

Tantilla brevicauda

Tantilla excelsa

Tantilla gottei

Tantilla hendersoni

Tantilla jani

Tantilla lempira

Tantilla olympia

Tantilla psittaca

Tantilla ruficeps

Tantilla stenigrammi

Tantilla taeniata

Tantilla tecta

Tantilla tritaeniata

Tantilla vermiformis

Trimorphodon quadruplex

Adelphicos daryi

Adelphicos ibarrorum

Adelphicos veraepacis

Atractus darienensis

Atractus depressiocellus

Atractus hostilitractus

Atractus imperfectus

Chapinophis xanthocheilus

Coniophanes joanae

Crisantophis nevermanni

Cubophis brooksi

Dipsas articulata

Dipsas bicolor

Dipsas nicholsi

Dipsas tenuissima

Enulius bifoveatus

Enulius roatanensis

Geophis bellus

Geophis brachycephalus

Geophis championi

Geophis damiani

Geophis downsi

Geophis dunni

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The endemic herpetofauna of Central America

Table 8 (continued). Environmental Vulnerability Scores (EVS) for the endemic members of the herpetofauna of Central America.

Bolitoglossa kaqchikelorum

Bolitoglossa la

Bolitoglossa lignicolor

Bolitoglossa longissima

Bolitoglossa magnifica

Bolitoglossa marmorea

Bolitoglossa meliana

Bolitoglossa minutula

Bolitoglossa mombachoensis

Bolitoglossa morio

Bolitoglossa nigrescens

Bolitoglossa ninadormida

Bolitoglossa nussbaumi

Bolitoglossa nvmpha

Bolitoglossa obscura

Bolitoglossa odonnelli

Bolitoglossa omniumsanctorum

Bolitoglossa oresbia

Bolitoglossa pacaya

Bolitoglossa pesrubra

Bolitoglossa porrasorum

Bolitoglossa psephena

Bolitoglossa pygmaea

Bolitoglossa robinsoni

Bolitoglossa robusta

Bolitoglossa salvinii

Bolitoglossa schizodactyla

Bolitoglossa sombra

Bolitoglossa sooyorum

Bolitoglossa splendida

Bolitoglossa striatula

Bolitoglossa subpalmata

Bolitoglossa suchitanensis

Bolitoglossa synoria

Bolitoglossa taylori

Bolitoglossa tenebrosa

Bolitoglossa tica

Bolitoglossa tzultacaj

Bolitoglossa xibalba

Bolitoglossa zacapensis

Cryptotriton monzoni

Cryptotriton nasalis

Cryptotriton necopinus

Cryptotriton sierraminensis

Cryptotriton veraepacis

Amphib. Reptile Conserv.

Geophis fulvoguttatus

Geophis godmani

Geophis hoffmanni

Geophis nephodrymus

Geophis ruthveni

Geophis talamancae

Geophis tectus

Geophis zeledoni

Hydromorphus concolor

Hydromorphus dunni

Imantodes phantasma

Leptodeira rhombifera

Leptodeira rubricata

Ninia celata

Ninia espinali

Ninia maculata

Ninia pavimentata

Ninia psephota

Omoadiphas aurula

Omoadiphas cannula

Omoadiphas texiguatensis

Rhadinaea calligaster

Rhadinaea pulveriventris

Rhadinaea sargenti

Rhadinaea stadelmani

Rhadinaea vermiculaticeps

Rhadinella anachoreta

Rhadinella hempsteadae

Rhadinella lisyae

Rhadinella montecristi

Rhadinella pegosalyta

Rhadinella pilonaorum

Rhadinella rogerromani

Rhadinella serperaster

Rhadinella tolpanorum

Sibon anthracops

Sibon argus

Sibon carri

Sibon lamari

Sibon longifrenis

Sibon manzanaresi

Sibon merendonensis

Sibon miskitus

Sibon noalamina

Sibon perissostichon

41 January 2019 | Volume 13 | Number 1 | e168

Table 8 (continued). Environmental Vulnerability Scores (EVS) for the endemic members of the herpetofauna of Central America.

Cryptotriton xucaneborum

Dendrotriton bromeliacius

Dendrotriton chujorum

Dendrotriton cuchumatanus

Dendrotriton kekchiorum

Dendrotriton rabbi

Dendrotriton sanctibarbarus

Nototriton abscondens

Nototriton barbouri

Nototriton brodiei

Nototriton costaricense

Nototriton gamezi

Nototriton guanacaste

Nototriton lignicola

Nototriton limnospectator

Nototriton major

Nototriton matama

Nototriton mime

Nototriton nelsoni

Nototriton oreadorum

Nototriton picadoi

Nototriton picucha

Nototriton richardi

Nototriton saslaya

Nototriton stuarti

Nototriton tapanti

Nototriton tomamorum

Oedipina alfaroi

Oedipina alleni

Oedipina altura

Oedipina berlini

Oedipina capitalina

Oedipina carablanca

Oedipina chortiorum

Oedipina collaris

Oedipina cyclocauda

Oedipina fortunensis

Oedipina gephyra

Oedipina gracilis

Oedipina grandis

Amphib. Reptile Conserv.

Mata-Silva et al.

Trimetopon barbouri

Trimetopon gracile

Trimetopon pliolepis

Trimetopon simile

Trimetopon slevini

Trimetopon viquezi

Urotheca guentheri

Urotheca myersi

Urotheca pachyura

Micrurus alleni

Micrurus hippocrepis

Micrurus mosquitensis

Micrurus ruatanus

Micrurus stewarti

Micrurus stuarti

Epictia ater

Epictia martinezi

Epictia pauldwyeri

Amerotyphlops costaricensis

Amerotyphlops stadelmani

Typhlops tycherus

Agkistrodon howardgloydi

Atropoides indomitus

Atropoides picadoi

Bothriechis guifarroi

Bothriechis lateralis

Bothriechis marchi

Bothriechis nigroviridis

Bothriechis nubestris

Bothriechis supraciliaris

Bothriechis thalassinus

Cerrophidion sasai

Cerrophidion wilsoni

Lachesis melanocephala

Lachesis stenophrys

Porthidium porrasi

Porthidium volcanicum

Rhinoclemmys funerea

Kinosternon angustipons

42 January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

EVS

160

146

140

ido 118

109

wo 97

2 100

S 80 74

iD]

5 60

40 33

5 4 6

o = = oy ra]

oO oO & oO o oO oO & > oO

Aw . ow £ & & ee ee ou ar er &

« =

Score

Fig. 4. Graph showing Central American endemic species and their corresponding Environmental Vulnerability Scores (EVS)

ranging from 10 to 20.

Conservation Priority

500

450 429

Number of Species

MN MN

150

100

27 21 c 23 21

3 5 4 4 ? 1 1

@ ,o @ RS @ b ens RS @ S o ae & s

eo « a S & 9 Ef Ke R s es é

Priority Level

Fig. 5. Graph of Central American endemic species allocated to the 14 conservation priority groups.

Amphib. Reptile Conserv. 43 January 2019 | Volume 13 | Number 1 | e168

Mata-Silva et al.

Priority Level Twelve (two); Priority Level Thirteen (one);

and Priority Level Fourteen (one). Even so, the next most

important conclusion of this study is that these 61 species

make up 9.8% of the total compendium of endemic species

in Central America. The number of species in these eight

groups also decreases sharply, as 69.8% of the 61 species

fall into the first two priority levels, 1.e., Seven and Eight.

When we examined the 623 endemic species relative to

the number of physiographic regions inhabited, the results

are as follows: one region (429+23 = 452); two regions

(73+21 = 94); three regions (27+5 = 32); four regions (21+4

= 25); five regions (9+4 = 13); six regions (3+2 = 5); seven

regions (1); and eight regions (1). Perusal of these data

supports another conclusion, 1.e., that 72.6% of the total

number of species occupy a single physiographic region.

Based on the assumptions of this study, these 452 species

can be expected to offer the major challenge in efforts to

protect the endemic component of the Central American

herpetofauna. The next most challenging group contains

the 94 species occupying two regions. Together, the single-

group and double-group species comprise 546 (87.6%) of

the total of 623 Central American endemic species.

Our analysis in this paper indicates that most of the 623

endemic Central American herpetofaunal species are judged

as high vulnerability based on the EVS methodology, and

are demonstrated to occupy relatively few physiographic

regions (one or two). The endemic component of the

Central American herpetofauna, just as with the Mexican

endemic component (Johnson et al. 2017), is of global

significance and constitutes the most significant challenge

to conservation professionals working within this segment

of the Mesoamerican herpetofauna. Johnson et al. (2017)

arrived at the same conclusion in their work on the

Mexican endemic herpetofauna. Considered as a whole, the

Mesoamerican endemic herpetofauna comprises the 789

Mexican endemic species dealt with by Johnson et al. (2017)

and the 623 Central American endemic species dealt with

here, as well as the 225 species restricted in distribution to

Mexico and Central America (1.e., Mesoamerica; Wilson et

al., 2017) for a total of 1,637 species. This figure represents

more than three quarters of the entire Mesoamerican

herpetofauna (Wilson et al., 2017). We examine the

parameters of the challenge facing conservation biologists

working in Central America in the following section.

Prognosis for the Endemic Central American

Herpetofauna

The same environmental issues impacting the Mexican

endemic herpetofauna, as discussed by Johnson et al.

(2017), also impinge upon the Central American endemic

herpetofauna. In light of this situation, we emphasize that

the survival of the 623 endemic species inhabiting Central

America ultimately depends on addressing the underlying

issues that lead to all environmental problems, including

biodiversity decline, that in turn stand in the way of

designing a sustainable existence for humanity’s tenure on

Amphib. Reptile Conserv.

Earth. Johnson et al. (2017: 609) explained what we face

as follows: “Fundamentally, humans have created and

maintain these environmental problems because of their

capacity for rational thought, i.e., their ability to connect

cause to effect through the passing of time, and adopting

an anthropocentric worldview that stresses the exploitation

of the world’s resources to support the burgeoning human

population. Such a worldview contrasts markedly with that

of environmentalists, who have adopted ‘a worldview that

helps us make sense of how the environment works, our

place in the environment, and right and wrong environmental

behaviors’ (Raven and Berg 2004: G-6). Obviously, the

present anthropocentric worldview held by most people

represents the fundamental reason why these environmental

problems exist, and continued human population growth

allows them to worsen over time.”

The anthropocentric worldview, also known as the

Western worldview, “includes human superiority and

dominance over nature, the unrestricted use of natural

resources, increased economic growth to manage an

expanding industrial base, the inherent rights of individuals,

and accumulation of wealth and unlimited consumption of

goods and services to provide material comforts” (Raven

and Berg 2004: 17). This worldview not only creates the

entire spectrum of environmental problems, but also the

entire panoply of human societal issues we see played

out every day in various media outlets. Ultimately, they

arise from a commitment to discriminate among groups

of people, i.e., on the basis of racial background, gender,

religion, economic wealth, political persuasion, and so

forth. Thus, not only is humanity poised against the rest

of the living world, but also varying groups of humans

are in conflict with one another. As the focus of humanity

decreases from larger to increasingly smaller realms of

interest, it can be argued that mental stability gives way

to instability, and eventually gives rise to the increased

incidence of the narcissistic personality disorder (NPD).

This disorder is highly variable in presentation and can

manifest across a broad spectrum of severity, but is generally

characterized by pervasive grandiosity, an excessive

need for admiration, and a lack of empathy (Caligor et al.

2015). Envisioning NPD as an extreme end-point of the

intensification of anthropocentrism might explain why

the potential causes of this disorder remain unknown and

that clinical guidelines have yet to emerge (Caligor et al.

2015). Given that none of the authors of this paper possesses

credentials in psychology or psychiatry, our idea about the

connection between the anthropocentric worldview and the

narcissistic personality disorder can be best understood as

a hypothesis remaining to be tested, hopefully by a cross-

discipline team of environmental scientists, deep ecology

philosophers, and biocentric psychologists/psychiatrists.

Studying such a connection could lie within the realm of

environmental psychology, defined as an interdisciplinary

field that focuses on the interplay between environments

and human cognition and behavior; considering the term

“environment” broadly, including both natural and human-

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The endemic herpetofauna of Central America

Table 9. Summary of EVS values for Central American endemic species, arranged by family. Shaded area encompasses high

vulnerability scores.

Number Environmental Vulnerability Scores

Bufonidae — 1 2 6 9 4 1 1 — — _-

Centrolenidae

Craugastoridae

Dendrobatidae

Eleutherodactyludae

Hylidae

Leptodactylidae

Microhylidae

Phyllomedusidae

Pipidae

Ranidae

Subtotals

Plethodontidae

Subtotals

Caeciliidae

Dermophiidae

Subtotals

Totals

Anguidae

Corytophanidae

Dactyloidae

Eublepharidae

Gymnophthalmidae

Helodermatidae

Iguanidae

Mabuyidae

Phrynosomatidae

Phyllodactylidae

Scincidae

Sphaerodactylidae

Sphenomorphidae

ep WN BN RK WD

—

ies)

Telidae

Xantusiidae

Subtotals

Anomalepididae

Colubridae

Dipsadidae

Elapidae

Leptotyphlopidae

Typhlopidae

Viperidae

Subtotals

Geoemydidae

Amphib. Reptile Conserv. 45 January 2019 | Volume 13 | Number 1 | e168

Mata-Silva et al.

Table 9 (continued). Summary of EVS values for Central American endemic species, arranged by family. Shaded area encompasses

high vulnerability scores.

Number

Families of

Species

Environmental Vulnerability Scores

ele fepe[«[«[«[r[™[o[>

Kinosternidae

Subtotals

Totals

Sum Totals

Category Totals

Plate 19. Lithobates warszewitschii (Schmidt, 1857). Warszewitsch’s Frog is a priority eleven species with an EVS of 10, found

on “the Atlantic versant from northeastern Honduras to central Panama, both slopes of the cordilleras of Costa Rica and western

Panama, lowlands of southwestern Costa Rica, and eastern Panama and gallery forests in nonpeninsular northwestern Costa Rica”

(Savage 2002: 405). This individual came from Nectandra Reserve, in the province of Alajuela, Costa Rica. Photo by Sean Michael

Rovito.

> 7"

Ves

Plate 20. Bolitoglossa alvaradoi Taylor, 1954. The Moravia Plate 21. Bolitoglossa centenorum Campbell, Smith, Streicher,

de Chirripd Salamander is a priority two species with an

EVS of 16, distributed on “the Atlantic versant of Costa

Rica” (Frost, 2018). This individual was found in Veragua

Rainforest , in the province of Limon, Costa Rica. Photo by

Victor Acosta-Chaves.

Amphib. Reptile Conserv.

Acevedo, and Brodie, 2010. This salamander is a priority one

species with an EVS of 18, which is known “only from the

type locality in the Sierra Cuchumatanes, Huehuetenango,

Guatemala” (Frost 2018). This individual was encountered at the

type locality, near San Mateo Ixtatan. Photo by Todd Pierson.

January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

made environments (De Young 2013). Since its conception,

research in environmental psychology has often targeted

human attitudes towards the natural environment, and

current trends are now shifting to a focus on sustainable

living in the context of environmental issues (De Young

2013).

Finding lasting solutions to environmental problems

must be based on a realistic, fact-based approach that

evaluates the symptoms of these problems until their

causes are identified (Wilson and McCranie 2004). Often,

the search for an ultimate cause stops when exposed to

the anthropocentric worldview. A worldview, however, 1s

a collection of basic values that “help us to make sense of

the world, understand our place in it, and determine right

and wrong behaviors” (Raven and Berg 2004: G-17). How

the values that characterize the anthropocentric worldview

have arisen through the evolution of human behavior to

become predominant, however, generally has not been

explored. Our working assumption, 1.e., our hypothesis, is

that the ultimate causes are deeply engrained in the origins

of human behavior and have become so pervasive as to

underlie our efforts to understand our world, and our place

in it. Even a discipline called environmental psychology

might not expose the steps in behavioral evolution that

would allow present-day humans to address the malady

known as the anthropocentric worldview. In particular, this

viewpoint is evident when considering that environmental

psychology adopts a broad array of theories, methods,

and interpretations from other disciplines as needed, and

this mosaic approach can make it difficult to understand

the field as a whole and the role it might play in these

societal issues moving forward. An encouraging sign is the

recent emergence of even more specific sub-fields, such as

conservation psychology and ecopsychology, which aim to

provide solutions or interventions for problems specifically

related to conservation of the natural world (Steg and Vlek

2009; De Young 2013).

There is clearly a critical need to develop novel

approaches for studying animal behavior and human

psychology that emphasize reasons why the anthropocentric

worldview has become so predominant, and what needs to

be done to replace it with the environmental worldview. If,

as we hypothesized, there is a psychological connection

among centrist forms of thinking at larger scales (i.e., the

anthropocentric worldview) and those at smaller scales

(1.e., narcissism), then we are faced with an even greater

challenge than commonly is envisioned.

Searching for Ultimate Solutions

Johnson etal. (2017: 613) offered some ideas about searching

for ultimate solutions to the problem of biodiversity decline,

based on opinions promulgated by Wilson and Townsend

(2010), Wilson (2016), and Kopnina (2016), and concluded

as follows: “Our opinion is that humans have the rational

capacity to design a sustainable world through cooperative

action, but our species’ attitudes and actions will have to

Amphib. Reptile Conserv.

change. Our preparedness will have to improve as well.

Such change will have to be based on realistic, fact-based

appraisals of where we are now and where we want to be

in the future. Biologists will have to commit to helping the

rest of us understand why the protection of biodiversity

is critical to enjoying a sustainable world. Cultural

anthropologists also will have to assist humanity at large to

understand why the maintenance of cultural diversity also

is essential to living sustainably. Educational reform will

have to be central to such efforts, to help people learn how

to think and act critically and base decisions on the way

things really are, and not how we might wish them to be by

denying reality. The devotion humans have for structuring

beliefs on the basis of little or no evidence, essentially

reversing the benefit of rationality, will have to surrender

to critical-thinking education established by top-to-bottom

educational reform.”

Critical-thinking educational reform, however, is much

easier to conceive than to bring into reality. A fundamental

question is why such reform has not been undertaken.

This question is not easy to answer, but perhaps the most

fundamental reason is that the educational systems currently

in existence are products of the anthropocentric worldview

and reflect its mindsets. These educational systems also have

developed within the current economic systems responsible

for the huge disparities between the rich and poor, and act to

reinforce these disparities.

Ultimate solutions will emerge only from a clear

understanding of the evolution of human psychology,

as confronted with the problems we face. If not, then the

endemic herpetofauna of Central America, as well as the

remainder of life on Earth, will become casualties of the

biodiversity crisis that eventually will envelop all humanity.

Conclusions and Recommendations

Conclusions

A. As concluded by Johnson et al. (2017), life on Earth exists

as a result of the interplay among the planet’s three abiotic

spheres, the atmosphere, hydrosphere, and lithosphere.

B. Environmental problems and the biodiversity crisis exist

because of the impact of humans on all of the planetary

spheres, including the biosphere, and extend along their

existing energy and materials pathways.

C. The biodiversity crisis impacts all life across the globe,

and all levels of its organization.

D. The endemic component of the Central American

herpetofauna is of global significance, and its importance

increases with the addition of new information. Forty-three

species have been added to this component within the last

two years, bringing the total to 623 species.

E. The percentage of endemism of the Central American

herpetofauna 1s 56.9, compared to 61.1 in Mexico, the other

major portion of Mesoamerica.

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Mata-Silva et al.

Plate 22. Bolitoglossa conanti McCranie and Wilson, 1993.

Conant’s Mushroomtongue Salamander is a priority one species

with an EVS of 16, with a distribution “on the Atlantic and

Pacific versants in western Honduras and eastern Guatemala”

(Townsend and Wilson 2008). This individual came from Aldea

San Joaquin, in the department of Copan, Honduras. Photo by

Sean Michael Rovito.

Plate 24. Bolitoglossa diaphora McCranie and Wilson, 1995.

The Cusuco Salamander is a priority one species with an EVS

of 18, which ranges on “the Atlantic versant in northwestern

Honduras” (Townsend and Wilson 2008). This individual was

found in Parque Nacional Cusuco, in the department of Cortés,

Honduras. Photo by Sean Michael Rovito.

Plate 26. Oedipina koehleri Sunyer, Townsend, Wake, Travers,

Gonzalez, Obando, and Quintana, 2011. This worm salamander

is a priority one species with an EVS of 16, restricted to “three

highland areas in northern Nicaragua” (Frost 2018). This

individual came from Reserva Natural Cerro Musun, in the

department of Matagalpa, Nicaragua. Photo by Javier Sunyer.

Amphib. Reptile Conserv.

Plate 23. Bolitoglossa cuchumatana (Stuart, 1943). The Oak

Forest Salamander is a priority one species with an EVS

of 14, which is found in the “departments of El Quiché and

Huehuetenango in the Sierra de Cuchumatanes, Guatemala”

(Frost, 2018). This individual came from near Laguna Maxbal,

in the department of Huehuetenango, Guatemala. Photo by

Todd Pierson.

Plate 25. Bolitoglossa subpalmata (Boulenger, 1896). This

salamander is a priority one species with an EVS of 15, which

occurs “on both slopes of the Cordillera de Guanacaste,

Cordillera de Tilaran, Cordillera Central, and their outliers in

central to northern Costa Rica” (Frost 2018). This individual

was encountered in Volcan Barva, Parque Nacional Braulio

Carrillo, in the province of, Heredia, Costa Rica. Photo by

Victor Acosta-Chaves.

Plate 27. Coloptychon rhombifer (Peters, 1876). The Isthmian

Alligator Lizard is a priority one species with an EVS of

16, distributed from “southwestern Costa Rica and adjacent

western Panama (Savage, 2002: 533). This individual was

found in San Juan de Rincon, Peninsula de Osa, in the province

of Puntarenas, Costa Rica. Photo by César Barrio-Amoros.

48 January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

Table 10. Conservation priority listing of the endemic herpetofaunal species in Central America based on the EVS categorization

and the range of physiographic occurrence.

Priority One: High Vulnerability Species in Single Physiographic Region (429 species)

Atelopus chiriquiensis

Atelopus chirripoensis

Atelopus limosus

Incilius aucoinae

Incilius epioticus

Incilius guanacaste

Incilius holdridgei

Incilius karenlipsae

Incilius majordomus

Incilius periglenes

Incilius peripatetes

Incilius porteri

Hyalinobatrachium talamancae

Hyalinobatrachium vireovittatum

Craugastor adamastus

Craugastor anciano

Craugastor andi

Craugastor angelicus

Craugastor aphanus

Craugastor azueroensis

Craugastor bocourti

Craugastor catalinae

Craugastor chingopetaca

Craugastor chrysozetetes

Craugastor coffeus

Craugastor cruzi

Craugastor cuaquero

Craugastor cyanochthebius

Craugastor daryi

Craugastor emcelae

Craugastor emleni

Craugastor escoces

Craugastor fleischmanni

Craugastor gabbi

Craugastor gulosus

Craugastor inachus

Craugastor jota

Craugastor melanostictus

Craugastor merendonensis

Craugastor milesi

Craugastor monnichorum

Craugastor myllomyllon

Craugastor nefrens

Craugastor olanchano

Craugastor omoaensis

Amphib. Reptile Conserv.

Nototriton tapanti

Nototriton tomamorum

Oedipina altura

Oedipina berlini

Oedipina capitalina

Oedipina carablanca

Oedipina chortiorum

Oedipina collaris

Oedipina cyclocauda

Oedipina fortunensis

Oedipina gephyra

Oedipina gracilis

Oedipina grandis

Oedipina kasios

Oedipina koehleri

Oedipina leptopoda

Oedipina maritima

Oedipina motaguae

Oedipina nica

Oedipina nimaso

Oedipina pacificensis

Oedipina paucidentata

Oedipina petiola

Oedipina poelzi

Oedipina quadra

Oedipina salvadorensis

Oedipina savagei

Oedipina stenopodia

Oedipina taylori

Oedipina tomasi

Oedipina tzutujilorum

Oedipina uniformis

Pseudoeurycea exspectata

Caecilia volcani

Oscaecilia elongata

Oscaecilia osae

Dermophis costaricensis

Dermophis gracilior

Abronia anzuetoi

Abronia aurita

Abronia campbelli

Abronia fimbriata

Abronia frosti

Abronia gaiophantasma

Abronia meledona

49 January 2019 | Volume 13 | Number 1 | e168

Mata-Silva et al.

Table 10 (continued). Conservation priority listing of the endemic herpetofaunal species in Central America based on the EVS

categorization and the range of physiographic occurrence.

Craugastor phasma

Craugastor podiciferus

Craugastor polyptychus

Craugastor punctariolus

Craugastor rayo

Craugastor rhyacobatrachus

Craugastor rivulus

Craugastor saltuarius

Craugastor stadelmani

Craugastor tabasarae

Craugastor talamancae

Craugastor taurus

Craugastor trachydermus

Craugastor underwoodi

Craugastor xucanebi

Pristimantis adnus

Pristimantis museosus

Pristimantis pirrensis

Strabomantis laticorpus

Ameerega maculata

Andinobates claudiae

Andinobates geminisae

Colostethus latinasus

Ectopoglossus astralogaster

Ectopoglossus isthminus

Oophaga arborea

Oophaga pumilio

Oophaga speciosa

Phyllobates lugubris

Phyllobates vittatus

Diasporus citrinobapheus

Diasporus darienensis

Diasporus hylaeformis

Diasporus igneus

Diasporus majeensis

Diasporus pequeno

Diasporus sapo

Diasporus tigrillo

Diasporus ventrimaculatus

Bromeliohyla melacaena

Dryophytes bocourti

Duellmanohyla legleri

Duellmanohyla lythrodes

Duellmanohyla rufioculis

Ecnomiohyla minera

Ecnomiohyla rabborum

Amphib. Reptile Conserv.

Abronia montecristoi

Abronia salvadorensis

Abronia vasconcelosii

Celestus adercus

Celestus bivittatus

Celestus cyanochloris

Celestus hylaius

Celestus laf

Celestus montanus

Celestus orobius

Celestus scansorius

Coloptychon rhombifer

Diploglossus montisilvestris

Mesaspis cuchumatanus

Mesaspis monticola

Mesaspis salvadorensis

Dactyloa casildae

Dactyloa kathydayae

Dactyloa microtus

Norops alocomyos

Norops altae

Norops amplisquamosus

Norops benedikti

Norops bicaorum

Norops campbelli

Norops cusuco

Norops datzorum

Norops fortunensis

Norops fungosus

Norops gruuo

Norops haguei

Norops heteropholidotus

Norops intermedius

Norops johnmeyeri

Norops kemptoni

Norops kreutzi

Norops leditzigorum

Norops magnaphallus

Norops marsupialis

Norops monteverde

Norops morazani

Norops muralla

Norops ocelloscapularis

Norops osa

Norops pachypus

Norops pijolensis

50 January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

Table 10 (continued). Conservation priority listing of the endemic herpetofaunal species in Central America based on the EVS

categorization and the range of physiographic occurrence.

Ecnomiohyla salvaje

Ecnomiohyla thysanota

Ecnomiohyla veraguensis

Exerodonta catracha

Exerodonta perkinsi

Isthmohyla calypsa

Isthmohyla debilis

Isthmohyla infucata

Isthmohyla insolita

Isthmohyla picadoi

Isthmohyla pictipes

Isthmohyla xanthosticta

Isthmohyla zeteki

Plectrohyla calvata

Plectrohyla dasypus

Plectrohyla exquisita

Plectrohyla psiloderma

Plectrohyla tecunumani

Plectrohyla teuchestes

Ptychohyla dendrophasma

QOuilticohyla sanctaecrucis

Scinax altae

Smilisca manisorum

Smilisca puma

Leptodactylus silvanimbus

Hypopachus pictiventris

Pipa myersi

Lithobates lenca

Lithobates miadis

Lithobates vibicarius

Bolitoglossa anthracina

Bolitoglossa aurae

Bolitoglossa aureogularis

Bolitoglossa bramei

Bolitoglossa carri

Bolitoglossa cataguana

Bolitoglossa celaque

Bolitoglossa centenorum

Bolitoglossa cerroensis

Bolitoglossa chucantiensis

Bolitoglossa compacta

Bolitoglossa conanti

Bolitoglossa copia

Bolitoglossa cuchumatana

Bolitoglossa cuna

Bolitoglossa daryorum

Amphib. Reptile Conserv.

Norops pseudokemptoni

Norops pseudopachypus

Norops purpurgularis

Norops roatanensis

Norops rubribarbaris

Norops salvini

Norops sminthus

Norops tenorioensis

Norops townsendi

Norops triumphalis

Norops tropidolepis

Norops utilensis

Norops villai

Norops wampuensis

Norops wermuthi

Norops woodi

Norops yoroensis

Bachia blairi

Potamites apodemus

Ctenosaura bakeri

Ctenosaura oedirhina

Ctenosaura palearis

Marisora alliacea

Marisora magnacornae

Marisora roatanae

Phyllodactylus insularis

Phyllodactylus palmeus

Phyllodactylus paralepis

Lepidoblepharis rufigularis

Lepidoblepharis victormartinezi

Sphaerodactylus alphus

Sphaerodactylus dunni

Sphaerodactylus graptolaemus

Sphaerodactylus guanaje

Sphaerodactylus homolepis

Sphaerodactylus leonardovaldesi

Sphaerodactylus pacificus

Sphaerodactylus poindexteri

Sphaerodactylus rosaurae

Scincella rara

Cnemidophorus duellmani

Holcosus miadis

Dendrophidion paucicarinatum

Oxybelis wilsoni

Tantilla albiceps

Tantilla bairdi

January 2019 | Volume 13 | Number 1 | e168

Mata-Silva et al.

Table 10 (continued). Conservation priority listing of the endemic herpetofaunal species in Central America based on the EVS

categorization and the range of physiographic occurrence.

Bolitoglossa decora

Bolitoglossa diaphora

Bolitoglossa diminuta

Bolitoglossa dunni

Bolitoglossa epimela

Bolitoglossa eremia

Bolitoglossa gomezi

Bolitoglossa gracilis

Bolitoglossa heiroreias

Bolitoglossa helmrichi

Bolitoglossa huehuetenanguensis

Bolitoglossa indio

Bolitoglossa insularis

Bolitoglossa jacksoni

Bolitoglossa jugivagans

Bolitoglossa kamuk

Bolitoglossa kaqchikelorum

Bolitoglossa la

Bolitoglossa longissima

Bolitoglossa magnifica

Bolitoglossa marmorea

Bolitoglossa meliana

Bolitoglossa minutula

Bolitoglossa mombachoensis

Bolitoglossa nigrescens

Bolitoglossa ninadormida

Bolitoglossa nussbaumi

Bolitoglossa obscura

Bolitoglossa omniumsanctorum

Bolitoglossa oresbia

Bolitoglossa pacaya

Bolitoglossa pesrubra

Bolitoglossa porrasorum

Bolitoglossa psephena

Bolitoglossa pygmaea

Bolitoglossa robinsoni

Bolitoglossa robusta

Bolitoglossa sombra

Bolitoglossa sooyorum

Bolitoglossa splendida

Bolitoglossa subpalmata

Bolitoglossa suchitanensis

Bolitoglossa synoria

Bolitoglossa taylori

Bolitoglossa tenebrosa

Bolitoglossa tica

Amphib. Reptile Conserv.

Tantilla berguidoi

Tantilla gottei

Tantilla hendersoni

Tantilla lempira

Tantilla olympia

Tantilla psittaca

Tantilla stenigrammi

Tantilla tecta

Tantilla tritaeniata

Tantilla vermiformis

Adelphicos daryi

Adelphicos ibarrorum

Adelphis veraepacis

Atractus dariensis

Atractus depressiocellus

Atractus hostilitractus

Atractus imperfectus

Chapinophis xanthocheilus

Coniophanes joanae

Cubophis brooksi

Dipsas nicholsi

Dipsas tenuissima

Enulius bifoveatus

Enulius roatanensis

Geophis bellus

Geophis championi

Geophis damiani

Geophis downsi

Geophis dunni

Geophis fulvoguttatus

Geophis godmani

Geophis nephodrymus

Geophis talamancae

Geophis zeledoni

Hydromorphus dunni

Imantodes phantasma

Leptodeira rubricata

Ninia celata

Ninia espinali

Omoadiphas aurula

Omoadiphas cannula

Omoadiphas texiguatensis

Rhadinaea calligaster

Rhadinaea pulveriventris

Rhadinella lisyae

Rhadinella pegosalyta

52 January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

Table 10 (continued). Conservation priority listing of the endemic herpetofaunal species in Central America based on the EVS

categorization and the range of physiographic occurrence.

Bolitoglossa tzultacaj

Bolitoglossa xibalba

Bolitolossa zacapensis

Cryptotriton monzoni

Cryptotriton necopinus

Cryptotriton sierraminensis

Cryptotriton veraepacis

Cryptotriton xucaneborum

Dendrotriton bromeliacius

Dendrotriton chujorum

Dendrotriton cuchumatanus

Dendrotriton kekchiorum

Dendrotriton rabbi

Dendrotriton sanctibarbarus

Nototriton abscondens

Nototriton barbouri

Nototriton brodiei

Nototriton costaricense

Nototriton gamezi

Nototriton guanacaste

Nototriton lignicola

Nototriton limnospectator

Nototriton major

Nototriton matama

Nototriton mime

Nototriton nelsoni

Nototriton oreadorum

Nototriton picadoi

Nototriton picucha

Nototriton richardi

Nototriton saslaya

Nototriton stuarti

Rhadinella pilonaorum

Rhadinella rogerromani

Rhadinella tolpanorum

Sibon lamari

Sibon manzanaresi

Sibon merendonensis

Sibon miskitus

Sibon noalamina

Sibon perissostichon

Trimetopon gracile

Trimetopon slevini

Trimetopon viquezi

Urotheca myersi

Micrurus mosquitensis

Micrurus ruatanus

Micrurus stuarti

Epictia martinezi

Epictia pauldwyeri

Typhlops tycherus

Atropoides indomitus

Bothriechis guifarroi

Bothriechis lateralis

Bothriechis marchi

Bothriechis nigroviridis

Bothriechis nubestris

Bothriechis thalassinus

Cerrophidion sasai

Cerrophidion wilsoni

Porthidium porrasi

Porthidium volcanicum

Kinosternon angustipons

Priority Two: High Vulnerability Species in Two Physiographic Regions (73)

Atelopus certus

Incilius signifer

Rhinella centralis

Hyalinobatrachium dianae

Craugastor aurilegulus

Craugastor campbelli

Craugastor charadra

Craugastor epochthidius

Craugastor evanesco

Craugastor fecundus

Craugastor lauraster

Craugastor obesus

Craugastor pechorum

Amphib. Reptile Conserv.

Celestus atitlanensis

Dactyloa ibanezi

Dactyloa kunayalae

Dactyloa maia

Dactyloa savagei

Norops aquaticus

Norops carpenteri

Norops charlesmyersi

Norops cryptolimifrons

Norops loveridgei

Norops polylepis

Norops wilsoni

Norops zeus

53 January 2019 | Volume 13 | Number 1 | e168

Mata-Silva et al.

Table 10 (continued). Conservation priority listing of the endemic herpetofaunal species in Central America based on the EVS

categorization and the range of physiographic occurrence.

Craugastor persimilis

Craugastor rostralis

Craugastor rugosus

Craugastor sabrinus

Craugastor sandersoni

Pristimantis altae

Pristimantis caryophyllaceus

Oophaga granulifera

Oophaga vicentei

Ecnomiohyla bailarina

Ecnomiohyla fimbrimembra

Ecnomiohyla sukia

Isthmohyla lancasteri

Agalychnis annae

Bolitoglossa alvaradoi

Bolitoglossa lignicolor

Bolitoglossa nvmpha

Bolitoglossa odonnelli

Bolitoglossa salvinii

Cryptotriton nasalis

Oedipina alfaroi

Oedipina alleni

Oedipina ignea

Oedipina stuarti

Dermophis occidentalis

Heloderma charlesbogerti

Ctenosaura melanosterna

Ctenosaura praeocularis

Ctenosaura quinquecarinata

Sceloporus lunaei

Mesoscincus managuae

Lepidoblepharis emberawoundule

Cnemidophorus ruatanus

Dendrophidion crybelum

Mastigodryas dorsalis

Tantilla jani

Tantilla taeniata

Geophis ruthveni

Ninia pavimentata

Rhadinaea sargenti

Rhadinella montecristi

Urotheca pachyura

Micrurus hippocrepis

Bothriechis supraciliaris

Lachesis melanocephala

Lachesis stenophrys

Rhinoclemmys funerea

Priority Three: High Vulnerability Species in Three Physiographic Regions (27)

Craugastor chac

Craugastor gollmeri

Craugastor stejnegerianus

Pristimantis cerasinus

Pristimantis pardalis

Silverstoneia flotator

Agalychnis saltator

Bolitoglossa schizodactyla

Oedipina pseudouniformis

Diploglossus bilobatus

Dactyloa insignis

Norops lionotus

Norops macrophallus

Norops quaggulus

Echinosaura panamensis

Ctenosaura flavidorsalis

Sceloporus malachiticus

Drymobius melanotropis

Scolecophis atrocinctus

Dipsas articulata

Dipsas bicolor

Rhadinaea vermiculaticeps

Sibon anthracops

Sibon carri

Trimetopon barbouri

Micrurus stewarti

Atropoides picadoi

Priority Four: High Vulnerability Species in Four Physiographic Regions (21)

Craugastor megacephalus

Craugastor mimus

Craugastor ranoides

Diasporus diastema

Bolitoglossa colonnea

Amphib. Reptile Conserv.

Norops wellbornae

Holcosus leptophrys

Holcosus quadralineatus

Leptophis nebulosus

Crisantophis nevermanni

54 January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

Table 10 (continued). Conservation priority listing of the endemic herpetofaunal species in Central America based on the EVS

categorization and the range of physiographic occurrence.

Bolitoglossa dofleini Rhadinella anachoreta

Dactyloa brooksi Sibon argus

Norops apletophallus Sibon longifrenis

Norops elcopeensis Micrurus alleni

Norops humilis Agkistrodon howardgloydi

Norops oxylophus

Priority Five: High Vulnerability Species in Five Physiographic Regions (9)

Cochranella granulosa Coleonyx mitratus

Craugastor noblei Marisora unimarginata

Bolitoglossa striatula Dendrophidion rufiterminorum

Gymnopis multiplicata Trimorphodon quadruplex

Basiliscus plumifrons

Priority Six: High Vulnerability Species in Six Physiographic Regions (3)

Norops limifrons Dendrophidion apharocybe

Sphaerodactylus millepunctatus

Priority Seven: Medium Vulnerability Species in Single Physiographic Region (23)

Atelopus senex Plectrohyla pokomchi

Incilius chompipe Plectrohyla quecchi

Incilius fastidiosus Lithobates juliani

Incilius melanochlorus Lithobates taylori

Rhinella chrysophora Bolitoglossa morio

Isthmohyla angustilineata Dactyloa ginaelisae

Isthmohyla graceae Norops cobanensis

Isthmohyla pseudopuma Ninia psephota

Ithmohyla rivularis Rhadinaea stadelmani

Isthmohyla tica Rhadinella hempsteadae

Plectrohyla chrysopleura Rhadinella serperaster

Plectrohyla glandulosa

Priority Eight: Medium Vulnerability Species in Two Physiographic Regions (21)

Atelopus zeteki Norops mecraniei

Incilius ibarrai Lepidophyma mayae

Incilius leucomyos Lepidophyma reticulatum

Craugastor bransfordii Helminthophis frontalis

Atlantihyla panchoi Tantilla brevicauda

Atlantihyla spinipollex Tantilla excelsa

Duellmanohyla salvadorensis Geophis tectus

Duellmanohyla salvavida Trimetopon pliolepis

Duellmanohyla soralia Trimetopon simile

Duellmanohyla uranochroa Amerotyphlops stadelmani

HAyloscirtus colymba

Priority Nine: Medium Vulnerability Species in Three Physiographic Regions (5)

Atelopus varius Tantilla ruficeps

Craugastor laevissimus Geophis brachycephalus

Ptychohyla hypomykter

Priority Ten: Medium Vulnerability Species in Four Physiographic Regions (4)

Lithobates warszewitschii Tantilla armillata

Amphib. Reptile Conserv. 55 January 2019 | Volume 13 | Number 1 | e168

Mata-Silva et al.

Table 10 (continued). Conservation priority listing of the endemic herpetofaunal species in Central America based on the EVS

categorization and the range of physiographic occurrence.

Leptodrymus pulcherrimus

Amerotyphlops costaricensis

Priority Eleven: Medium Vulnerability Species in Five Physiographic Regions (4)

Mastigodryas alternatus

Ninia maculate

Urotheca guentheri

Epictia ater

Priority Twelve: Medium Vulnerability Species in Six Physiographic Regions (2)

Norops cupreus

Geophis hoffmanni

Priority Thirteen: Medium Vulnerability Species in Seven Physiographic Regions (1)

. RESIS yA ol ieee! . = Sea

Plate 28. Dactyloa insignis (Cope, 1871). The Decorated Anole

is a priority four species with an EVS of 14, with a distribution

restricted to “the Cordillera Tilaran and Cordillera Central of Costa

Rica” (Poe and Ryan 2017: 6). This individual was encountered in

Estacion Pocosol, Bosque Eterno de los Nifios, in the province of

Alajuela, Costa Rica. Photo by Victor Acosta-Chaves.

Plate 30. Norops carpenteri (Echelle, Echelle, and Fitch, 1971).

Carpenter’s Anole is a priority two species with an EVS of 16,

distributed “on the Atlantic versant from northeastern Honduras

to northwestern Panama” (McCranie and Kohler 2015: 45). This

individual was found in Reserva El Copal, Jiménez, Costa Rica.

Photo by Victor Acosta-Chaves.

Amphib. Reptile Conserv.

Plate 29. Dactyloa kunayalae (Huleback, Poe, Ibafiez, and

Williams, 2007). This anole is a priority two species with an EVS

of 15, restricted in distribution to “central Panama” (Kohler 2008:

100). This individual came from Parque Nacional General de

Division Omar Torrijos Herrera, in the province of Coclé, Panama.

Photo by Abel Batista.

Plate 31. Norops kemptoni (Dunn, 1940). Kempton’s Anole is a

priority one species with an EVS of 15, found in the “highlands

of Chiriqui, Panama” (Kohler 2008: 106). This individual is from

Alto Chiquero, Parque Nacional Volcan Baru, in the province of

Chiriqui, Panama. Photo by Javier Sunyer.

January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

Plate 32. Coleonyx mitratus (Peters, 1863). The Central American

Banded Gecko is a priority six species with an EVS of 14, which

ranges along the “Atlantic lowlands of northeastern Guatemala

and northwestern Honduras [and the] Pacific lowlands from

Guatemala to southwestern Costa Rica” (Savage 2002: 482). This

individual was located in Sector Santa Rosa, Parque Nacional

Santa Rosa, in the province of, Guanacaste, Costa Rica. Photo by

Victor Acosta-Chaves.

Plate 34, Sceloporus Saale ntnedis oe 1864. The Green oe

Lizard is a priority three species with an EVS of 10, distributed

from “El Salvador and Honduras across Nicaragua and Costa Rica

to Panama” (Kohler 2008: 152). This individual was encountered

in Cerro de la Muerte, in the province of San José, Costa Rica.

Photo by Victor Acosta-Chaves.

Plate 36. hcinaed Maieates (Cope, 1876) The Thick Graceful

Brownsnake is a priority one species with an EVS of 14, distributed

in “the Cordillera de Tilaran, Cordillera Central, and Cordillera de

Talamanca of Costa Rica and on Volcan Tenorio in the Cordillera

de Guanacaste...to extreme western Panama” (Savage 20002:

623). This individual came from San Gerardo de Dota, in the

province of San José, Costa Rica. Photo by Victor Acosta-Chaves.

Amphib. Reptile Conserv.

Plate 33. Heloderma Rea Campbell and Vanini, 1988.

The Guatemalan Beaded Lizard is a priority two species that

inhabits “the Rio Motagua Valley, in the Atlantic versant of eastern

Guatemala” (Reiserer et al. 2013: 81). This individual was found

at Cabafias, in the department of Zacapa, Guatemala. Photo by

Andres Novales.

Plate 35. Scolecophis atrocinctus (Schlegel, 1837). priority three

species with an EVS of 13, which ranges along the “Pacific

versant from southeastern Guatemala to northwestern Costa Rica;

the species also is found on the Atlantic versant in southwestern

Honduras, western Nicaragua, and northwestern Costa Rica”

(Wilson and Mata-Silva 2015: 422). This individual was found

in Tilaran, in the province of Guanacaste, Costa Rica. Photo by

Victor Acosta-Chaves.

Plate 37. Bothriechis lateralis Peters, 1862. The Coffee Palmviper

is a priority one species with an EVS of 16. distributed in “the

cordilleras of Costa Rica and western Panama (Savage, 2002:

724). This individual came from La Nevera, Serrania de Tabasara,

Panama. Photo by Abel Batista.

January 2019 | Volume 13 | Number 1 | e168

Mata-Silva et al.

F. The Central American endemic herpetofauna is

distributed unevenly among the 10 physiographic regions

we recognize in this portion of Mesoamerica. The number

of endemic species in these regions ranges from six in the

Yucatan Platform to 254 in the Isthmian Central American

highlands. Most of the 623 endemic species are limited to

a single physiographic region, 1.e., 450 (72.2%). The next

largest number of 95(15.2%) is for those occupying two

regions. Thus, 545 (87.5%) of the species occur in only one

or two of the 10 regions.

G. An implementation of the EVS system of conservation

assessment demonstrates that all of the scores for the 623

endemic species in Central America lie within the medium

vulnerability (63) or high vulnerability (560) categories.

H. We used the same means as Johnson et al. (2017) for

the Mexican endemic species to prioritize the conservation

significance of the Central American endemic species, 1.e.,

by combining the data on physiographic distribution with

that on EVS. This procedure allowed us to identify 14

priority levels, of which six are high vulnerability and eight

are medium vulnerability groupings.

I. The number of species occupying the six high vulnerability

levels decrease markedly and consistently from 429 in

Priority Level One (high vulnerability species occupying

a single physiographic region) to one in Priority Level Six

(high vulnerability species occurring in six physiographic

regions. The total number of species allocated to the seven

priority levels amounts to 562, 90.2% of the 623 Central

American endemic species.

J. The number of species occurring in the eight medium

vulnerability levels also decreases consistently, but not

as markedly, from 23 in Priority Level Seven (medium

vulnerability species occupying single physiographic

regions) to one in Priority Level Fourteen (medium

vulnerability species distributed in eight physiographic

regions). The total number of species placed in these eight

priority levels is 61, 9.8% of the total number of Central

American endemics.

K. Our analysis demonstrates that most Central American

endemic species are assessed as high vulnerability species

that occupy only one or two physiographic regions.

A significant number of species (44) are of medium

vulnerability, and they also inhabit one or two physiographic

regions.

L. Protecting the endemic component of the globally

significant Central American herpetofauna represents the

most significant challenge for conservation professionals

working in this portion of Mesoamerica.

M. One of the conclusions of the conservation analyses

we have conducted in recent years, including in this

paper, is that perpetual protection of the Mesoamerican

herpetofauna presently is a goal far from realization. Our

prognosis is that this goal will not be attained until humans

are willing to address the widespread problems created

by the anthropocentric worldview, and the impediment

Amphib. Reptile Conserv.

it represents for allowing the environment “to function

indefinitely without going into a decline from the stresses

imposed by human society on natural systems such as fertile

soil, water, and air” (Raven and Berg 2004: G-15).

N. Ifthereis any merit to our hypothesis that anthropocentrism

is part of a cascade of psychological ailments, which extend

through ethnocentrism and culminate in the narcissistic

personality disorder, it might predict that the critical-

thinking educational reform called for by Johnson et al.

(2017) will have to be recognized as requiring species-

wide psychotherapy to treat a species-wide mental disease.

If so, addressing this disease will be the largest problem

undertaken by humanity during its existence on planet

Earth.

Recommendations

A. The recommendations presented in the Johnson et al.

(2017: 616) study on the Mexican endemic herpetofauna

dealt with the establishment of a global coalition “to

document all of Earth’s inhabitants within the 21* century,

and to provide for their perpetual protection.” This

recommendation applies here as well.

B. Johnson et al. (2017) also recommended that the system

of prioritization they developed could help determine how

funding for a countrywide system of sustainable reserves

could be best utilized to protect the Mexican endemic

herpetofauna. The same can be said for the Central

American endemic herpetofauna, with respect to the seven

nations comprising this region of the world.

C. We propose that until such steps are taken, the Central

American endemic herpetofauna, as well as the entire

planetary biota, will become a casualty of anthropocentrism.

Nonetheless, these steps only will constitute stopgap

measures, as the underlying anthropocentric worldview held

by most humans will continue to accelerate the degradation

of the planetary life-support systems, and eventually will

render the planet unsuitable to support life.

“.. Each loss of a species is a loss of a companion on the

long journey of evolution ...”’

—David W. Orr (2016).

Acknowledgments.—We thank the people who gra-

ciously allowed us to use their beautiful tmages of various

Central American endemic species to illustrate this paper,

including Victor Acosta-Chaves, César Barrio-Amoros,

Abel Batista, Andres Novales, Todd Pierson, Sean Michael

Rovito, and Javier Sunyer. We also are appreciative of the

assistance of Alan Resetar of the Field Museum of Natural

History in providing us with photographs of FMNH 74376,

the specimen tentatively identified as Dipsas viguieri by

Cadle (2005). Abel Batista, Sebastian Lotzkat, and Louis W.

Porras significantly improved the quality of the manuscript

with their outstanding reviews. We are indebted hugely to

these colleagues for sharing their expertise with us.

January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

Paar / OD ate +o : Ree ¥ x mai | \ ea, Set ~) ™ 4 s : 2 min .

Plate 38. Bothriechis nigroviridis Peters, 1859. The Black-speckled Palm-pitviper is a priority one species with an EVS of 17, which

ranges along “the cordilleras of Costa Rica and western Panama” (Savage, 2002: 725). This individual was found at Jurutungo, in the

province of Chiriqui, Panama. Photo by Javier Sunyer.

Plate 39. Lachesis stenophrys Cope, 1875. The Central American Bushmaster is a priority two species with an EVS of 17, distributed

from “southeastern Nicaragua to central Panama” (Kohler 2008: 330). This individual came from Parque Nacional Braulio Carrillo, in the

province of Heredia, Costa Rica. Photo by César Barrio-Amoros.

a . lt

= = e <e “pg ls ¢

= : e x se

Pe Ae ; pe Spy :

Plate 40. Porthidium porrasi Lamar, 2003. The White-tailed Hog-nosed Pitviper is a priority one species with an EVS of 18, with a

distribution restricted to the Peninsula de Osa and adjacent areas of southwestern Costa Rica (Solorzano 2004). Pictured here is an

individual from this region, in the province of Puntarenas, Costa Rica. Photo by César Barrio-Amoros, courtesy of Roel de Plecker.

Amphib. Reptile Conserv. 59 January 2019 | Volume 13 | Number 1 | e168

Mata-Silva et al.

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January 2019 | Volume 13 | Number 1 | e168

The endemic herpetofauna of Central America

Addendum (changes past conclusion of analyses)

We chose a cut-off date of 30 April 2018 to discontinue

revising the huge number of calculations dealing with the

623 endemic herpetofaunal species documented in Central

America. Past this date, we added pertinent taxa and

publications to this addendum until the correction of the

proof, as follows:

(1) Craugastor aenigmaticus. Arias et al. (2018) described

this new species of rainfrog from the Cordillera de

Talamanca in southern Costa Rica. This frog is known from

several localities in montane rainforest, and its EVS can be

calculated as 5+8+4=17, placing it in the middle portion of

the high vulnerability category. This species is limited to a

single physiographic region, the Isthmian Central American

highlands, and can be placed in conservation priority level one.

(2) Craugastor castanedai. McCranie (2018) described

this new species of rainfrog from the Parque Nacional Pico

Bonito in north-central Honduras. This frog is known from

only two localities on either side of the Quebrada de Oro,

and its EVS can be calculated as 6+8+4=18, placing it in

the upper portion of the high vulnerability category. This

species is limited to a single physiographic region, the

eastern nuclear Central American highlands, and can be

placed in conservation priority level one.

(3) Craugastor gutschei. McCranie (2018) also described

this new species of rainfrog from the western portion of

the Cordillera Nombre de Dios in north-central Honduras.

Its EVS can be calculated as 5+7+4=16, placing it in the

middle portion of the high vulnerability category. This

Species 1s restricted to the eastern nuclear Central American

highlands, and can be allocated to conservation priority

level one.

(4) Hemiphractus elioti. Hill et al. (2018) described this new

species of horned frog from the Cordillera de Talamanca in

western Panama. Its EVS can be determined as 5+7+5=17,

placing it in the middle portion of the high vulnerability

category. This species 1s restricted to the Isthmian Central

American highlands, and can be allocated to conservation

priority level one.

(5) Hemiphractus kaylockae. Hill et al. (2018) named this

new species of horned frog from the highlands of eastern

Amphib. Reptile Conserv.

Panama. Its EVS can be estimated as 6+8+5=19, placing it

in the upper portion of the high vulnerability category. This

Species 1s restricted to the eastern Panamanian highlands,

and can be placed in conservation priority level one.

(6) Hemiphractus panamensis. Hill et al. (2018) revalidated

this taxon of horned frogs from the eastern highlands of

Panama. Its EVS can be determined as 5+8+5=18, placing it

in the upper portion of the high vulnerability category. This

species is restricted to the western portion of the eastern

Panamanian highlands in the central portion of the country,

and can be placed in conservation priority level one.

(7) Hemiphractus fasciatus. Hill et al. (2018) determined

that this species, formerly considered to be the single

representative of this peculiar hylid genus in Central

America, should be considered to be a South American

taxon not resident in Panama.

(8) Norops caceresae. Hofmann and Townsend (2018)

described this anole from the Lenca highlands of

southwestern Honduras. Its EVS can be estimated as

5+7+3=15, placing in the lower portion of the high

vulnerability category. This species is limited to the eastern

nuclear Central American highlands, and can be placed in

conservation priority level one.

(9) Rhadinella xerophila. Ariano-Sanchez. and Campbell

(2018) described this dipsadid snake from dry forest and

thorn scrub of the Valle del Motagua, Guatemala. Its EVS

can be calculated as 6+8+2=16, placing it in the middle

portion of the high vulnerability category. This species 1s

restricted to a single physiographic region, the Caribbean

lowlands of eastern Guatemala and northern Honduras, and

can be placed in conservation priority level one.

(10) The genera Coloptychon and Gerrhonotus. Garcia-

Vazquez et al. (2018), in a paper on the molecular

systematics and historical biogeography of the anguid genus

Gerrhonotus, left open a number of taxonomic questions

about the alligator lizards, but at least seemed to conclude

that the monotypic genus Coloptychon (and its species

rhombifer) should be returned to the genus Gerrhonotus.

The genus Coloptychon has had a history of wobbling

between recognition as distinct or not from Gerrhonotus

over the course of its 142-year history. In this addendum,

we follow the decision of Garcia-Vazquez et al. (2018).

January 2019 | Volume 13 | Number 1 | e168

Mata-Silva et al.

Vicente Mata-Silva is a herpetologist originally from Rio Grande, Oaxaca, Mexico. His interests include

_ ecology, conservation, natural history, and biogeography of the herpetofaunas of Mexico, Central America,

and the southwestern United States. He received his B.S. degree from the Universidad Nacional Autonoma de

México (UNAM), and his M.S. and Ph.D. degrees from the University of Texas at El Paso (UTEP). Vicente

is an Assistant Professor of Biological Sciences at UTEP in the Ecology and Evolutionary Biology Program,

and Assistant Director of UTEP’s 40,000 acre Indio Mountains Research Station, located in the Chihuahuan

Desert of Trans-Pecos, Texas. To date, Vicente has authored or co-authored over 100 peer-reviewed scientific

publications. He also was the Distribution Notes Section Editor for the journal Mesoamerican Herpetology.

Dominic L. DeSantis is currently a Ph.D. candidate and National Science Foundation-Graduate

Research Fellow at the University of Texas at El Paso. He received his Bachelor’s degree at Texas

State University where he also completed multiple research projects on the antipredator behavior of

the critically endangered Barton Springs Salamander (Eurycea sosorum). His ongoing dissertation

@s research integrates multiple field monitoring technologies to study snake movement and behavioral

§ ecology. Dominic accompanied Vicente Mata-Silva, Eli Garcia-Padilla, and Larry David Wilson on

survey and collecting trips to Oaxaca in 2015, 2016, and 2017 and is a co-author on numerous natural

history publications produced from those visits, including an invited book chapter entitled: “Conservation of Herpetofauna in Disturbed

Habitats: Perspectives from Short-term Surveys in the Sierra Madre del Sur, Oaxaca, Mexico.” Overall, Dominic has co-authored over

50 peer-reviewed scientific publications.

Eli Garcia-Padilla is a herpetologist primarily focused on the study of the ecology and natural history of

the Mexican herpetofauna. His research efforts have centered on the Mexican states of Baja California,

Tamaulipas, Chiapas, and Oaxaca. His first experience in the field was researching the ecology of the

insular endemic populations of the rattlesnakes Crotalus catalinensis, C. muertensis (now allocated to C.

pyrrhus) and C. tortugensis (now allocated to C. atrox) in the Gulf of California. For his Bachelor’s degree

he presented a thesis on the ecology of C. muertensis (now allocated to C. pyrrhus) on Isla El Muerto, Baja

California, Mexico. To date, he has authored or co-authored over 75 peer-reviewed scientific publications.

Currently, he is Saplees as a formal Curator of Amphibians and Reptiles from Mexico in the electronic platform “Naturalista” of the

Comision Nacional para el Uso y Conocimiento de la Biodiversidad (CONABIO; www.naturalista.mx). One of his main passions is

environmental education, and for several years he has been working on a variety of projects that include the use of audiovisual media as

a powerful tool to reach large audiences and to promote the importance of the knowledge, protection, and conservation of biodiversity

in Mexico. Eli’s interests include wildlife and conservation photography, and his art has been published in several recognized scientific,

artistic, and educational books, magazines, and websites. Presently, he is collaborating on a research project evaluating the Jaguar (Panthera

onca) as an umbrella species for the conservation of the herpetofauna of Nuclear Central America.

Jerry D. Johnson is Professor of Biological Sciences at The University of Texas at El Paso, and has

extensive experience studying the herpetofauna of Mesoamerica, especially that of southern Mexico.

Jerry is the Director of the 40,000-acre “Indio Mountains Research Station,” was a co-editor on

Conservation of Mesoamerican Amphibians and Reptiles and co-author of four of its chapters, co-editor

of Mesoamerican Herpetology: Systematics, Zoogeography, and Conservation, and co-author of Middle

American Herpetology: A Bibliographic Checklist. He also is the senior author of the recent paper “A

conservation reassessment of the Central American herpetofauna based on the EVS measure” and is

Mesoamerica/Caribbean editor for Geographic Distribution section of Herpetological Review. Johnson

has authored or co-authored over 120 peer-reviewed papers, including other studies on the conservation

status of the Mesoamerican herpeteofauna. One species, Zantilla johnsoni, has been named in his honor.

For several years, he was an Associate Editor and Co-chair of the Taxonomic Board for the journal

Mesoamerican Herpetology.

Larry David Wilson is a herpetologist with lengthy experience in Mesoamerica. He was born in

Taylorville, Illinois, United States, and received his university education at the University of Illinois at

Champaign-Urbana (B.S. degree) and at Louisiana State University in Baton Rouge (M.S. and Ph.D.

degrees). He has authored or co-authored over 400 peer-reviewed papers and books on herpetology,

including numerous papers on the conservation status of Mesoamerica and its constituent parts. Larry is

the senior editor of Conservation of Mesoamerican Amphibians and Reptiles and the co-author of seven

of its 21 chapters. His other books include The Snakes of Honduras, Middle American Herpetology, The

: Amphibians of Honduras, Amphibians & Reptiles of the Bay Islands and Cayos Cochinos, Honduras, The

Amphibians and Reptiles of the Honduran Mosquitia, and Guide to the Amphibians & Reptiles of Cusuco National Park, Honduras.

To date, he has authored or co-authored the descriptions of 71 currently recognized herpetofaunal species, and seven species have

been named in his honor, including the anuran Craugastor lauraster, the lizard Norops wilsoni, and the snakes Oxybelis wilsoni,

Myriopholis wilsoni, and Cerrophidion wilsoni. For several years, Larry was an Associate Editor and Co-chair of the Taxonomic

Board for the journal Mesoamerican Herpetology.

Amphib. Reptile Conserv. 64 January 2019 | Volume 13 | Number 1 | e168

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

13(1) [General Section]: 65-77 (e169).

Short Communication

Ecological interactions between arthropods and small

vertebrates in a lowland Amazon rainforest

‘Rudolf von May, 7Emanuele Biggi, *Heidy Cardenas, ‘M. Isabel Diaz, **Consuelo Alarcon, ‘Valia

Herrera, *Roy Santa-Cruz, *Francesco Tomasinelli, ‘°Erin P. Westeen, ‘Ciara M. Sanchez-Paredes,

‘Joanna G. Larson, ‘Pascal O. Title, ‘°Maggie R. Grundler, ‘Michael C. Grundler, ‘Alison R. Davis

Rabosky, and ‘Daniel L. Rabosky

'Museum of Zoology, Department of Ecology and Evolutionary Biology, University of Michigan, Biological Sciences Building, 1105 N. University,

Ann Arbor, Michigan 48109-1085, USA 2International League of Conservation Photographers *Area de Herpetologia, Museo de Historia Natural

de la Universidad Nacional de San Agustin (MUSA), Av. Alcides Carrion s/n, Arequipa, PERU ‘Universidad Nacional de San Antonio Abad

del Cusco, Cusco, Peru y Museo de Biodiversidad del Peru, Cusco, PERU ‘Department of Biology, John Carroll University, 1 John Carroll

Boulevard, University Heights, Ohio 44118, USA °Museo de Biodiversidad del Peru, Urbanizacion Mariscal Gamarra A-61, Zona 2, Cusco, PERU

’Departamento de Herpetologia, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Av. Arenales 1256, Jesus Maria, Lima,

PERU 8Milan, ITALY °Department of Environmental Science, Policy, and Management, University of California, Berkeley, California 94720, USA

'0Laboratorio de Estudios en Biodiversidad, Universidad Peruana Cayetano Heredia, Lima, PERU "Environmental Resilience Institute, Indiana

University, 717 E 8" St, Bloomington, Indiana 47408, USA

Abstract.—Ecological interactions such as those involving arthropod predators and parasitoids and their prey

or hosts provide evidence for selective pressures influencing small vertebrate populations, and are key to

understanding the many connections that shape food webs in tropical rainforests. Here, we document 15

predator-prey interactions involving different types of arthropod predators and vertebrate prey including frogs,

lizards, snakes, and a mammal. Documented also are three cases of fly myiasis in frogs, and provide further

evidence of a commensal relationship involving a tarantula and a narrow-mouthed frog in lowland Amazonian

Peru.

Keywords. Amazonia, amphibians, centipedes, commensalism, frogs, lizards, myiasis, opossums, parasitoids, preda-

tor-prey, reptiles, snakes, spiders, stingless bees, water bugs

Citation: von May R, Biggi E, Cardenas H, Diaz MI, Alarcén C, Herrera V, Santa-Cruz R, Tomasinelli F, Westeen EP, Sanchez-Paredes CM, Larson

JG, Title PO, Grundler MR, Grundler MC, Rabosky ARD, Rabosky DL. 2019. Ecological interactions between arthropods and small vertebrates in a

lowland Amazon rainforest. Amphibian & Reptile Conservation 13(1) [General Section]: 65—77 (e169).

4.0 International (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in any

medium, provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced, are

as follows: official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org.

Received: 25 May 2018; Accepted: 12 September 2018; Published: 28 February 2019

Many groups of arthropods are predators of vertebrates

and play a critical role in the structure and functioning

of food webs (McCormick and Polis 1982). Spiders are

among the most diverse predaceous arthropods in the

tropics and exhibit high levels of both family and spe-

cies richness in this climatic region (Cardoso et al. 2011).

Reports of spider predation on vertebrates include prey

from all major vertebrate taxonomic groups including

fishes, amphibians, reptiles, birds, and mammals (Mc-

Cormick and Polis 1982; Greene 1988; Das et al. 2012:

Nyffeler and Knornschild 2013; Nyffeler and Pusey

2014). In addition to spiders, other predaceous arthro-

pods common in terrestrial environments include scorpi-

ons, centipedes, ants, and beetles, while those in aquatic

Correspondence. !rvonmay@umich.edu

Amphib. Reptile Conserv.

environments include water bugs, dragonfly larvae, div-

ing beetles, and other invertebrates (Corey 1988; Toledo

2005; Wells 2007; von May 2009a; Biggi and Tomasinel-

li 2017). Most predaceous arthropods rely on specialized

trophic structures and venom to capture and paralyze ver-

tebrate prey (McCormick and Polis 1982). Morphologi-

cal adaptations include modified jaws, enlarged beaks,

and massive chelicerae. Some taxa have evolved dozens

of venom proteins that are injected at once during prey

capture. For example, the venom of some species of gi-

ant water bugs (Belostomatidae) contains a powerful mix

of nearly 130 venom proteins, including cytolytic toxins,

antimicrobial peptides, and enzymes, that they inject in

their fish prey (Walker et al. 2018).

February 2019 | Volume 13 | Number 1 | e169

von May et al.

> ls »

> Ermanvele Bi ap ls " re 4 > fal — ° »

ue afr a or

Fig. 1. (A) The spider Ancylometes sp. (Ctenidae) preying upon an adult Dendropsophus leali;, (B) the spider Phoneutria sp.

(Ctenidae) preying on a sub-adult Hamptophryne boliviana. Photos by Emanuele Biggi (A) and Francesco Tomasinelli (B).

Predation of small vertebrates by arthropods has been __ predator-prey observations from Villa Carmen Biologi-

documented in several lowland rainforest sites (e.g., cal Station (12°53’43.8”S, 71°24’13.7”°W, 520 m elev.),

Corey 1988; Menin et al. 2005; Toledo 2005; Das et also located in Madre de Dios region, and one parasitoid

al. 2012; Nyffeler and Knornschild 2013; Nyffeler and —_ observation from Madre Selva Research Station, Loreto

Pusey 2014), yet our knowledge of these interactions re- = region, northern Peru (3°37°39.1”S, 72°14’24.4’°W, 105

mains limited, especially given the diversity of vertebrate | _m elev.). The predator-prey observations are presented

prey and potential arthropod predators in species-rich — in chronological order and involve prey in different life

tropical communities. It is valuable to document these stages including eggs, larvae, juveniles, and adults. In

predator-prey interactions in the field, because they pro- § most cases, the prey specimens were not collected be-

vide a snapshot of the many connections that shape food cause they were consumed by the predators. Identifi-

webs and provide evidence for selective pressures influ- = cation of prey was primarily done in the field and was

encing small vertebrate populations. This is especially | subsequently confirmed based on photographs presented

important in tropical rainforests, given their exceptional herein. The observations on parasitoid infections and

levels of biodiversity. Documenting predation by spiders |= commensalism are also presented in chronological order.

and other arthropods in these ecosystems is essential

even if many predaceous arthropods remain undescribed | Predator-prey Interactions

and are mostly classified as morphospecies (Cardoso et

al. 2011), because they provide insights into an impor- On 20 February 2008, at 1947 h, we observed a spider of

tant source of vertebrate mortality that appears to be less —_ the genus Ancylometes (Ctenidae) preying upon an adult

common in extra-tropical communities. Dendropsophus leali (Hylidae; Figure 1A). The spider

Here, documented are 15 predator-prey interactions held its prey tightly by the back. This observation took

involving different types of arthropod predators and ver- __ place in the floodplain forest, at a site located approxi-

tebrate prey including frogs, lizards, snakes, anda mam- _—_— mately 2 km from the station.

mal. Also documented are three cases of fly myiasis (live

parasitic infestation by maggots) in frogs, and provide On 22 February 2008, at 2056 h, we observed a spider of

further evidence of a commensal relationship involving — the genus Phoneutria (Ctenidae) preying on a sub-adult

a tarantula and a narrow-mouthed frog in lowland Ama- = Hamptophryne boliviana (Microhylidae; Figure 1B).

zonian Peru. The spider held its prey tightly by the body and the frog

Our main study site, Los Amigos Biological Sta- did not display any movement. This observation took

tion (12°34’07”S, 70°05’57”W, 250 m elev.), is located place in the floodplain forest, at a site located 1.5 km

in Madre de Dios region, southeastern Peru. A general from the station.

overview of the amphibian and reptile fauna, the habi-

tats, and the local climate at this site was provided by On 23 February 2008, around 2110 h, we observed a fish-

von May et al. (2006, 2009b, 2010a,b), and Whitworth — ing spider of the genus 7haumasia (Pisauridae) preying

et al. (2016) provided a preliminary list of the reptile | upona tadpole (unidentified) at a temporary pond located

taxa recorded at this site. Additionally, we report two __ in terra firme forest (Figure 2A). The pond is located near

Amphib. Reptile Conserv. 66 February 2019 | Volume 13 | Number 1 | e169

Ecological interactions between arthropods and small vertebrates

nm . -

Emanue e Bigg - Ahuratit”

Fig. 2. (A) The fishing spider Thaumasia sp. (Pisauridae) preying upon a tadpole (unide

tified) ata temporary pond located in terra

firme forest; (B) a ctenid spider (genus undetermined; Ctenidae) preying upon a subadult Boana sp. G. Photos by Emanuele Biggi

(A) and Francesco Tomasinelli (B).

(~50 m) the station laboratories and is used by at least a

dozen amphibian species during the wet season (R. von

May, pers. obs.).

On 25 February 2008, at 2122 h, we observed a ctenid

spider (genus undetermined; Ctenidae) preying upon a

subadult Boana sp. G (Hylidae; Figure 2B). The spider

was on top and the frog did not display any movement.

This observation took place in the terra firme forest, at a

site located approximately 1 km from the station.

On 25 February 2008, around 2200 h, we observed a th-

eraphosid spider (Theraphosidae), possibly in the genus

Pamphobeteus, preying upon an adult Hamptophryne

boliviana (Microhylidae; Figure 3A). Both individuals

were found in the leaf litter, and the spider was holding

the frog by the posterior section of the body. This obser-

: —~»

bates ail - —— x“ 2 © ./%

Fig. 3. (A) A theraphosid spider, cf. Pamphobeteus sp. (Theraphosidae), preying upon Hamptophryne boliviana; (B) a ctenid spider

vation took place in the floodplain forest, at a site located

approximately 1.5 km from the station.

On 11 March 2016, during a night survey, we observed a

wandering spider (Ctenidae) preying upon an individual

of Leptodactylus didymus (Leptodactylidae). Both indi-

viduals were found on a leaf, and the spider was holding

the frog by the back (Figure 3B). We encountered both

individuals in the terra firme forest approximately 1 km

from the station.

On 17 March 2016, at 2130 h, during a leaf-litter plot sur-

vey, we observed a spider of the genus Crenus (Ctenidae)

holding an individual of Cercosaura eigenmanni (Gym-

nophthalmidae) by the middle of its body (Fig. 4). Upon

capture, the spider held its prey tight; the lizard tail kept

moving for several minutes after capture. At the time of

~ &

—_

(Ctenidae) preying upon Leptodactylus didymus. Photos by Emanuele Biggi (A) and Pascal Title (C).

Amphib. Reptile Conserv.

February 2019 | Volume 13 | Number 1 | e169

von May et al.

Fig. 4. The spider Crenus sp. (Ctenidae) preying upon a subadult Cercosaura eigenmani. Photo by Mark Cowan.

preservation of both individuals, the spider had released

its prey, which was already dead. This observation took

place in the terra firme forest, at a site located 3.5 km

from the station.

On 24 October 2016, at ~2100 h we disturbed a scolo-

pendrid centipede (Chilopoda, Scolopendridae) that was

consuming a live juvenile snake, Dipsas catesbyi (Col-

ubridae), in the leaf litter near Villa Carmen Biological

Station. A precise identification of the predator is not

possible because it abandoned its prey and quickly re-

treated into the leaf litter before we had time to take a

photograph. The snake had a gaping wound in its right

side, where the scolopendrid had eaten through the skin

and muscle of the body wall (Fig 5A). Due to the sever-

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ity of its injuries, the snake was humanely euthanized,

preserved as a voucher specimen, and deposited in the

herpetological collection at the Museum of Natural His-

tory, Universidad Nacional Mayor de San Marcos, Lima,

Peru (MUSM 37084). It had an SVL of 245 mm and a tail

length of 84 mm.

On the morning of 7 November 2016, we found a juve-

nile snake, Micrurus obscurus (Elapidae) deceased in a

funnel trap, being consumed by a scolopendrid centipede

(Chilopoda, Scolopendridae). By the time we removed

the predator from the trap, 1t had decapitated the snake

and removed the skin and muscle from ~20 mm of the

vertebral column (Fig 5B). This observation took place

in a forest with short and narrow stemmed trees with

Fig. 5. (A) Juvenile snake Dipsas catesbyi with lesion caused by scolopendrid centipede (red arrow); (B) juvenile snake Micrurus

obscurus, missing head and soft tissues on most anterior part of body as a result of predation by scolopendrid centipede. Photos by

Joanna Larson (A—B).

Amphib. Reptile Conserv. 68 February 2019 | Volume 13 | Number 1 | e169

Ecological interactions between arthropods and small vertebrates

ee OS

Fig, 6. (A) Theraphosid aside Pamphobeteus sp. (Theraphosidae) preying upon the 1 mouse opossum MEE OSESE cf. noctivagus,

(B) The same individual of Pamphobeteus sp. dragging the mouse opossum on the leaf litter. Photos by Maggie Grundler (A—B).

little understory and rocky soil near Villa Carmen Bio-

logical Station. The snake had an SVL of >248 mm anda

tail length of 16 mm. The snake specimen was deposited

in the herpetological collection at the Museum of Natu-

ral History, Universidad Nacional Mayor de San Marcos,

Lima, Peru (MUSM 37349).

On 18 November 2016, at 2313 h during a night survey,

we observed a theraphosid spider, Pamphobeteus sp.

(Theraphosidae) preying upon a mouse opossum, Mar-

mosops Cf. noctivagus (Didelphidae). The spider was on

the ground in the leaf litter holding the opossum by the

neck region (Fig. 6); a video of the interaction is archived

in the University of Michigan Deep Blue Data repository

(doi.org/10.7302/Z2862DP1). When we first encoun-

tered the pair, the opossum was still responsive and kick-

ing weakly. We observed the interaction for approxi-

mately 5 minutes after which time the opossum ceased

all movement and the spider dragged it away around a

tree root. This observation took pigee in age floodplain

forest, approximately 2 km from the station. It 1s worth

noting that this appears to be the first documentation of a

large mygalomorph spider (infraorder Mygalomorphae)

preying upon opossums (R. Voss, pers. comm.). Opos-

sum species in the genus Marmosops are primarily found

on the forest floor, are active at night, and live in different

habitats including old growth forest, secondary forest,

and open areas (Emmons and Feer 1997). In the lowlands

of southeastern Peru, Marmosops noctivagus 1s the only

local species with clear-white underparts, and the grayish

dorsal pelage of the individual preyed upon by the spider

(Fig. 6) suggests that it was a juvenile or subadult (R.

Voss, pers. comm. ).

On 22 November 2017, at 2254 h during a standardized

night survey, we observed a wandering spider (Ctenidae)

preying upon an individual of Hamptophryne boliviana

(Microhylidae). Both individuals were found in the leaf

litter, and the spider was holding by the rear right leg

(Fig. apy We observed as the spider manipulated the

Fig. 7. (A) A wandering spider (Ctenidae) preying upon EEE cane ae (B) the spider Ancylometes sp. (eienidae) preying

upon an adult Dendropsophus sarayacuensis, (C) giant water bug (Belostomatidae) preying upon an adult Dendropsophus minutus;

the belostomatid was guarding a clutch of eggs (likely its own clutch). Photos by Erin Westeen (A) and Maria Isabel Diaz (B—C).

February 2019 | Volume 13 | Number 1 | e169

Amphib. Reptile Conserv.

von May et al.

Fig. 8. (A) Stingless bees in the genus 7rigona (Apidae) preying upon a clutch of tree frog eggs (Hylidae) at a temporary pond

located in terra firme forest; (B) the spider Phoneutria sp. (Ctenidae) preying upon an adult Dendropsophus kamagarini. Photos by

Rudolf von May (A) and Roy Santa-Cruz (B).

frog and maintained a hold of its leg for 20 minutes be-

fore continuing with the survey. Both individuals were

encountered in the leaf litter in the terra firme forest, ap-

proximately 400 m from the station. The spider superfi-

cially resembles those in the genus Ancylometes (Cteni-

dae); however, the genus assignment is tentative because

no voucher specimen 1s available.

On 26 November 2017, at 2313 h, during a survey of

frog breeding activity in a temporary pond, we observed

a spider of the genus Ancylometes (Ctenidae) holding an

individual of Dendropsophus sarayacuensis (Hylidae)

by the head and the anterior part of the body (Fig. 7B).

Both individuals were on the trunk of a tree located in an

inundated area near the pond’s edge, 0.35 m above the

water surface. This observation took place in the terra

firme forest, at a site located approximately 600 m from

the station.

On 27 November 2017, at 2219 h, we observed a giant

water bug (Belostomatidae) holding an individual of

Dendropsophus minutus (Hylidae) by the body (Fig. 7C).

The belostomatid was perched on the stem of a small

plant and was guarding a clutch of eggs (likely its own

clutch), ca. 0.2 m above the water surface. This observa-

tion took place in the terra firme forest, at a site located

approximately 600 m from the station.

On 29 November 2017, at 1410 h, we observed stingless

bees of the genus 7rigona (Apidae) preying upon a clutch

of tree frog eggs (Hylidae) at a temporary pond located

in terra firme forest (Fig. 8A). The pond is located ap-

proximately 600 m from the station. The egg clutch was

attached to a small tree branch, ca. 1.0 m above the water

surface, and was in partial shade. Prior to this observa-

tion, several species of tree frogs of the genus Dendrop-

sophus (Hylidae) were observed breeding at this pond

Amphib. Reptile Conserv.

over several nights (H. Cardenas and M.I. Diaz, pers.

obs.).

On 29 November 2017, at 2110 h, we observed a spider

of the genus Phoneutria (Ctenidae) preying upon an adult

Dendropsophus kamagarini (Hylidae). Though the only

photo available of this event (Fig. 8B) does not show the

spider holding onto the frog, the actual predation event

appeared to have had taken place moments prior to find-

ing the spider and frog in the field (as suggested by the

frog posture in the image). After taking the photo, the

frog did not display any movement and the spider started

eating it (no additional image is available). This observa-

tion took place in the terra firme forest, at a site located

approximately 100 m from the station.

Most predation events documented here involve spiders

that belong to different families and those in the fam-

ily Ctenidae were the most frequently seen predators

(Appendix 1). Ctenid spiders, also known as wandering

spiders, are ubiquitous across Neotropical rainforests

and forage in different microhabitats including the leaf-

litter, tree trunks, and the canopy (Gasnier et al. 1995;

Pétillon et al. 2018). Ctenid spiders are typically sit-and-

wait predators, which choose a leaf or a branch for their

nightly hunt and are very sensitive to air and ground vi-

brations, but also their eyes seem to play a role in prey

and motion detection (Neuhofer et al. 2009). They use

specialized hairs (also known as trichobothria; Barth et

al. 1993) on the legs and pedipalps to detect air vibra-

tion and the direction of prey. The principal eyes are re-

sponsible for object discrimination (Schmid 1998) and

the secondary eyes are responsible for motion detection

(Neuhofer et al. 2009). Depending on the species, they

tend to hunt terrestrially (e.g., many Ctenus species) or

arboreally (e.g., Phoneutria and Cupiennius). However,

these strategies are not strict and we found these spiders

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Ecological interactions between arthropods and small vertebrates

Fig. 9. (A) Osteocephalus cf. leprieurii infected by several fly larvae; part of the skin of the infected area was removed to show

cavity with degraded tissue and one fly larva (on right); (B) Dendropsophus leali and fly larvae (Diptera) that emerged through the

frog’s mouth; (C) Ranitomeya uakarii infected by a maggot that emerged from a small round lesion on its back. Photos by Rudolf

von May (A), Daniel Rabosky (B), and Valia Herrera (C).

almost everywhere from the ground up to the canopy (E.

Biggi and F. Tomasinelli, pers. obs.).

Parasitoid Infections

Though technically distinct from predators, parasitoids

may also represent an important source of mortality in

small Amazonian vertebrates. Several studies have docu-

mented lethal cases of flesh fly myiasis affecting small

frogs (Crump and Pounds 1985; Hagman et al. 2005;

Eizemberg et al. 2008). Below, we document three cases

of fly myiasis in lowland Amazonian frogs.

On 12 January 2012, around 2210 h, we found a juve-

nile Osteocephalus cf. leprieurii infected by several fly

larvae. The frog was captured during a visual encounter

survey in the floodplain forest, approximately 1 km from

the station. The frog was placed in a perforated plastic

container and observed over the following day. At mid-

afternoon, the frog had died and we removed three fly

larvae; the cutaneous lesions on its back were conspicu-

ous (Fig. 9A). On 20 November 2016, we captured an

adult Dendropsophus leali (Hylidae) at approximately

0930 h, sheltering under leaf litter that had been placed

atop a funnel trap in floodplain forest; the animal was

placed in a plastic bag and retained for processing. The

frog was observed alive at 0830 h the following day, but

had died with mouth agape by 1130 h. Upon removing

the frog from the container, a large dipteran larva exited

the mouth of the frog; the larva was approximately 8 mm

long (versus 20 mm SVL for the frog). Efforts were made

to rear the larva; however, the adult escaped following

pupation. Immediately following the exit of the larva, the

skin around the thoracic and abdominal region of the frog

appeared to be shrunken, as though significant visceral

loss had occurred (Fig. 9B). The frog was preserved as

a specimen and deposited in the University of Michigan

Museum of Zoology (UMMZ 246153). This case is no-

table relative to other myiases reported for frogs, because

no external or cutaneous lesions were noted, as is charac-

Amphib. Reptile Conserv.

teristic for many flesh fly infections, and because of the

large size of the larva relative to the frog. In addition to

these two tree frogs, we observed one case of fly myiasis

in the poison frog Ranitomeya uakarii (Dendrobatidae).

On 20 January 2017, at 1230 h, we observed an indi-

vidual of R. uakarii hopping on the leaf litter; we cap-

tured it and placed it in a plastic bag, and took it to the

lab to process it later the same day. When we returned

from fieldwork around 1630 h, the frog was dead and

we noticed a maggot that periodically emerged from a

small lesion on its back (Fig. 9C); a video of the inter-

action is archived in the University of Michigan Deep

Blue Data repository (do1.org/10.7302/Z2862DP1). This

observation took place in terra firme forest near Madre

Selva Biological Station (3°37°14.8’S, 72°14°48.5”W),

Loreto region, northern Peru. The host exhibited a small

round wound on the back, similar to those observed in

other poison frogs (Hagman et al. 2005).

Commensalism between Spiders and Frogs

In addition to the predation and parasitoid infection

events described above, and countless more that take

place every day in the rainforest, a more congenial rela-

tionship between spiders and frogs exists in southwest-

ern Amazonia. This relationship involves a tarantula and

a narrow-mouthed frog that uses the same retreat site

used by the spider. The spider was originally identified

as Xenesthis immanis (Theraphosidae) and the frog as

Chiasmocleis ventrimaculata (Microhylidae), respec-

tively (Cocroft and Hambler 1989). Field observations

and experiments showed that the spider 1s able to capture

and eat several other frog species, while it rejects C. ven-

trimaculata (Cocroft and Hambler 1989; Csakany 2002).

Here we update the identification of both participants in

this interaction, based on our observations at Los Ami-

gos Biological Station. The spider belongs to the genus

Pamphobeteus (Theraphosidae; Fig. 10) and the frog is

Chiasmocleis royi (Microhylidae; Fig. 10), a species de-

scribed recently (Peloso et al. 2014). If an individual of

February 2019 | Volume 13 | Number 1 | e169

von May et al.

> es

Fig. 10. A more congenial relationship: the spider Pamphobeteus sp. (Theraphosidae) and Chiasmocleis royi. Photo by Emanuele

Biggi.

C. royi approached or was presented to young or adult — of C. royi around a different spider burrow occupied by

individuals of Pamphobeteus, the spiders always felt the | 6-8 spiderlings. The frogs were 5—50 cm from the en-

air movement and tried to grasp the frog as it wasa prey, — trance of the burrow. We captured and measured six of

but they always released it without even trying to bite the — these juvenile C. royi (SVL in mm were as follows: 14.6,

frog (Cocroft and Hambler 1989; Biggi and Tomasinelli, 15.7, 14.6, 14.6, 16.2, 14.9).

pers. obs.).

Similar associations between spiders and frogs have been

In one evening in early March 2008, right after dusk set, | documented in North America, India, and Sri Lanka. In

we observed the emergence of three C. royifromasingle | North America, Blair (1936) described the association

burrow (first ones to emerge), followed by the emergence involving another species of narrow-mouthed frog, Gas-

of more than 10 spiderlings (second group), and, lastly,an —_ trophryne olivacea (Microhylidae), and the spider Apho-

adult female Pamphobeteus. The frogs dispersed through = nopelma hentzi (Theraphosidae). Field observations

an area surrounding the burrow (<2 m7) while all spiders = showed that one or more frogs may occupy a spider bur-

stayed closer to the burrow (E. Biggi and F. Tomasinelli, | row and the resident spiders did not prey upon the frogs

pers. obs.). The female Pamphobeteus appeared to react (Blair 1936). Subsequent studies of G. olivacea showed

to minimum air movement more rapidly than the spider- __ that the frogs also occupy burrows used by rodents, and

lings. In addition, we saw multiple individuals of C. royi researchers hypothesized that the underground micro-

near other spider burrows. This observation took place in _— habitats probably protect the frogs from desiccation in

the floodplain forest, at a site located approximately 1.5 = xeric environments (Fitch 1956; Hunt 1980). Another

km from the station. Furthermore, on 8 March 2008, we association involves the Tungara Frog Engystomops

found five individuals of C. royi next to a spider burrow — pustulosus (Leptodactylidae) and a theraphosid spider

with multiple entrances; the first and second individual (Powell et al. 1984). As in the previous case, researchers

of C. royi were <2 cm from two separate entrances, anda — found multiple frogs inside spider burrows and observed

third individual of C. royi was ca. 15cm fromathirden- _ that the spiders did not prey upon the frogs. Powell et

trance. This third entrance led to a section of the burrow al. (1984) hypothesized that skin secretions probably

occupied by at least two Pamphobeteus spiderlings. We — deter the spiders from predating the frogs. In India, the

captured and measured one adult and three juvenile C. —_ narrow-mouthed frog Uperodon taprobanicus (Microhy-

royi (SVL inmm were as follows: 19.9, 13.3, 13.2,15.2). __ lidae) has been observed in tree holes used by the thera-

This observation also took place in the floodplain forest, | phosid spider Poecilotheria hanumavilasumica (Siliwal

at a site located approximately 1.5 km from the station. | and Ravichandran 2008). On multiple night surveys con-

In the same evening, we found seven juvenile individuals § ducted in Rameshwaram Island, the researchers never

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Ecological interactions between arthropods and small vertebrates

a z Pal f

Fig. 11. Juvenile Pamphobeteus sp. infested by fly larva

observed P. hanumavilasumica preying upon the frogs

(Siliwal and Ravichandran 2008). In Sri Lanka, a closely

related frog species, Uperodon nagaoi (Microhylidae),

has been observed in tree holes used by two species of

theraphosid spiders, Poecilotheria ornata and Poeciloth-

eria cf. subfusca (Karunarathna and Amarasinghe 2009).

These researchers observed the frogs and the spiders

sharing the same tree holes on multiple occasions, yet no

predation events were recorded.

In their study, Cocroft and Hambler (1989) proposed

the hypothesis that chemical defenses in the skin of C.

royi prevent spider predation and suggested that the frog

might be a commensal because it obtains protection

against predators. In this study, we observed that some

spiderlings had fly larvae attached to their bodies (Fig.

11; E. Biggi and F. Tomasinelli, pers. obs.). These ob-

servations prompted some questions. Is the frog’s pres-

ence in the burrow beneficial to the spiders? Are the frogs

keeping the number of pests inside the nest in check? If

the frog had chemical defenses, are these defenses se-

creted by skin glands (e.g., skin alkaloids), or do they

have a dietary origin, or are they produced by the micro-

biome on the frog’s skin? All of these hypotheses remain

untested.

Conclusion

The collection and dissemination of natural history data

are critical for understanding how invertebrate predators

and parasitoids impact small vertebrates communities in

tropical rainforests. Future studies aimed at quantifying

the frequency of ecological interactions (e.g., predation,

Amphib. Reptile Conserv.

commensalism) involving arthropods and small verte-

brates across tropical and temperate forest habitats will

shed light on patterns of commonness and rarity of these

organisms among regions, and their effect on the struc-

turing and functioning of food webs.

Acknowledgements.—We thank the Amazon Con-

servation Association and the staffs at Los Amigos and

Villa Carmen biological stations for facilitating our work

at the stations. We also thank Project Amazonas and the

staff at Madre Selva Research Station for assisting us in

our work at that station. Field research was supported by

a fellowship from the David and Lucile Packard Founda-

tion (to DLR), and by the Amazon Conservation Associa-

tion (to RVM), the Wildlife Conservation Society (RVM),

Rosemary Grant Award from the Society for the Study of

Evolution (JGL), Edwin C. Hinsdale UMMZ Scholarship

(JGL), and University of Michigan startup finds (ARDR).

Research and collecting permits were issued by the Insti-

tuto Nacional de Recursos Naturales (INRENA), the Di-

reccion General Forestal y de Fauna Silvestre (DGFEFS),

and the Servicio Nacional Forestal y de Fauna Silves-

tre (SERFOR), Peru (R.D. 11-2008-INRENA-IFFS-

DCB, 120-2012-AG-DGFFS-DGEFFS, 064-2013-AG-

DGFFS-DGEFFS, 292-2014-AG-DGFFS-DGEFFS,

R.D.G. —_029-2016-SERFOR-DGGSPFFS, R.DG.

405-2016-SERFOR-DGGSPFES). We thank I. Holmes,

M. Cowan, I. Russell, P. Cerda, T.Y. Moore, J.C. Cusi,

E.S. Vargas Laura, C. Macahuache Diaz, R. Villarcorta

Diaz, E. Durand Salazar, E.M. Iglesias Antonio, N. Ta-

fur Olortegui, O.L. Huacarpuma Aguilar, and Y. Casa-

nca Leon for assistance in field data collection. We also

February 2019 | Volume 13 | Number 1 | e169

von May et al.

thank R. Voss (American Museum of Natural History)

and J. Patton (Museum of Vertebrate Zoology) for their

help with the opossum indentification. We thank Jaime

Villacampa, Chris Beirne, and one anonymous reviewer

for providing constructive comments on the manuscript.

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Appendix 1. Summary of predator-prey interactions reported in this study.

Arthropod Predator Vertebrate Prey

Se a Family Deena as Family eine

Ancylometes sp. Ctenidae Dendropsophus leali Hylidae 1A

Phoneutria sp. Ctenidae Hamptophryne boliviana Microhylidae 1B

Thaumasia sp. Pisauridae Tadpole N/A (Anura) 2A

Ctenid spider sp. Ctenidae Boana sp. G Hylidae 2B

Theraphosid spider Theraphosidae Hamptophryne boliviana Microhylidae 3A

Ctenid spider Ctenidae Leptodactylus didymus Leptodactylidae 3B

Ctenus sp. Ctenidae Cercosaura eigenmanni Gymnophthalmidae 4

Centipede Scolopendridae Dipsas catesbyi Colubridae SA

Centipede Scolopendridae Micrurus obscurus Elapidae 5B

Pamphobeteus sp. Theraphosidae Marmosops cf. noctivagus Didelphidae 6

Ctenid spider Ctenidae Hamptophryne boliviana Microhylidae 7A

Ancylometes sp. Ctenidae Dendropsophus sarayacuensis Hylidae yas:

Water bug Belostomatidae Dendropsophus minutus Hylidae Te

Trigona sp. Apidae Frog eggs Hylidae 8A

Phoneutria sp. Ctenidae Dendropsophus kamagarini Hylidae 8B

Amphib. Reptile Conserv.

February 2019 | Volume 13 | Number 1 | e169

Amphib. Reptile Conserv.

von May et al.

Rudolf von May is a Postdoctoral Research Fellow at the Museum of Zoology and the Department

of Ecology and Evolutionary Biology at the University of Michigan. His current research seeks to

understand how biological communities are structured across habitats and elevations, with special focus

on amphibians and reptiles living in the Andes-Amazon region.

Emanuele Biggi is an Italian naturalist with a Ph.D. in Environmental Sciences, Associate Fellow of the

International League of Conservation Photographers, and focuses on the smaller creatures, conservation

of nature, and science-at-work photography. . Emanuele is the author and curator of scientific expositions

that bring nature to people, and raise awareness about the natural world. Website: www.anura. it

Heidy Cardenas is a Research Associate at Area de Herpetologia del Museo de Historia Natural (MUSA)

de la Universidad Nacional de San Agustin de Arequipa, Peru. Her current research interests include

taxonomy, natural history, and conservation of reptiles

M. Isabel Diaz has a B.S. degree in biological sciences and is associated with Natural History Museum of

Universidad Nacional San Antonio Abad del Cusco, Peru (MHNC) and Museum of Biodiversity of Peru

(MUBI). Her research interests include taxonomy, evolution, ecology, and conservation of amphibians

and reptiles.

Consuelo Alarcén Rodriguez is a graduate student at the Department of Biology at John Carroll

University. Her research interests are distribution and diversity of amphibians and reptiles in Peru. She is

currently studying contemporary and paleodistribution of Pseudoboine snakes.

Valia Herrera Alva is a Peruvian biologist and a researcher at the Herpetological Collection of the

Museum of Natural History of the Universidad Nacional Mayor de San Marcos (MUSM). Her research

interests include ecology, conservation, and physiology of amphibian and reptiles.

Roy Santa-Cruz is a Research Associate at Area de Herpetologia del Museo de Historia Natural (MUSA)

de la Universidad Nacional de San Agustin de Arequipa, Peru. His current research interests include

taxonomy, natural history, and conservation of amphibian and reptiles. Roy currently coordinates several

research projects focusing on threatened species of Andean frogs.

Francesco Tomasinelli has a degree in Environmental Science and works as a freelance biologist,

journalist, and photographer. As an ecologist he conducts surveys and assessments on local fauna and

flora in agricultural and semi-urbanized areas. As a photojournalist he joined several scientific expeditions

in the tropics and covered the activity of conservation and environmental programs in Italy and abroad.

He currently creates exhibits for museums, involving live invertebrates, photos, and activities for visitors.

Website: www. isopoda.net

Erin P. Westeen is a graduate student at the University of California, Berkeley, and was previously

a post-graduate researcher at the Museum of Zoology and Department of Ecology and Evolutionary

Biology at the University of Michigan. Her interests are in the evolution of squamate reptiles and has

recently completed a study focusing on jaw morphology of opisthoglyphous snakes.

76 February 2019 | Volume 13 | Number 1 | e169

Amphib. Reptile Conserv.

Ecological interactions between arthropods and small vertebrates

Ciara M. Sanchez-Paredes is a Peruvian biologist associated with the Laboratory of Studies in

Biodiversity from Universidad Peruana Cayetano Heredia. Her research interest is the impact of human

interactions in animal wellbeing from the perspective of environmental management and sustainable

development.

Joanna G. Larson is a graduate student at the Department of Ecology and Evolutionary Biology and the

Museum of Zoology at the University of Michigan. She is interested in understanding the processes that

have shaped the distribution of amphibian species richness across space and clades. Her studies integrate

phylogenetic, ecological, phenotypic, and dietary data.

Pascal O. Title is a postdoctoral research fellow at Indiana University, and he recently completed his

doctoral studies at the Museum of Zoology and Department of Ecology and Evolutionary Biology at the

University of Michigan. He is interested in the spatial distribution of diversity, and how that may or may

not reflect evolutionary history and landscape effects on diversification.

Maggie R. Grundler is a graduate student at the University of California, Berkeley, and she was previously

a post-graduate researcher at the Museum of Zoology and Department of Ecology and Evolutionary

Biology at the University of Michigan. She is interested in the evolution of diet and ecological speciation

in reptiles and amphibians.

Michael C. Grundler is a graduate student in the Museum of Zoology and the Department of Ecology

and Evolutionary Biology at the University of Michigan. He is interested in the natural history of

squamate reptiles and in the evolution of their diverse trophic strategies.

Alison R. Davis Rabosky is an Assistant Professor and Curator of Herpetology in the Museum of

Zoology at the University of Michigan. She uses interdisciplinary approaches that combine ecological

physiology, behavioral ecology, functional genomics, and macroevolutionary comparative analysis to

understand the origin and stability of novel phenotypes in nature.

Daniel L. Rabosky is an Associate Professor and Curator of Herpetology in the Museum of Zoology

at the University of Michigan. He studies macroevolution, global diversity gradients, the ecology and

evolution of squamate reptile communities, and conducts fieldwork on reptiles and amphibians in

Australia and the Neotropics.

Fk, February 2019 | Volume 13 | Number 1 | e169

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

13(1) [General Section]: 78-89 (e164).

Description of head scalation variation, hemipenis,

reproduction, and behavior of the Indian Smooth Snake,

Coronella brachyura (Gunther 1866)

Dikansh S. Parmar

Department of Biosciences, Veer Narmad South Gujarat University, Surat, Gujarat, INDIA

Abstract.—The first confirmed record of the Indian Smooth Snake, Coronella brachyura, with nine supralabials

on both sides is reported from Indian state Gujarat, with taxonomic and morphological details, and information

on etymology, behavior, diet, hemipenis, report of gravid female with eggs, and distribution of species.

Information provided is based on examination of three individuals, two live and one dead. Present study

includes details of examined individuals’ data and observations, along with references to published literature.

Keywords. Reptilia, Squamata, Serpentes, Colubridae, rare, endemic, taxonomy, Gujarat, Surat

Citation: Parmar DS. 2019. Description of head scalation variation, hemipenis, reproduction, and behavior of the Indian Smooth Snake, Coronella

brachyura (Gunther 1866). Amphibian & Reptile Conservation 13(1) [General Section]: 78-89 (e164).

ternational (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in any medium,

provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced, are as follows:

Official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org.

Received: 10 August 2017; Accepted: 05 April 2018; Published: 24 February 2019

Introduction

The Indian Smooth Snake, Coronella brachyura (Gutn-

ther 1866) is an endemic, rare, and harmless Colubridae

with smooth scales, a slender almost tubular body and a

short tail. This snake occurs in two colors: glossy black

or brown dorsally, and uniform creamy white ventrally

(Fig. 1A, B). Small in size, reaching a maximum length

of 640 mm, this snake 1s reclusive and mild-tempered. It

is reported from the Tapi district of India, which shares

a border with the Dangs district (part of Western Ghats,

one of the hotspot regions of biodiversity in the world),

from Surat district in South Gujarat, and from three

other states in India: Maharashtra, Madhya Pradesh,

and Chhattisgarh. From Gujarat, this species was first

obtained by the author in 2006 from the Surat district.

After identification and scale counts were recorded it was

handed over to R. Vyas for further investigation, and he

subsequently deposited it as a voucher specimen in the

museum of Bombay Natural History Society, Mumbai

(BNHS 3407).

The Indian Smooth Snake is often mistaken for the

Wallace’s Striped Snake (Wallaceophis gujaratensis),

Banded Racer (Argyrogena fasciolata), Glossy Bellied

Racer (Platyceps ventromaculatus), or hatchlings of the

Checkered Keelback (Xenochrophis piscator) which

have a dark color or faint checkered pattern. Vyas and Pa-

tel (2007) presented a photograph of Wallaceophis guja-

Correspondence. ophiophagus_hannah10@yahoo.com

Amphib. Reptile Conserv.

ratensis as Coronella brachyura. Later Patel et al. (2015)

reported that specimens identified by Vyas and Patel

(2007) from two localities (Ahmedabad and Bhavnagar)

were incorrect, and the misidentified specimens from

Bhavnagar were Wallaceophis gujaratensis not Coro-

nella brachyura but they stated records from these two

localities remains in question. Then Mirza, Vyas, Patel,

Maheta, and Sanap (2016) established the misidentified

photograph from Bhavnagar locality as Wallaceophis gu-

Jaratensis. In all, the same record was presented as two

different species in three different publications by a total

of seven authors claiming to have identified it by examin-

ing photographs. Among these seven, one mutual author

identified the specimen three times, first as Coronella

brachyura in 2007, then as Wallaceophis gujaratensis

in 2015 (but stating that this record remained in ques-

tion), and finally as Wallaceophis gujaratensis in 2016.

This sequence of events shows the confusion among au-

thors in identifying specimens and localities of Coronella

brachyura and Wallaceophis gujaratensis. However, Pa-

tel et al. (2015) stated that specimens from two localities

(Ahmedebad and Bhavnagar) remained in question, but

the current author (DSP) identified these specimens from

the published photographs. Photograph labeled in Sauria

as “Fig. 2 The specimen of Coronella brachyura from

Sagwadi, Bhavnagar, Gujarat” is here confirmed as Wal-

laceophis gujaratensis based on morphological features:

absence of crown-like mark on head at parietal region,

February 2019 | Volume 13 | Number 1 | e164

Parmar

1 Mag

presence of two distinct dark lateral stripes, and reddish

iris (not visible in the published photograph). Photograph

labeled in Sauria as “Fig. 3 The specimen of Coronella

brachyura from Jodhapur [sic], Ahmedebad, Gujarat” is

here confirmed as Coronella brachyura based on mor-

phological features: upper edges of supralabials marked

with black streak from first supralabial on snout to supra-

labials in contact with eye, indistinct dark stripes on lat-

eral sides, crown like pattern on head seen only on sharp

observation, two thin parallel indistinct lines on verte-

bral region from head (right behind the parietal shields)

to neck, morphology of scales on body and eye entirely

black.

In the literature, Coronella brachyura is considered

poorly-known. Herein are reported additional data and

observations on etymology, taxonomy, morphology, be-

havior, diet, reproduction, habitat, distribution, and scale

variations in this poorly-known species.

Methods

Data are based on three individuals: Two were live cap-

tured individuals and one was found dead on the road

(roadkill). Recovered individuals are mentioned as A, B,

>" f Caps I

“nee

‘pees

and C, according to dates and years of their recovery (Ta-

Fig. 1. Two colors of Coronella brachyura. (A) Black color. (B) Brown color. Photo credit Dikansh S. Parmar

ble 1). Individual A was found on | January 2008 from

the Department of M.Sc. (IT), Veer Narmad South Guja-

rat University at 0730 hours. Individual C was found on

16 November 2010 from Piplod area Saru Nagar Society

at 1600 hours. Individuals A and C were live female indi-

viduals. Road kill individual B was found on 8 June 2010

from University road at 2200 hours, and was male (Fig.

2A, B). Equipment used in taking data includes: a stan-

dard ruler to the nearest mm, magnifying lens, vernier

caliper, pen, record book, string, and a camera. Ventral

scale counts were taken according to the method pro-

posed by Dowling (1951). Subcaudals were counted on

one side by excluding the terminal scute.

Sex determination was completed using the pop-

ping method (Gregory 1983). In this method the thumb

is placed under the cloaca, applying gentle pressure to-

ward the vent. Gentle pressure 1s applied with the other

thumb moving from the tip of the tail towards the cloaca.

Squeezing it gently toward the cloaca the hemipenis of

the male pops out by the increased internal pressure in

the base of the tail. This technique should be done only

by an expert due to the risk of damaging the sexual or-

gans.

Snout to vent length and tail length were measured

by marking the length on a piece of string. Snout to vent

a Z . ay

Fig. 2. (A) Male individual found in road kill. (B) Close up of its hemipenis. Photo credit Dikansh S. Parmar.

Amphib. Reptile Conserv.

February 2019 | Volume 13 | Number 1 | e164

Indian Smooth Snake (Coronella brachyura)

length was measured up to the posterior margin of the

anal plate. The last infralabial is defined as the infralabial

still completely covered by the last supralabial. Numbers

of dorsal scale rows were counted at one head length be-

hind the head, at midbody, and at one head length before

the tail. Measurements and scale counts taken to describe

the scalation and characteristics were: total body length

(TBL), snout to vent length (SVL), tail length (TL), dor-

sal scales (DS), ventrals (V), subcaudals (SC), supral-

abials (SL), infralabials (INL), right and left side (RL),

number of supralabials in contact with eye (SLCE), Su-

praoculars (SO), preoculars (PRO), postoculars (PO),

loreals (L), supralabials in contact with loreal (SLCL),

temporals (T), and anal plate (AP).

Results

Etymology: Coronella brachyura. Coronella is a Latin

word meaning “small crown,” in reference to the pattern

around the head of snake. Brachyura is modern Latin

from the Greek words brachys, “short,” and oura, “tail”

(Wiktionary 2017). The full name means a snake with a

small crown-like pattern on head and a short tail. Its ver-

nacular name in Gujarati language is Suvaro saap (Desai

2011, 2017). Suvaro means “smooth” or “silky” and saap

means “snake.”

Taxonomy: This species belongs to the family Colub-

ridae, subfamily Colubrinae, genus Coronella, and spe-

cies C. brachyura. Scientific name is binomial Coronella

brachyura, and a synonym is Zamenis brachyurus Gin-

ther, 1866. Coronella brachyura was described by Gin-

ther in 1866 as Zamenis brachyurus (Smith 1943) based

on specimens collected from Western Ghats in Poona

(Pune) district of Maharashtra (Smith 1943). Coronella

is closely related to Lampropeltis, the American king

snakes. Both groups were once classified within the same

genus, then separated later and placed in different genera

(Mehrtens 1987). Hoser (2012) separated Wallophis from

the genus Coronella on the basis of phylogenetic studies

and by the presence of 23 rows of scales on dorsal mid-

body and triangular frontal scale. Hoser placed C. brachy-

ura within a monotypic genus Wallophis. Similarly, after

Hoser (2012), Mirza and Patel (2017) transferred the ge-

nus Coronella to Wallophis on the basis of phylogenetic

study. This phylogenetic analysis was based on partial

sequencing of nuclear and mitochondrial genes using a

road kill specimen. The scalations of supralabials, num-

ber of supralabials touching eye, temporals, ventrals, and

subcaudals described by Mirza and Patel (2017) in diag-

nosis vary from the scalations described in this present

study. Patel et al. (2015) mentioned eight to nine supral-

abials in Coronella brachyura, but have not provided any

individual with nine supralabials on both sides and fifth

and sixth supralabials touching eye on both sides. Patel

et al. (2015) provided evidence of: eight supralabials on

both sides, eight supralabials on one side and nine supra-

Amphib. Reptile Conserv.

labials on other side, fourth and fifth supralabials touch-

ing eye on both sides, and fourth and fifth supralabials

touching eye on one side and fifth and sixth supralabials

touching eye on other side. Later on in diagnosis, Mirza

and Patel (2017) mentioned eight to nine supralabials and

fifth and sixth supralabials touching eye rarely, without

any photographic evidence or data of any individual with

nine supralabials on both sides and fifth and sixth supra-

labials on both sides, neither have they provided phylo-

genetic analysis of any specimen with these variations.

This type of variation in supralabials and number of su-

pralabials touching eyes on left and right side have been

noted by author (DSP) in many snakes, including: Com-

mon Trinket Snake, Coelognathus helena helena, Glossy

Marsh Water Snake, Gerarda prevostiana, Common

Vine Snake, Ahaetulla nasuta, Common Wolf Snake,

Lycodon aulicus, and Barred Wolf Snake, Lycodon stria-

tus. This type of variation may occur due to mutations,

breaking of scales, injury at birth to hatchling emerging

from its egg, or variations in temperature during incuba-

tion (Parmar 2017). Variations in scalations for example

right side (eight) and left side (nine or more) are common

in many species, some of them are mentioned above but

specimens having variations with equal numbers of sca-

lations for example right side (nine) and left side (nine)

are rarely found. However, Mirza and Patel (2017) have

mentioned eight to nine supralabials and fifth and sixth

touching eye in diagnosis without presenting a record of

any individual with such variations on both sides. More-

over, Patel et al. (2015) did not accept the allocation of

the genus Wallophis by Hoser (2012) and commented

that Hoser (2012) did not provide any valid taxonomic

characters to support partitioning the genus Coronella.

The case is the same with Mirza and Patel (2017) as they

have not yet reported phylogenetic analysis or morphol-

ogy and scale counting studies of individuals with nine

supralabials on both sides, fifth and sixth supralabials

touching eye on both sides, two + one temporals, and

greater numbers of ventrals and subcaudals. The Reptile

Database website has posted a comment (viewed on 13

Jul 2018) on this issue that:

“Synonym: Wallophis was revived in a non-peer re-

viewed journal (Litteratura Serpentium), without ad-

equate justification (by Ken Welch) and more recently

by Mirza and Patel 2017. Although Mirza and Patel

provided a diagnosis for Wallophis, they did not in-

clude any other members of Coronella in their mo-

lecular analysis despite the availability of DNA se-

quences, hence we do not recognize the genus for the

time being.”

According to us in order to accept this replacement of

genus more evidence and detailed studies of additional

specimens from India are required. Required also would

be molecular analysis of other members of Coronella due

to the many morphological and scale count variations in

February 2019 | Volume 13 | Number 1 | e164

Parmar

Table 1. Details and measurements (mm) of Coronella brachyura, individuals A, B, and C.

Individuals A

1 January 2008

M.Sc. (IT) Surat

Date of collection

Name of locality

Gender F

TBL 458

SVL 395

TL 63

DS PREP te)

Vv 241

AP Entire

SC 48

SL RL- 9

INL RL- 10

SLCE RL- 2 (5" and 6")

SO 1

PRO y

PO )

1 1

SLCL RL- 3 (2"4, 3, and 4")

T 242

the species. In order to change a genus a complete study

of species is required. A genus should not be changed

on the basis of variation in one single individual, par-

ticularly in rare, poorly known and a species with highly

variable scalations in head (labials, oculars, temporals,

loreals, etc.), dorsal body, ventrals and subcaudals. Mirza

and Patel (2017) mentioned the variations in supralabi-

als and number of supralabials touching eye (without

providing any individual with such variations on both

sides). However, in individuals with nine suprlabials

on both sides and fifth and sixth supralabials touching

eye on both sides, the author has found that unreported

variations in suprlabials, number of supralabials touch-

ing loreal, postoculars and temporals. Therefore, it is not

possible to accept allocation of genus Wallophis without

a detailed comparison of morphological and genetic dif-

ferences in individuals with both normal and varied sca-

lation. Only after all individuals with normal and varied

scalations are evaluated should any appropriate changes

to the genus be considered.

Morphology and coloration: Details of all three in-

dividuals are given in Table 1. These snakes are small,

glossy black- or brown-colored, tubular or round-bodied

with uniform creamy white belly and large ventrals. Head

is slightly broader than neck, less wide and more elongat-

ed, with round eye pupil. Eye entirely black. Short tail 1s

less than one sixth of the total length, pointed at the end.

Snout 1s not tapered, but slightly round. Nostril occupies

whole depth of the sutures between nasals. Supralabi-

als are distinctly white or pale white, or sometimes light

Amphib. Reptile Conserv.

RL- 2 (4 and 5")

B C

8 June 2010 16 November 2010

University Road Surat Piplod Surat

M F

469 49]

396 427

73 64

23:23:19 Zei2a49

219 215

Entire Entire

50 57

RL- 8 RL- 9

RL- 10 RL- 10

RL- 2 (5" and 6")

=S NO Fe

RL- 2 (2™ and 3")

RL- 3 (2%, 3 and 4")

242

brown in color. Upper edges of supralabials are marked

with a black streak, which begins from the first supral-

abial on snout and ends on the first supralabial in contact

with the eye. Scales on dorsal body are often edged with

pale white or white dots, displaying a slightly checkered

pattern on the forebody. Two indistinct thin parallel lines

(appears in the from of dots or short dashes) on nape are

usually present. Indistinct or faint ventrolateral stripes on

body run from right behind the eye to the end of the tail.

Scalations: According to data cited in the literature, mid-

body scales are in rows of 23:23:19 and 200-224 ventral

scales. Anal plate entire. Paired subcaudals 46-53. Eight

supralabials (fourth and fifth touching eye), one loreal,

one preocular, two postoculars, and two + two temporals

(Whitaker and Captain 2004, 2008). Scalation of individ-

uals examined here differs from the scalation mentioned

above. Individual A has 241 ventrals, more than recorded

in the literature (Whitaker and Captain 2004; Vyas and

Patel 2007; Patel et al. 2015; Mirza and Patel 2017), and

individual C has 57 subcaudals more than documented

in the literature (Whitaker and Captain 2004, 2008; Vyas

and Patel 2007; Patel et al. 2015; Mirza and Patel 2017).

Rescued individuals also have one side with nine supra-

labials, fifth and sixth touching eye, and the other side

with eight supralabials, fourth and fifth touching eye. In

the present study, individuals A (Fig. 3A, B) and C (Fig.

4A, B) have nine supralabials on both sides, fifth and

sixth supralabials touching eye on both sides, while in-

dividual B (Fig. 5A, B, C) has eight supralabials on both

sides, fourth and fifth touching eye on both sides. The in-

February 2019 | Volume 13 | Number 1 | e164

Indian Smooth Snake (Coronella brachyura)

—_— oe

Fig. 3. Individual A, both sides nine supralabials, fifth and sixth supralabials in contact with eye, and second, third, and fourth

e vit tt ; : ;

an : a

"* RAN YS = a ee of .

= oh : 1@ SP es : ey |

7 4

supralabials in contact with loreal scale. (A) Left side. (B) Right side. Photo credit Dikansh S. Parmar.

dividual (B) with eight supralabials on both sides has the

fifth supralabial in contact with the temporal and postoc-

ular, while the individuals with nine supralabials on both

sides have the sixth supralabial in contact with the tem-

poral and postocular. Individuals in this species have one

preocular, but individual A has variations in preoculars,

with two preoculars instead of one. At first, the left side

preocular looked different from the right. However, on

close observation with a magnifying lens, the presence of

two preoculars was confirmed on both sides in individual

A. Loreal is in good contact with postnasal, prefrontal,

preocular, and second and third supralabials in individual

(B) with eight supralabials on both sides. In individuals

(A and C) with nine supralabials, loreal is in good con-

tact with postnasal, prefrontal, and preocular, but differs

by touching three supralabials: second, third, and fourth,

instead of only second and third supralabials. Loreal is

not in contact with internasal in any case and is distinctly

separated by contact of postnasal and prefrontals. Frontal

is triangular shaped. Rostral is broad and high in contact

with first supralabial, prenasal, and internasal. The indi-

vidual obtained in 2006 (and subsequently handed over

to R. Vyas) had variations in temporals, dorsals and ven-

trals: it had temporals two + one instead of two + two,

—

ed a

. = ai Mog

“ii i

Fig. 4, Individual C, both sides nine supralabials, fifth and sixth supralabial

dorsal midbody scales 23:23:21 versus 23:23:19 and 237

ventrals instead of 224, more than noted in the published

literature (Whitaker and Captain 2004, 2008). Reduction

in midbody scales are also marked with 23:23:17 dorsal

midbody scales (Patel et al. 2015).

Behavior and diet: The Indian Smooth Snake is a cre-

puscular burrowing snake (Whitaker and Captain 2004,

2008), but is often seen active during the day time. It is

terrestrial, but 1t can climb well. It lives under stones, brick

piles, and trees, but is also found in open-ground arid areas

and grasses. It does not bite on handling, as it is a mild-

tempered snake, but it hunts fiercely. As an active hunter

with constricting abilities, it coils around its prey (Fig. 6).

Adults and juveniles of Hemidactylus brookii were offered

to captive individual A, which readily preyed upon them.

This snake 1s active throughout the year, but it is more ac-

tive from late winter to early monsoon season, in hotter

days between February and June. Diet consists of geckos

and new-born garden lizards and skinks (Whitaker and

Captain 2008). Apart from these, frogs were offered to a

captive individual, but it did not show any interest in hunt-

ing frogs (Fig. 7), thus frogs are excluded from diet.

, ; ca, ae i

TR Sa oa me a

Ss in contact with eye, and

supralabials in contact with loreal scale. (A) Left side. (B) Right side. Photo credit Dikansh S. Parmar.

Amphib. Reptile Conserv.

February 2019 | Volume 13 | Number 1 | e164

Parmar

Table 2. New distributional records of Coronella brachyura in India. Localities in map are shown state-wise in numerals from 1

to 16.

Serial Number State Localities in Map

l Maharashtra 1

2 Maharashtra 2

3 Maharashtra 3

4 Maharashtra 4

5 Maharashtra 5

6 Maharashtra 6

7 Maharashtra 7

8 Maharashtra 8

9 Maharashtra 9

10 Maharashtra 10

11 Gujarat 11

12 Gujarat 12

13 Gujarat 13

14 Madhya Pradesh 14

15 Madhya Pradesh 15

16 Chhattisgarh 16

Reproduction: Snakes and lizards are classified in the

order Squamata (scaled reptiles). For the first time a grav-

id female was obtained from Indian state Gujarat, district

Surat, Taluka Oldpad. It laid four eggs on 15 April 2017

(Fig. 8). Length of female was 640 mm. Measured size

of eggs in mm were length x width a- 31.3-12.5, B - 32.3-

13.4, y - 31.6-12.7, 6 - 33.4-13.9. Eggs were marked with

red pen by rescuers in order to make note of proper side

and were kept for artificial incubation at room tempera-

ture. Laid eggs started shrinking and appeared to be af-

fected by hypoosmosis in early days of incubation (Par-

mar 2018). On 8 June 2010, the first authentic record of a

male was made in Gujarat, district Surat. Male squamates

have intromittent organs which function to deliver sperm

during copulation. According to an earlier description of

a damaged specimen, hemipenes in this species extends

up to 13th caudal plate, the distal half of hemipenes is

calyculate, the cups large with scalloped edges, and the

proximal half spinose (Boulenger 1890). Roadkill indi-

vidual obtained by us was sexed using the popping meth-

od. It has a 10 mm long red, spiky, and unforked hemipe-

nis with visible su/cus spermaticus. The spikes are in two

colors: white dorsally and red from mid to shaft. Two or

three spikes at the base are much larger than others (Fig

2A, B). The spikes on hemipenes help males to fix the

hemipenis in place during mating and the sulcus sper-

maticus 1s a groove that helps to conduct sperm into the

female during copulation. Size and shape of hemipenis

varies from species to species, and male hemipenis and

female cloacas are compatible with each other accord-

ing to species. If males have spiky hemipenis, females

of the same species have thicker cloacal walls compared

to those of species with males with less spikes or no

spikes on hemipenis. For species of males with branched

Amphib. Reptile Conserv.

District Locality Coordinates

Pune Pune L831 Ne 73:01 -E

Pune Talegaon 18.72°N, 73.68°E

Yavatmal Wani 20.03°N, 78.57°E

Solapur Kurduwadi 18.08°N, 75.43°E

Ahmednagar Visapur 18.48°N, 74.35°E

Nashik Nashik 20.00°N, 73.78°E

Amravati Melghat 2). 26°N 77 er,

Latur Latur 18.23°N, 76.36°E

Pune Khed 18.56°N, 73.43°E

Jalna Jalna 19.83°N, 75.88°E

Surat Surat 2) ONS 73837

Tap Tap ANQEN, /3760°R

Ahmedabad Ahmedabad D3202°N 25 7 r

Bhopal Bhopal 235 SPN GFE25 7 E

Ujjain Ujjain 23.10°N, 75.47°E

Rajnandgaon Rajnandgaon 21.09°N, 81.03°E

hemipenis, females have branched cloacas. These similar

structures of hemipenis and cloacas result in a lock and

key mechanism during mating which ensures that mating

occurs only between a male and female of the same spe-

cies (see Gunther 1866 and Dufour 1844).

Distribution: Current type locality, Poona (Pune), was

first discovered by Gunther (1866). The second specimen

was reported by Blanford (1870) near Wun in southeast-

ern Berar. Later records were provided from Anderson

(1871), Theobald (1876), Boulenger (1890), Scarlet

(1891), Wall (1923), Lindberg (1932), Gharpurey (1935),

Smith (1943), Whitaker and Captain (2004), Mistry

(2005), Vyas and Patel (2007), Nande and Deshmukh

(2007), Shyam Kamble and Rahul Deshmane (2009),

Ingle and Sarsavan (2011), Ghadage et al. (2013), and

Patel et al. (2015). This species was previously found in

three states: Gujarat, Maharshtra, and Madhya Pradesh.

Now tt is also reported in 2015 from the Churtya block of

Rajnandgaon district in Chhattisgarh state by Nova Na-

ture Welfare Society (Fig. 9) [Table 2]. Records of this

Species are from three districts and twelve localities in

Gujarat, eight districts and ten localities in Maharashtra,

two districts and two localities in Madhya Pradesh, and

one district and one locality in Chhattisgarh, as shown

in the distributional graph of this species (Fig. 10).

One of the reasons behind the distribution of this spe-

cies in only four states could be the location of this area

around the Tropic of Cancer, which crosses the country

at 23.43701°N, as well as the type of forests in the area.

The Tropic of Cancer passes through eight states in India.

Among these eight states, the species is reported from

three states crossed by the Tropic of Cancer: Gujarat,

Madhya Pradesh, and Chhattisgarh. As this snake is found

February 2019 | Volume 13 | Number 1 | e164

Indian Smooth Snake (Coronella brachyura)

= SR ei

ane may

tise

Fig. 5, Individual B, both sides eight supralabials, fourth and fifth in contact with eye, and second and third supralabials in contact

with loreal scale. (A) Left side. (B) Right side. (C) Live individual with same scalation. Photo credit Dikansh S. Parmar.

active mostly in summer or hotter days from February to

June, its population is also found in regions where sun

rays fall straight, in the states of the central western part

of India. Even though the Tropic of Cancer does not pass

through Maharashtra, this species and its type locality are

reported from Maharashtra, possibly because Maharash-

tra shares interstate borders with Gujarat in the north-

west, Madhya Pradesh in the north and Chhattisgarh in

the east. All four states have hilly forest regions. Another

possibility is that the species might have migrated from

Maharashtra to warmer regions of the other three states

that have forests where the climate 1s moderate and more

suitable for the species to survive. Another geographic

factor responsible for the distribution range of this spe-

cies is the presence of rivers. The Tapi and Narmada

rivers flow in three states: Gujarat, Maharashtra, and

Madhya Pradesh. The origin of the river Narmada is at

Amarkantak in Madhya Pradesh, less than 30 km from

the fourth state, Chhattisgarh. The river Tapi originates

from the hills of Mahadev in Madhya Pradesh and flows

through Maharashtra and Gujarat. In 2006 Gujarat was

affected by floods due to heavy rainfall. Surat district

was especially severely affected because the Tapi river

cs Ae

Fig. 6. Coronella brachyura preying upon gecko, coiling

around it. Photo credit Dikansh S. Parmar.

Amphib. Reptile Conserv.

Fig. 8. Eggs of Coronella brachyura in hypoosmotic condition.

Photo credit Vedant Lala.

flows through the central part of the city and meets the

Arabian Sea in the west of the city. Tapi is the only major

river that flows in Surat. Indian Smooth Snake is reported

from parts of Gujarat from 2006 onwards. Along with

the Indian Smooth Snake, another rare species endemic

to India, the Stout Sand Snake, Psammophis longifrons,

distributed in Gujarat and Maharashtra (Whitaker and

Captain 2008), was also obtained for the first time from

Surat district after the floods. The origins of the Tapi and

Narmada rivers are in Madhya Pradesh, and they flow

Fig. 7. Captive individual did not eat frogs when offered. Photo

credit Dikansh S. Parmar.

(axa ey"

February 2019 | Volume 13 | Number 1 | e164

Parmar

Table 3. Distribution of Coronella brachyura in Gujarat state, Surat, Tapi, and Ahmedabad districts, showing number of individuals

and their detailed localities along with abbreviation of localities in map. G1 to G12 indicates localities of species in map of Gujarat.

Localities Number of Total Number

Serial Number District Localities Shown in Map Coordinates Individuals — of Individuals

1 Surat Adajan Gl DL LOSING Te 19 a 1 1

2 Surat Ambheta G2 21.26°N, 72.74°E 1 1

3 Surat Bhatpor G3 21030" NED 952 1 1

4 Surat Bhestan G4 2 LAPS IN29 2.852 E: | 1

5 Surat Oldpad G5 21.34°N, 72.75°E 1 1

6 Surat Piplod G6 215° N, 72 77E 6 6

7 Surat Sachin G7 21.08°N, 72.88°E 1 1

8 Surat ra G8 21.14°N, 72.76°E 1 1

Safal Square

9 Surat VNSGU M.Sc. (IT) G9 21.15°N, 72.78°E 1 1

10 Tapi Ambapani G10 20.97°N, 73.50°E 1 1

11 Ahmedabad Bopal G11 23,02°N,.72:57°E 1 1

2 Ahmedabad Jodhpur G12 23.03°N, 72.40°E 1 1

from Madhya Pradesh through Maharashtra and enter

into Gujarat. In Gujarat, distributional records of the

Indian Smooth Snake species are from Surat, Tapi, and

Ahmedabad districts. Records from Gir Somnath district

need confirmation. One individual was obtained from

a forest near Ambapani village of Tapi district, which

is only a few kilometers from The Dangs district. The

Dangs district shares a border with the Tapi district and

Maharashtra state, thus the species may possibly exist in

The Dangs district. The number of individuals obtained

from Surat district includes: six individuals from Piplod,

one from Veer Narmad South Gujarat University M.Sc.

(IT) Surat, one from Adajan, one from Oldpad, one from

Ambheta, one from University road (Safal square), one

from Bhatpor, one from Bhestan, and one from Sachin

(Table 3). A total of 14 individuals were obtained in 11

years from 2006 to 2017 in and around Surat city.

Discussion

In Gujarat, an Indian Smooth Snake was rescued by the

author (Dikansh S. Parmar) in 2006. It was with normal

scalation (except 237 ventrals, 23:23:21 dorsals, and two

+ one temporals). From 2006 to 2018, the author has col-

lected more complete and accurate information from ev-

ery individual of this species when obtained. In 2008, the

author obtained the first Indian Smooth Snake with nine

supralabials on both sides and two preoculars. In 2010, a

second individual with nine supralabials was obtained. In

total, the author recovered six specimens of this species.

All of them were released to their natural habitat after

recording data. The purpose behind studying this species

is to collect more information regarding the taxonomy,

morphology, activity, diet, behavior, reproduction, and

distribution. These records will be helpful in revealing

Amphib. Reptile Conserv.

the reasons behind the low population of this snake, its

sex ratio, its endemism to India, threats to the species, its

biological importance, and the role of the species in the

environment.

Whitaker and Captain (2004, 2008) and Ingle and

Sarsavan (2011) reported that Coronella brachyura feeds

on juvenile geckos in captivity but Patel et al. (2015) re-

ported keeping live individuals for few days but when

offered juveniles of Hemidactylus sp. they were not ac-

cepted. In the present study, Coronella brachyura readily

fed on either adult or juvenile Hemidactylus brookii. Re-

ported variations in scalation also raise questions about

its evolution, subspecies, ancestors, and distribution in

certain types of localities. A phylogenetic study and DNA

sequencing are required to know the relationship between

the Indian smooth snake with nine supralabials and In-

dian smooth snake with eight supralabials. Our present

study found variations in supralabials, preoculars, num-

ber of scales touching eye, number of supralabials in

contact with loreal, number of supralabials in contact

with postoculars, number of supralabials in contact with

temoprals, ventrals, and subcaudals. These variations in

scalation are signs of a new sister species or subspecies

of Indian smooth snake, which we are looking forward

to discovering by phylogenetic study and DNA sequenc-

ing in individuals that we will obtain in the future. To

get answers to many questions regarding this species,

more observations, examinations, and study of different

specimens of this species are required. The questionable

records in previous reports (Vyas and Patel 2007; Patel et

al. 2015; Mirza et al. 2016) on the identification and lo-

calities are clarified here with enough justification, so the

records from Bhavnagar were Wallaceophis gujaratensis

but the records from Ahmedabad were Coronella brachy-

ura. Recently an individual of Wallaceophis gujaratensis

February 2019 | Volume 13 | Number 1 | e164

Indian Smooth Snake (Coronella brachyura)

INDIA

o_— Rivers

==== Tropic of CANCER

— State wise distribution of species

@ Tropic of Cancer passing through states

Y hn

Fig. 9. Distribution shown in map in all four states of India along with Tropic of Cancer, Narmada and Tapi rivers. Distribution and

localities shown separately in Gujarat state, Surat and Tapi districts. Map prepared by Nitin Patel and Smita Ramkumar.

Amphib. Reptile Conserv. 86 February 2019 | Volume 13 | Number 1 | e164

Parmar

Graph Showing Distribution Of

Coronella brachyura

. ==

Chhattisgarh Madhya Pradesh

Gujarat Maharashtra

a Districts mLocalities

Fig. 10. Graph showing distribution of Coronella brachyura. Prepared by Dikansh S. Parmar.

obtained from Saldi village, Amerli district (neighboring

district of Bhavnagar district) was mistaken as Coronel-

la brachyura. Later when the rescuer sent photographs

(Fig. 11), the author (DSP) identified it as Wallaceophis

gujaratensis. This is further evidence that specimens re-

ported from Bhavnagar were Wallaceophis gujaratensis

not Coronella brachyura. Four individuals of Wallaceo-

phis gujaratensis were also reported from Saldi village,

Amerli district by Mirza et al. (2016).

In India, Coronella brachyura is on schedule IV of the

Wildlife Protection Act of 1972 (Srinivasulu et al. 2013).

This species is rare but is listed under Least Concern

status according to the IUCN (International Union for

Conservation of Nature and Natural resources) Red List,

which justifies that this species is listed as Least Concern

in view of its wide distribution in the western and cen-

tral states of India, and the absence of recognized major

threats (Srinivasulu et al. 2013). However, the author has

observed certain threats to this species by humans, in-

cluding urbanization, expansion of roads, roadkill, habi-

tat loss, and intentional killing of snakes due to fear and

lack of awareness. There are also some natural threats

such as floods and predators. Although this species is re-

ported from four states of India, the populations are very

small. According to our findings this species should be

listed under Vulnerable status in order to provide ben-

efits for its protection. Due to rapid urbanization, forest,

agriculture lands, grasslands, etc. are being replaced by

concrete jungles. The natural habitats of animals includ-

ing reptiles are lost in most of the region. National gov-

ernment or international organizations can restrict un-

necessary construction by passing laws and limiting the

number of buildings in key habitat areas.

Acknowledgements.—The author is indebted and

grateful to Amphibian & Reptile Conservation for en-

Amphib. Reptile Conserv.

couraging submission by waiving all publication fees.

Mr. Bhavesh Trivedi (wildlife photographer and nature

activist) for providing photographs and information on

records of Wallace’s Striped Snake in Saurashtra prov-

ince of Gujarat state. Mr. Jenis Patel (Zoologist, Phd

scholar and researcher) for discussion and support. Mr.

Kaushal Modi for providing measurements of eggs and

female length. Mr. Mehul Thakur (volunteer in NGO

“Team Prayas”) for sharing valuable information about

the rescue records and additional localities of the spe-

cies in Surat district, and for his valuable comments. Mr.

Nitin Patel (involved in nature conservation programs)

for preparing the map, and Ms. Smita Ramkumar (pur-

suing M.Sc. in microbiology in Veer Narmad South

Gujarat University, Surat) for making corrections to the

map. Special thanks to Dr. Shantilal K. Tank (H. O. D.

of Department of Biosciences, Veer Narmad South Guja-

eas

Fig. 11. Wallaceophis gujaratensis with two distinct lateral

stripes, from Saldi village, Amerli district, Gujarat. Photo

credit Bhavesh Trivedi.

February 2019 | Volume 13 | Number 1 | e164

Indian Smooth Snake (Coronella brachyura)

gestions. Mr. Vasudev Limbachia (animal activist, snake

rescuer, and dog handler) for providing one additional

locality record of species in Surat. Mr. Vedant Lala (res-

cuer) for providing photographs of eggs.

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February 2019 | Volume 13 | Number 1 | e164

Parmar

583 p.

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Dikansh S. Parmar has been involved in rescue, rehabilitation, and research of birds,

animals, reptiles, and amphibians since childhood. He is interested in environmental

education and herpetology, and has conducted numerous snake awareness programs,

tree plantations, bird watching, and adventure activities including mountaineering,

artificial wall climbing, trekking, and camping. He worked on the environmental

education bus project proposed by C.E.E. (Center for Environment Education) in

schools of three districts of Gujarat: Surat, Bharuch, and The Dangs. He worked on

a bird census project, a vulture conservation project, and a project entitled “Survey of

river pollution due to industrial waste and its effect on aquatic life.” His research is

mainly focused on serpents. He is writing a book on snakes of Gujarat entitled The Cold

Blooded Gujarat which he is about to complete. He has contributed to two consecutive

editions of the Sarp Sandarbh book, 5" edition (2011) and 6" edition (2017). He also

contributed photographs, information, and as voluntary rescuer in ‘Snake Lovers Club’

app to educate people about the reptiles and ensure these are rescued successfully. He

is studying to receive a M.Sc. in Zoology in Veer Narmad South Gujarat University in

the department of Biosciences, in Gujarat, Surat and recently passed the written exam

February 2019 | Volume 13 | Number 1 | e164

Amphibian & Reptile Conservation

13(1) [General Section]: 90-103 (e170).

Official journal website:

amphibian-reptile-conservation.org

First record of Gracixalus quangi Rowley, Dau, Nguyen, Cao

& Nguyen, 2011, from Hoa Binh Province, Vietnam, including

the first documentation of advanced larval stages and an

extended tadpole description

12Cuong The Pham, *Anna Rauhaus, ‘Thang Dai Tran, *Christian Niggemann, ‘Phuong Huy Dang,

456Minh Duc Le, '?Truong Quang Nguyen, and *”*Thomas Ziegler

‘Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Hanoi, VIETNAM *Graduate

University of Science and Technology, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Hanoi, VIETNAM *AG Zoologischer

Garten K6ln, Riehler Strasse 173, D-50735 Cologne, GERMANY *Faculty of Environmental Sciences, Hanoi University of Science, Vietnam

National University, 334 Nguyen Trai Road, Hanoi, VIETNAM? Central Institute for Natural Resources and Environmental Studies, Hanoi National

University, 19 Le Thanh Tong, Hanoi, VIETNAM °Department of Herpetology, American Museum of Natural History, Central Park West at 79th

Street, New York, New York 10024, USA ‘Institute of Zoology, University of Cologne, Ziilpicher Strasse 47b, D-50674 Cologne, GERMANY

Abstract.—We provide the first record of Gracixalus quangi from Hoa Binh Province, Vietnam, based on

morphological and molecular evidence. The species was originally described from Nghe An Province and

subsequently recorded from Son La and Thanh Hoa provinces in Vietnam. The first documentation of advanced

larval stages up to Gosner stage 45 is provided from observations on the development of two egg clutches

at the indoor amphibian facility of the Me Linh Station for Biodiversity. The clutches were collected in Hang

Kia - Pa Co and Ngoc Son - Ngo Luong Nature Reserves in Hoa Binh. An extended description of a tadpole

in stage 35 is presented including the morphology of the oral disc and the first complete larval staging table

(comprising stages 14 to 45) for G. quangi. This is the first record of the species from a karst forest and we

provide the first natural history data of G. quangi from such a habitat type.

Keywords. Morphology, molecular biology, tadpole matching, new record, karst forest, Southeast Asia, development,

tree frog

Citation: Pham CT, Rauhaus A, Tran TD, Niggemann C, Dang PH, Le MD, Nguyen TQ, Ziegler T. 2019. First record of Gracixalus quangi Rowley,

Dau, Nguyen, Cao & Nguyen, 2011, from Hoa Binh Province, Vietnam, including the first documentation of advanced larval stages and an extended

tadpole description. Amphibian & Reptile Conservation 13(1) [General Section]: 90—103 (e170).

NoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any medium, provided

the original author and the official and authorized publication sources are recognized and properly credited. The official and authorized publication

credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation; official journal website <amphibian-

reptile-conservation.org>.

Received: 22 September 2017; Accepted: 15 March 2018; Published: 27 February 2019

Introduction lus quangi is listed as Vulnerable in the IUCN Red List

of Threatened Species, being known only from few lo-

The Quang’s Tree Frog, Gracixalus quangi, was recent-

ly described by Rowley et al. (2011) from the montane

evergreen forest of Pu Hoat Nature Reserve in western

Nghe An Province, Vietnam. The species has also been

recorded from Copia Nature Reserve in Son La Prov-

ince by Pham et al. (2012a) and from Xuan Lien Nature

Reserve in Thanh Hoa Province by Pham et al. (2012b).

The Quang’s Tree Frog inhabits dense and relatively un-

disturbed montane rainforest between 600 and 1,300 m

above sea level. Reproduction occurs from April to No-

vember when males aggregate at breeding sites. Eggs are

deposited on vegetation overhanging shallow, muddy

ponds, into which the larvae fall after hatching. Gracixa-

Correspondence. *ziegler@koelnerzoo.de

Amphib. Reptile Conserv.

calities in Vietnam. Its habitat continues to decline both

in quality and extent (IUCN SSC Amphibian Specialist

Group 2015).

Recent herpetological surveys conducted in Hang Kia

- Pa Co and Ngoc Son - Ngo Luong nature reserves al-

lowed a range gap in the distribution of G. qguangi to be

filled by providing the first record of the species from

Hoa Binh Province (Fig. 1), based on morphological and

molecular evidence, and recorded the species for the first

time from karst forest habitat.

In general, the importance of detailed morphological

larval descriptions have been underestimated, and

only limited information on morphology and natural

February 2019 | Volume 13 | Number 1 | e170

Pham et al.

Fig. 1. Adult Gracixalus quangi from Hoa Binh Province.

Photography by T: Ziegler.

history is available for anuran larvae, compared to adult

stages. However, the biphasic life cycle of anurans

imposes dramatically different selective regimes on the

aquatic larval stage, the metamorphic stage, and the

terrestrial post-metamorphic stage (Haas and Das 2011).

Knowledge about tadpole development is crucial for the

proper determination of anuran larval stages in the field,

e.g., for assessments of habitat, ecological adaptations,

and population density, which all provide important

information for potential conservation actions such as

conservation breeding programs in the context of the

global amphibian crisis.

Thus, detailed descriptions of developmental stages

from egg until metamorphosis are important additions to

larval descriptions, which are usually limited to one de-

velopmental stage that represents a very short period of

the anuran life cycle. Rowley et al. (2011) observed lar-

vae of G. quangi within clutches up to stage 24 of devel-

opment (Gosner 1960) and provided some measurements

and a photograph of a tadpole at stage 24. However, tooth

rows were not yet obvious in that early developmental

stage, so the oral disc was not described adequately.

Rowley et al. (2015) published new information on the

breeding habitats, eggs, embryos, and tadpoles of three

Gracixalus spp. and a description of the tadpole of G.

quangi up to stage 26. To provide a more complete de-

scription of larval development, including the advanced

larval stages, two egg clutches were transferred from

the newly recorded population in Hoa Binh Province to

the indoor amphibian facility of the Me Linh Station for

Biodiversity (Ziegler et al. 2016). Based on the larvae of

G. quangi from Hoa Binh which subsequently hatched

and further developed, we herein provide the first docu-

mentation of advanced larval stages together with an ex-

tended tadpole description, including the morphology of

the oral disc.

Material and Methods

Field surveys. Field surveys were conducted in Hoa Binh

Province in April 2014 (Fig. 2). Minimum and maximum

air temperatures and humidity were measured daily with

a thermo-hygrometer (TFA Dosmann/Wertheim Kat.

Amphib. Reptile Conserv.

Elevation (mi

0 75 1450

+++

Kilometers

Fig. 2. Map of Vietnam showing previous distribution records

(1: Pu Hoat Nature Reserve, Nghe An; 2: Copia Nature

Reserve, Son La; 3: Xuan Lien Nature Reserve, Thanh Hoa)

after Rowley et al. (2011), Nguyen et al. (2012), and Pham et al.

(2012a, b), as well as the new findings from Hoa Binh Province

(4: Hang Kia - Pa Co and 5: Ngoc Son - Ngo Luong nature

reserves).

Nr.30.5015). Of the twelve egg clutches observed near

small ponds in the limestone forest one egg clutch was

collected on 11 April 2014 from Hang Kia - Pa Co Nature

Reserve and one clutch was collected on 17 April 2014

from Ngoc Son - Ngo Luong Nature Reserve.

For proper species identification of larvae, four syntop-

ically occurring adult frogs (three males and one female)

were collected in Hang Kia - Pa Co and Ngoc Son - Ngo

Luong in April 2014, euthanized with ethylacetate, pre-

served in 70% ethanol, and deposited in the herpetologi-

cal collection of the Institute of Ecology and Biological

Resources (IEBR): One female IEBR 4329 (field number

HB 2014.25) and one male IEBR 4330 (field number HB

2014.26) collected by T.Q. Nguyen, C.T. Pham, and H.N.

Ngo on 12 April 2014 in Hang Kia - Pa Co Nature Reserve,

Hoa Binh Province, at an elevation of 1,376 m a.s.l.; two

adult males IEBR 4331, 4332 (field numbers HB 2014.73,

2014.74) collected by C.T. Pham, H.N. Ngo, and H.T. An

on 18 April 2014 in Ngoc Son - Ngo Luong Nature Re-

serve, Hoa Binh Province, at an elevation of 605 ma.s.1.

February 2019 | Volume 13 | Number 1 | e170

Gracixalus quangi distribution and larval description

Abbreviations: a.s.1. = above sea level; EN = distance

from anterior corner of eye to the nostril; HL = head

length; HW = head width; IND = internarial distance;

IOD = interorbital distance; NS = distance from nostril

to the tip of the snout; SVL = snout-vent length; UEW =

maximum width of upper eyelid.

Husbandry conditions and rearing. To further inves-

tigate the development of the eggs and larvae, the two

collected egg clutches were transferred to the indoor am-

phibian room at the Me Linh Station for Biodiversity.

The amphibian facilities at Me Linh were established

to keep and breed threatened or poorly known species

from Vietnam, both for research on husbandry, repro-

duction, and natural history and for building up ex situ

breeding programs and captive assurance populations

(see Ziegler et al. 2016). Room temperature was about

25-28 °C, and the humidity was 70-90%. Egg clutches

were left on leaves and kept in covered petri dishes be-

fore hatching. After hatching, tadpoles were housed in

plastic boxes measuring 36.5 x 26 x 13.5 cm (length x

width x height); the water depth was about 3 cm, and

air pumps provided for additional oxygen supply. Water

was changed manually every day. Feeding occurred 4—5

times a day with fish flakes (SERA vipan). Water param-

eters were: pH 6.6, KH (carbonate hardness) 9, GH (gen-

eral hardness) 4°, NO, 0 mg/l, NO, 0 mg/l.

Larval staging and description. Identification of larval

stages followed Gosner (1960) as reproduced in Mc-

Diarmid and Altig (1999). The two egg clutches were

photographed daily. Either daily or every few days after

hatching, up to five tadpoles from each clutch were pho-

tographed in a small aquarium and the total length was

measured (from the tip of the snout to end of the tail)

with a caliper. The day when the first clutch was found is

referred to as “day 1” 1n the Results section. One larva in

an early developmental stage (IEBR 4333a, field number

TZ 2014c) was preserved for genetic identification. Five

additional tadpoles (EBR 4333b, c, d, ZFMK 101032,

101033) in stages 35—38 were preserved for comparisons

of their oral discs with that of IEBR 4333a and for sub-

sequent morphological description. Preserved specimens

were deposited in the herpetological collections of the

Institute of Ecology and Biological Resources (IEBR)

of the Vietnamese Academy of Science and Technology,

Hanoi, Vietnam and the Zoologisches Forschungsmuse-

um Alexander Koenig, Bonn, Germany.

Measurements of preserved larvae were made with a

sliding caliper. Terminology for morphometric data and

abbreviations followed McDiarmid and Altig (1999), Al-

tig (2007), and Haas and Das (2011). The Labial Tooth

Row Formula (LTRF) with A (keratodont rows on upper

labium) and P (keratodont rows on lower labium) was ap-

plied as in McDiarmid and Altig (1999), and the ecomor-

phological assignment of larval types followed Altig and

Johnston (1989) and Orton (1953) as extended in Mc-

Amphib. Reptile Conserv.

Diarmid and Altig (1999). Abbreviations are as follows:

BH = body height; BL = body length (from snout to the

point where the axis of the tail myotomes meets the body

wall); BS = body end to center of spiracle; BW = maxi-

mum body width; ED = eye diameter; ES = eye-snout

distance; IND = internarial distance (center to center);

IOD = interorbital distance; LFH = lower fin height (at

MTH); MTH = maximum tail height; NE = distance from

center of naris to center of eye; ODW = oral disc width;

SN = distance of naris (center) to snout; SS = distance

of snout to center of spiracle; TAL = tail length (total

length minus body length); TMH = tail muscle height

at body-tail junction, where ventral line of musculature

meets trunk contour; TMW = tail muscle width at the

same level as TMW; TTL = total length (from the tip of

the snout to end of the tail); UFH = upper fin height.

Molecular analysis. Total genomic DNA was extracted

from a tissue sample of larva IEBR 4333a using a com-

mercially available DNeasy Tissue Kit following manu-

facturer’s instructions (QIAGEN Inc., Valencia, Califor-

nia, USA). A fragment of 16S gene was amplified with

the primer pair 16Sar + 16Sbr (Palumbi et. al. 1991).

The standard PCR conditions for 16S were 95 °C for 5

min, 40 cycles of [95 °C for 30 sec., 50 °C for 45 sec,

72 °C for 60 sec] and 72 °C for 6 min. All PCR products

were visualized on a gel before purification. Successful

amplification was purified to eliminate PCR components

with a GeneJETTM PCR Purification kit (Fermentas,

Canada). The purified PCR product was sent to Ist Base

(Malaysia) for sequencing. The sequence was compared

to those available for other species with a BLAST search

of GenBank.

Results

Morphology of adults. Morphological characters of the

four preserved adults from Hoa Binh Province are similar

to those in the original description of Gracixalus quangi

provided by Rowley et al. (2011): Males smaller than

females (SVL 22.8—24.0 mm male, 29.2 mm female);

head longer than wide (HL 8.7—10.2 mm, HW 8.3-9.2

mm male, HL 11.0 mm, HW 10.2 female); snout pointed

in dorsal view and in profile, projecting beyond margin

of the lower jaw; canthus rostralis distinct, loreal region

slightly concave; nostril oval, closer to tip of snout than

eye (NS 1.6—1.7 mm, EN 1.8—1.9 mm male, NS 1.8 mm,

EN 2.4 mm female); interorbital distance wider than 1n-

ternarial distance and upper eyelid (IOD 3.3-3.6 mm,

IND 2.5—2.8 mm, UEW 2.3—2.4 mm male, IOD 3.9 mm,

IND 3.4 mm, UEW 3.2 mm female); tympanum distinct,

rounded, about 40% eye diameter; vomerine teeth absent;

tongue notched posteriorly; external subgular vocal sac.

Forelimbs moderately robust; tips of fingers enlarged

into round discs with circummarginal grooves; relative

length of fingers I<II<IV<III; fingers free of webbing;

males with nuptial pad on finger I.

February 2019 | Volume 13 | Number 1 | e170

Pham et al.

Fig. 3. Adult couple of Gracixalus quangi from Hoa Binh

Province (male above, female below). Photo credit C.T; Pham.

Hindlimbs: tips of toes enlarged into round dics

with circummarginal grooves; relative length of toes

I<II<HI<V<IV; discs of toes slightly smaller than those

of fingers; webbing formula I1-11/211/2-21II1/2-21V2-

OV; inner metatarsal tubercle present; outer metatarsal

tubercle absent.

Dorsal surface of head, body, thigh, and shank with

small tubercles; largest and most concentrated on eyelids;

supratympanic fold present; throat and chest smooth;

ventral surface of thighs and belly coarsely granular.

Coloration in life (Fig. 3): Dorsal surface olive-green,

with brighter pale green on dorsal surface of upper arms;

brown interorbital bar covering eyelids and faint darker

X across back; black supratympanic line underneath su-

pratympanic fold, extending from eye to axilla; line of

large, black spots from supratympanic line to flanks; ven-

tral surface of throat, chest and belly opaque white with

translucent pale green margins.

Molecular analysis. According to the BLAST search of

GenBank, the 556 bps sequence of the amplified DNA

fragment was 99% similar to that of the Gracixalus quan-

gi holotype (GenBank accession number JN862537) and

thus the tadpole was clearly identified as conspecific.

Amphib. Reptile Conserv.

However, the new sequence differed from the type se-

quence in five positions.

Natural history. Adult frogs and 12 egg clutches were

found in the early rainy season in four breeding sites of

G. quangi in Hoa Binh Province, near two small ponds

(each ~1 m? in area) in the limestone forests of Hang

Kia - Pa Co and Ngoc Son - Ngo Luong nature reserves

(Figs. 4, 5). Clutches were small and consisted of eggs

in prominently clear jelly deposited in a clump at the tip

of green leaves between 0.3 to 0.7 m above the water

surface of the small ponds. In Hang Kia - Pa Co five egg

clutches containing 8 to 14 eggs each were found at an

elevation of 1,376 ma.s.l., and in Ngoc Son - Ngo Luong

seven egg clutches containing 6 to 18 eggs each were

found at an elevation of 650 m a.s.l. The mean clutch size

for the 12 clutches was 11 eggs (minimum 6, maximum

18). The habitat was located in the secondary karst forest

of medium and small hardwoods mixed with shrubs and

vines. The surrounding vegetation consisted of ferns and

other plants, less than one meter tall. Air temperature was

19.9-28.0 °C and relative humidity was 69-90%.

Developmental stages. The first clutch was found in

stage 14 (neural folds visible); stages 15-18 were ob-

served within the next two days (see Table 1 and Fig.

6 for early developmental stages). Developing bodies of

the larvae had a slightly darker coloration than the large,

white to cream-colored yolk mass. Stage 19 (gill buds,

distinct tail elongation) was photographed on day four;

stage 20 (gill circulation, further tail elongation) on day

five. The second clutch was found on that day in stage

20. At stage 22 (tail fins transparent), the pigmentation

of the larvae darkened slightly and the yolk sac started

to reduce. At stage 23, the body coloration had darkened

further, and the eyes had significantly darkened. The lar-

vae inside the eggs had grown continuously as the yolk

sac further reduced. At stage 25, the external gills had

atrophied and the yolk sac was largely absorbed. The

intestinal loop was visible on the lateral side of the tad-

poles inside the eggs. The grey coloration had extended

laterally as dark spots scattered on the sides and the ab-

Meee x

Fig. 4. Karst forest habitat in Hoa Binh Province. Photo credit

C.T! Pham.

February 2019 | Volume 13 | Number 1 | e170

Gracixalus quangi distribution and larval description

Table 1. Early developmental stages and morphological characters of Gracixalus quangi larvae from Hoa Binh Province; stage

diagnostic characters according to Gosner (1960) in italics. n: number of individuals observed at the respective stage; Age (days):

day the first clutch was found herein defined as day 1.

Stage Diagnostic characters n Age (days)

14 Neural folds 14 1

15-16 Elongation, rotation, neural tube, gill plates 13 2

17-18 — Tail bud, olfactory pits, large yolk mass, pigmentation 13 3

19 Gill buds, tail elongation, tail separated from yolk mass, tadpoles turned around inside eggs 4

20 Gill circulation, tail elongation 21 5

21 Cornea transparent, mouth opens 1] 6

pp) Tail fins transparent, pigmentation darkens, yolk mass starting to reduce 11 7

23-24 Labia and teeth differentiate, operculum covers gill bases / closes on right, coloration on body 11 8-9

and eyes darkened

2 External gills atrophied, mouth parts obvious, yolk mass largely resorbed, pigmentation 8 10

extended

domen in most larvae. On 4 May 2014 (24 days after

the first clutch was found), all larvae had hatched and

ingested food regularly; the tadpoles varied from stage

26 to stage 32 (see Table 2 and Figs. 7, 8 for advanced

developmental stages). The bodies of the tadpoles were

almost entirely translucent, with small dark grey and

white pigmented markings, that were concentrated dor-

sally and became more scattered and isolated towards the

sides. Although the dark grey pigments were arranged

more densely in some areas, they remained isolated dots,

while the white markings merged into conjunctions re-

sembling ice crystals in several individuals. In general,

the amount of grey and white pigments varied consider-

ably between the tadpoles, forming a distinct marble pat-

tern in some larvae, while in others only white or almost

no grey pigments were found throughout development.

The inner organs were clearly discernible and colored

white to rosy, the intestinal coils appeared white to light

grey. The dark grey and white spots were also found on

the tail, where especially the dark pigments were aggre-

gated and formed irregular large blotches in some larvae.

Those blotches were mostly found on the tail muscula-

ture and sometimes on the upper tail fin, the lower fin of

all tadpoles was completely translucent or only covered

by few white pigments. The v-shaped myotomes of the

tail musculature and blood vessels of the tail were well

discernible. The hind limb buds were white. The pupils

were black and round, and surrounded by a thin white to

light-yellow ring. The iris was light blue and covered by

skin with varying amounts of black and white pigments.

The keratinized structures of the oral disc were well dis-

cernible through the skin. The openings of the nares were

surrounded by dark pigmentation. In several individuals,

a triangular shaped white spot was found on the snout

tip. Around stage 28, the ground coloration of the skin

began to turn a pale green in some larvae. At stage 29

(length of hind limb buds > 1% d), in some individuals

the dark grey and white pigmentation had developed fur-

ther and formed a more distinct pattern, while in others

it remained as scattered markings. In most individuals,

Amphib. Reptile Conserv.

the white pigments had increased and extended laterally

and ventrally. The development of the hind limbs pro-

gressed from the foot paddle at stage 31 and stage 37,

all toes were separated and the legs began to bend at the

knee and foot joints. Dark grey pigmentation marked

the toes and limb joints. A larger part of the iris was ex-

posed and no longer covered by skin; the exposed part

around the pupil was pale yellow, while the outermost

ring was still blue with black pigmentation. At stage 39,

the foot was distinctly elongated; the keratinized mouth

pinrt

Fig. 5. Breeding sites of Gracixalus quangi in Hoa Binh

Province. Photo credit C.T: Pham.

February 2019 | Volume 13 | Number 1 | e170

Pham et al.

Table 2. Developmental stages and morphological characters of Gracixalus quangi larvae from Hoa Binh Province from stage 26

to 39; total length (TTL) in mm (mean + sd; range in parentheses), stage diagnostic characters according to Gosner (1960) in italics.

n: number of individuals observed at the respective stage. d: number of days in which stage could be observed within one clutch

(dates in parentheses); A: clutch found on 11 April 2014, B: clutch found on 17 April 2014; *= TTL of preserved larva in stage 36

excluded, as part of the tail was missing.

Stage Diagnostic characters

26-27 Larvae hatched, free-swimming and feeding,

blue iris coloration, light yellow ring around

pupil, body with dark grey and white pigments

in variable concentration, skin mostly

translucent, hind limb bud development (buds

light white)

28 Hind limb bud development (1 =d), pale green

dorsal ground coloration in some individuals

29 Hind limb bud development (I=11 d); dark

grey and white pigments extended in some

individuals

30 Hind limb bud development (1=2 d)

31 Foot paddle developed

32 Toe indentation 4-5

33 Toe indentation 3-4

34 Toe indentation 2-3

35 Toe indentation 1-2

ao" Toes 3-5 separated; forelimbs with elongated

fingers discernible under skin of preserved larva

3h All toes separated, legs begin to bend at knee

and foot joints, dark grey pigmentation on toes

and joints, larger part of iris exposed (pale

yellow); forelimbs with finger projections

discernible under skin of preserved larva

38 Meratarsal tubercle, toe elongation, toe joints

bending; forelimbs with elongated fingers and

well developed discs discernible under skin of

preserved larvae

39 Subarticular patches (slightly visible); foot

elongation

structures were still visible. Toe webbings and toe discs

were developed in stage 40 (see Table 3 and Fig. 8 for

stages of metamorphs). In stage 41, the coloration on the

back and hind limbs had become a pale green, while the

skin was still mostly translucent. The spiracle was still

present, the vent tube was gone. The eye coloration had

become light yellow. In several individuals, the dark 1n-

terorbital bar, which is also present in adults, appeared.

The fore limbs were well discernible under the skin, the

oral disc was reduced. Stage 45 (mouth posterior to eye,

tail stub) was only documented in one individual which

had entered the land section of the tank on 1 June 2014.

Fore and hind limbs were light green and speckled with

white dots; coloration on the body darker green to olive.

Amphib. Reptile Conserv. 95

TTL (mm) d (A) d(B)

17.5424 3 (May 4-6)

(14.6-21.4), n=6

nee: 5 (May 4-8)

(17.9-23.8), n=6

21.8406 5 (May 4-8)

(21.4-22.2), n=2

20.3 +2.0 7 (May 5-11)

(18.5-23.5), n=5

22.4420 5 (May 6-10)

(20.6-25.6), n=5

249429 2 (May 4-5)

(22.4-29.9), n=4

23.8424 2(May 4-5) 4 (May 7-10)

(20.9-26.9), n=6

25.6424 4. (May 4-7) 3 (May 9-11)

(20.8-29.5), n=9

26.6412 9 (May 4-12) 2 (May 9-10)

(25.7-28.6), n=7

ESE th 9 (May 6-14) 1 (May 11)

(26.1-29.4), n=6

Dect 7 (May 7-13) 2 (May 15-16)

(23.3-29,9), n=19

i pO We

(27.0-27.3), n=3

2 (May 17-18)

30.0+0.1

(30-30.1), n=3

2 (May 13-14) 1 (May 21)

Skin was still slightly translucent with inner organs vis-

ible. A light ochre to yellow, triangular shaped marking

was present on the forehead from the eyes to the snout

tip. The pupils were horizontal (round in larval stages);

the iris was colored golden-brown, with a bright yellow

ring around the pupil.

Assuming that the clutches were collected a few days

after egg deposition, the whole embryonal and larval de-

velopment required about 7—8 weeks.

Larval description. The following larval description is

based on a single tadpole (IEBR 4333b) in stage 35 (Figs.

9-11; for measurements see Table 4).

February 2019 | Volume 13 | Number 1 | e170

Gracixalus quangi distribution and larval description

Table 3. Developmental stages and morphological characters of Gracixalus quangi metamorphs from Hoa Binh Province; stage

diagnostic characters according to Gosner (1960) in italics. n: number of individuals observed at the respective stage. d: number of

days in which stage could be observed; dates in parentheses.

Stage Diagnostic characters d

40 Foot tubercles, vent tube present, toe webbings and toe discs developed Z 2 (May 19-20)

4] Forelimbs visible, mouthparts atrophy, vent tube gone, spiracle still present, ca. 20 14 (May 19—June 1)

back and hind limbs pale green, iris pale yellow, dark interorbital bar in

several larvae

45 Mouth posterior to eye, tail stub, froglet on land, fore and hind limbs light ] 1 (June 1)

green, speckled with white dots; body coloration darker green to olive, skin

slightly translucent; light ochre to yellow, triangular shaped marking on fore

head between eyes and snout tip; pupils horizontal; iris golden-brown, bright

yellow ring around pupil

Coloration in life: Skin translucent, dorsal body surface

pale green, laterally becoming whitish-transparent; ven-

trally completely translucent, inner organs (gills, heart,

gut coil) well discernible through the skin. Slight pig-

mentation consisting of small dark grey dots, density

decreasing from dorsal to ventral surface. Body dorsally

and laterally marbled with white, ice crystal shaped pig-

ments; on ventral side white marbling is only present in

intestinal coil area, while gill area is unpigmented. Spi-

racular tube is transparent. Soft mouthparts, hind limb,

and tail musculature are pale grey to white; tail muscu-

lature fades from anterior to posterior, becoming almost

transparent in the distal third of tail. V-shaped myosepts

of tail and large, red-colored blood vessels in the proxi-

mal third of tail are well discernible. Small aggregations

of dark dots forming diffuse blotches are present on the

proximal third of tail musculature. Tail fins are translu-

cent, with aggregations of the white pigments forming

irregular blotches on upper tail fin and in distal quarter

of lower tail fin. Ground coloration of iris is pale yellow,

with scattered light blue and black pigments; pupil is sur-

rounded by a unicolored, pale yellow ring. Skin which

partially covers eyes forms a darker ring around the eye,

which is irregularly patterned with bluish and dark grey

pigments.

Coloration in preservative: Ground coloration of body

and tail musculature yellowish-beige to light apricot, skin

distally translucent in dorsal view. Inner organs visible

through the skin appear reddish-brown. White pigments

on skin are not visible; dorsal and ventral pigmentation

consists of small grey spots; pigment aggregations on tail

musculature appear darker and almost black in preserved

specimen. Tail fins are transparent. Snout region, vent

tube, and hind limbs are whitish-translucent; few small

dark pigments scattered dorsally on hind limbs. Eyes and

keratinized mouth structures are black; pupils are white.

Description in dorsal view: Body shape oval, elongat-

ed (maximum body width 0.58 of body length), snout

rounded. Widest portion of body at the gill region (pos-

terior to eyes) in dorsal view; with a slight constriction

of body contour near location of spiracle opening. Oral

Amphib. Reptile Conserv.

disc is positioned anteroventrally and not visible in dor-

sal view. Eyes are of moderate size (eye diameter 0.23

of body width), directed laterally and positioned dorso-

laterally at the first body third (interorbital distance 0.58

of maximum body width), not visible in ventral view.

Nares are small, anterodorsally positioned and directed:

located nearer to snout tip than to pupil (SN 0.29 of NE);

internarial distance is 0.32 of interorbital distance. Tail

musculature is moderately developed (width of tail mus-

culature at base 0.42 of maximum body width).

Description in lateral view: Body slightly depressed

(maximum body height 0.81 of maximum body width),

Fig. 6. Early developmental stages of Gracixalus quangi from

Hoa Binh Province (development stages indicated). Photo

credit C.T. Pham.

February 2019 | Volume 13 | Number 1 | e170

Pham et al.

Table 4. Measurements of preserved tadpoles of Gracixalus quangi from Hoa Binh Province in mm; (for abbreviations see Material

and Methods) *= part of tail missing; ** = specimen used for genetic analysis with tail and part of body missing.

Specimen TEBR 4333a ~~ TEBR 4333b.——s TEBR 4333c~=— ZFMK 101032. —- IEBR 4333d_ Ss ZFMK 101033

Stage ? 35 36 37 38 38

BH 2.6 4.2 44 3.8 4.0 5.0

BL : 92 8.5 8.6 9.9 97

BS : 3.5 2.4 39 3.4 3.2

BW 43 53 5.9 4.7 57 5.9

ED 0.8 1.2 1.8 1.3 1.6 1.6

ES 1.5 1.7 1.9 1.8 1.9 1.9

IND {2 1.0 1.3 1.2 1.4 Ll

IOD 1.9 31 3.8 3.2 37 3.4

LFH : Ld : 0.6 0.5 0.8

MTH . 41 : 3.6 37 3.6

NE 12 1.4 1.8 1.4 1.5 2.0

ODW 1.6 aS) 2.4 2.1 2.1 2.1

SN 03 0.4 0.5 0.5 0.7 0.5

SS 43 5.7 6.1 5.4 6.5 6.5

TAL - 16.5 : 17.6 17.4 17.3

TMH J 2.8 3.0 2.9 3.0 2.5

TMW : i) 2.9 2.5 2.0 2.2

TTL 6.9 25.7 19.9 26.2 273 27.0

UFH : 1.4 : 1.2 Ll 11

LTRF 4(2-4)/3 5(2-5)/3 5(2-5)/3 5(2-5)/3(1) 5(2-5)/3(1) 5(2-5)/3(1)

with maximum height approximately on spiracle axis,

snout rounded. Tubular spiracle sinistral, laterally posi-

tioned at second third of body (distance from snout tip

to opening of spiracle 0.61 of body length), opening pos-

terolaterally and visible in dorsal view; spiracular open-

ing is oval. Outer wall of the medial vent tube is attached

to lower tail fin; cloacal aperture is dextral. Tail relatively

long (tail length 0.64 of total length) and of moderate

height (maximum height of tail, including fins, is 0.25

of tail length), v-shaped mysepta of tail musculature are

well discernible (tail muscle height 0.67 of maximum

body height, and 0.68 of maximum tail height). Tail mus-

culature almost parallel at proximal half, then gradually

tapering to end of tail, not reaching tail tip. Upper fin

originates at body-tail junction, lower fin inserts axially

parallel, connecting with abdomen. Point of maximum

tail height is located at the second half of the tail; dorsal

fin is higher than ventral fin (maximum height of upper

fin 0.34 of maximum tail height; maximum height of

lower fin 0.79 of maximum height of upper tail fin), tail

tip is narrowly rounded.

Oral disc anteroventrally positioned, of nearly trap-

ezoidal shape and laterally emarginated (oral disc width

0.42 of maximum body width). Oral disc framed by

finger-shaped marginal papillae, upper labium with large

medial gap. Lower labium and lower half of upper la-

bium with one row of submarginal papillae (about 50

submarginal papillae under posterior keratodont row P3).

Amphib. Reptile Conserv.

Labial tooth row formula 5(2-5)/3; AS greatly reduced.

About 43 keratodonts per 0.5 mm. Jaw sheaths black,

slightly serrated. Upper jaw sheath forming smooth arc,

lower jaw sheath V-shaped.

Measurements (in mm): BH 4.2; BL 9.2: BS 3.5; BW

5.3; ED 1.2; ES 1.7; IND 1.0; IOD 3.1; LFH 1.1; MTH

4.1; NE 1.4; ODW 2.2; SN 0.4; SS 5.7; TAL 16.5; TMH

2.8; TMW 2.2; TTL 25.7; UFH 1.4.

Variation within the larvae series. Oral disc: Generally

uniform in all preserved specimens; except for a medial

gap in P1 in stage 37 and 38 tadpoles (LTRF 5(2-5)/3[1 ]).

AS is very short in all specimens examined and not yet

present in the specimen used for genetic comparisons

(IEBR 4333a), which was preserved in an earlier lar-

val stage. The shape of the tail fins, especially the tail

tip, varies from broadly to narrowly rounded / bluntly

pointed. The coloration is highly variable; especially the

amount and composition of blotches formed by the dark

grey and white pigments. The nares are surrounded by

dark grey ridges in some larvae, and sometimes a distinct

white, nearly triangular fleck is present reaching from the

nares to the snout tip. The greenish dorsal coloration is

more intense in the advanced stages.

Proportions vary as follows (ratios for the tadpole for

genetic comparison are excluded here, as this specimen

was preserved in an early larval stage, and the tail and

part of the body were dissected for molecular analy-

February 2019 | Volume 13 | Number 1 | e170

Gracixalus quangi distribution and larval description

Fig. 7. Advanced developmental stages of Gracixalus quangi from Hoa Binh Province (development stages indicated). Photo credit

T. Ziegler.

a. ie

Fig. 8. Advanced developmental stages of Gracixalus quangi from Hoa Binh Province (development stages indicated). Photo credit

T. Ziegler and T:D. Tran.

Amphib. Reptile Conserv. 98 February 2019 | Volume 13 | Number 1 | e170

Pham et al.

vine eh

pet

Fig. 9. Drawing of a larva of Gracixalus quangi from Hoa Binh Province (IEBR 4333b) in development stage 35 in dorsal and

lateral view. Drawing by C. Niggemann.

ses): BW 0.54—0.69 of BL; ED 0.27-0.3 of BW; IOD

0.58-0.68 of BW; SN 0.25—0.47 of NE; IND 0.32-0.38

of IOD (0.63); TMW 0.35-0.53 of BW; BH 0.70-0.85

of BW; TAL 0.64—0.66 of TTL, TMH 0.63-0.76 of BH;

TMH 0.69-0.81 of MTH:; UFH 0.30-0.33 of MTH; LFH

0.46-0.73 of UFH; ODW 0.36-0.45 of BW.

In the four preserved stage 36-38 tadpoles, the fore-

limbs in different stages of development were well dis-

cernible under the skin in ventral view (see Table 3).

Tadpoles of Gracixalus quangi are exotroph: lentic:

benthic larvae of Orton’s Type IV.

Discussion

In this study, we confirmed the findings of Rowley et al.

(2015) that Gracixalus quangi deposits its eggs in clear

jelly clumps on the surface of live leaves, near the leaf

tip, in moderate heights above non flowing water. Clutch

sizes recorded herein accorded well with observations by

Rowley et al. (2015). The maximum height of clutches

above the water surface was <50 cm according to Row-

ley et al. (2015), while we found them at heights of up

to 70 cm. Rowley et al. (2011, 2015) examined tadpoles

up to stages 24 and 26, respectively, which generally cor-

responded well with our findings (e.g., large yolk mass in

embryonic stages). Rowley et al. (2011, 2015) assumed

the actual tooth row formula could only be observed in

advanced stages, which we herein confirm.

Grosjean (2005) recommended assessing tadpoles of

stages 32-40 for larval descriptions possessing the com-

plete set of taxonomically relevant characters in this range

of development. The keratodont row formula 5(2-5)/3[1]

was constant in our specimens in stages 35-38; while

the individual used for genetic comparison, which was

preserved 1n an earlier stage, was lacking the fifth tooth

Amphib. Reptile Conserv.

row in the upper labium. However, as this tooth row was

greatly reduced in all analyzed specimens and this indi-

vidual was obtained from the same egg clutch and corre-

sponded well morphologically with the other specimens

— —

Qe =

= Es]

a =

all —_

rat =

Zz =

a i

ot lage

te te

ee ¢ uae Cary pee casey ‘ a

a we ae

a (ey aN gua ny m At fe 2

ae HEMET ELL WAU Mathias ty fi sycagegnra mua eciegacte "” ==,

— PUI a

— “Aun, arene =

— CETTE ries =

WLU Lies Nea vee =

aie AOA aL nada yiiensiees sete run a

F we

fi ie

i td dA vere Wee ecas

OA Lg ee Lab?

Meda ia AAU LUG MMO

Fig. 10. Drawing of the oral field of a larva of Gracixalus

quangi from Hoa Binh Province (IEBR 4333b) in development

stage 35: natural view on top, schematic drawing below).

Drawing by C. Niggemann.

February 2019 | Volume 13 | Number 1 | e170

Gracixalus quangi distribution and larval description

pry

~_ * ie : - ’

" ae tol “ 7

; = Ce dl =) 7) Py vs a, * 2

fw 4a we? : : |) >.

Fig. 11. Larva of Gracixalus quangi from Hoa Binh Province (I

2014). Photo credit T. Ziegler.

photographed in comparable stages, it does not belong to

a different species. The gap in P1, which was present in

the specimens in stages 37 and 38, is likely to represent

intraspecific variation rather than being caused by onto-

genetic atrophy of the oral disc, which usually occurs in

later stages; however, the mechanism of mouth part atro-

phy is poorly studied and it is not known whether defined

patterns of tooth loss within the tooth rows occurs during

ontogeny (McDiarmid and Altig 1999).

While Gosner (1960) defined tadpoles in stages 20—25

as “hatchlings,” the larvae of G. guangi remained longer

inside the eggs. A lack of data acquisition in late April

prevented photographing larvae directly after hatching,

but since tadpoles of the first clutch remained inside the

eggs in stage 25 on day 10 and the first free swimming

larvae were documented in stages 26—27 on day 24, we

assume that hatching occurred in stage 25 and the maxi-

mum period of this stage was 13 days. A delayed hatch

may be advantageous for avoiding predation inside the

water body in early hatchling stages, as tadpoles in these

stages are quite active and able to swim when they drop

into the water. This could also explain the relatively large

yolk mass during the embryonic stages, which was com-

pletely resorbed at the time of hatching. However, the

time of hatching seems to be somewhat variable, as Row-

ley et al. (2011) found hatched larvae in stage 24.

The transfer of the eggs and the different climatic con-

Amphib. Reptile Conserv.

eer ae :

> | ieee

fy a. 4

EBR 4333b) in development stage 35 in life (photo taken 4 May

ditions in the Me Linh Station (i.e., relatively constant

temperatures of 25—28 °C versus fluctuations between

19-28 °C in the field) could have led to a delayed hatch

in our study. Rowley et al. (2015) also discussed the pos-

sible influence of the substrate on which eggs were de-

posited (live vs. dead leaves) on the embryonic develop-

ment. In general, the number of days the tadpoles were

found in a certain stage varied between 0.5—1 day for the

embryonic stages and between 2—14 days for the larval

and metamorph stages, but due to the small sample sizes

in some stages, these values may not be representative

of the general development. The hatchling in stage 26

described by Rowley et al. (2015) was relatively small

compared to our observations (10.6 mm TTL vs. 15.8

mm as the smallest length measured in stage 26 here).

The hatchling size could be influenced by the time of

hatching and various environmental parameters such as

water temperature, larval rearing density, and food avail-

ability, which were possible influences both on the tim-

ing of larval development and growth in other anuran

tadpoles (e.g., McDiarmid and Altig 1999; Alvarez and

Nicieza 2002). We found larvae from each clutch with

a maximum developmental distinction of five stages per

day and size differences up to 8.3 mm per stage, indi-

cating that variation in developmental time and growth

within a clutch reared under the same conditions occurs

at a certain level. The development of the forelimbs is

February 2019 | Volume 13 | Number 1 | e170

Pham et al.

clearly visible in several of the photographed and pre-

served tadpoles from stage 36 onwards due to the trans-

parency of the ventral skin. There is no defined sequence

for forelimb development in Gosner’s staging system

compared to hind limb development, but according to

McDiarmid and Altig (1999) forelimb development lags

behind that of the hind limbs by about one stage. How-

ever, the fingers were noticeably further elongated and

shaped in the larva preserved in stage 36 than in the one

in stage 37 (see Table 2), suggesting there is also some

variation in development before forelimbs emerge.

The white spot on the snout tip, which was listed as a

diagnostic character for tadpoles of the genus Gracixalus

by Delorme et al. (2005), included only the two species

G. supercornutus and G. gracilipes. However, it was not

detected in the study of Rowley et al. (2011), who sug-

gested it could be either a variable feature among species

within the genus, or it is present only in more well-devel-

oped tadpoles. As we found tadpoles with and without

the white spot on the snout, it seems to be a variable fea-

ture and not related to ontogenetic change. In general, we

found the coloration during the larval stages to be quite

variable (see Figs. 7, 8).

Finally, our first record of G. guangi from a karst for-

est environment indicates a broader adaptation poten-

tial of the species to different habitats than previously

thought, and thus also implies the possibility of a wider

distribution range.

Acknowledgements.—We are grateful to the direc-

torates of the Forest Protection Department of Hoa Binh

Province, Ngoc Son - Ngo Luong, and Hang Kia - Pa Co

nature reserves for support of our field work and for is-

suing relevant permits. We thank H.T. An and H.N. Ngo

(Hanoi) for their assistance in the field, and A. Hoch (Co-

logne) for a preliminary assessment of the larval mor-

phology. M. van Schingen and U. Schepp (Bonn), and A.

Vassilieva (Moscow) kindly helped to improve an earlier

version of the manuscript. We thank E. Sterling (New

York) and K. Koy (Berkeley) for providing the map. This

research was supported by the National Foundation for

Science and Technology Development (NAFOSTED,

Grant No. 106.05-2017.329) and the Cologne Zoo (Ger-

many).

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Cuong The Pham is a researcher at the Institute of Ecology and Biological Resources (IEBR) - Vietnam

Academy of Science and Technology (VAST). He recently obtained a doctoral degree at the Graduate

University of Science and Technology, Hanoi. He is a member of the Cologne Zoo’s Biodiversity and

Nature conservation projects in Vietnam and has published several papers, mainly dealing with Vietnams’

herpetodiversity. Cuong is experienced in biodiversity and field research and has conducted numerous field

surveys in Vietnam.

Anna Rauhaus started her career at the Aquarium/Terrarium Department of the Cologne Zoo in May

2011 and has been head keeper of the Terrarium section since 2014. Her focus of expertise is keeping

and breeding of amphibians, monitor lizards, snakes, and crocodilians as well as behavioral training. She

provides keeper trainings and helps to develop amphibian and reptile facilities within the frame of Cologne

Z0o’s conservation projects in Vietnam. Since 2011 she has been involved in 44 herpetological publications

with a focus on zoo biology, among which 14 have dealt with captive breeding, larval development and

diversity of tropical, in particular, Vietnamese amphibians.

Thang Dai Tran is a researcher at the Me Linh Station for Biodiversity, IEBR. His research interests are

herpetology and biodiversity conservation and has published several papers concerning the herpetofauna of

the Me Linh Station and reproductive biology of amphibians.

Christian Niggemann joined the Aquarium/Terrarium Department of the Cologne Zoo in 2014. He is a zoo

keeper in the Terrarium section with long term experience in amphibian and reptile husbandry. Christian

has artistic skills, and has created the comic strip “Shini” (representing the threatened Vietnamese crocodile

lizard) which guides visitors and children through the environmental exhibition of IEBR’s Me Linh Station

for Biodiversity in Vietnam, in addition to comic strips for the fire salamander and the green toad.

Phuong Huy Dang is Head of the Me Linh Station for Biodiversity, IEBR. His research interests are

mammalogy and biodiversity conservation. Phuong has conducted many field surveys on mammals in

Vietnam and is co-author of four books and 48 papers.

Minh Duc Le has worked on conservation-related issues in Southeast Asia for more than 15 years. His

work focuses on biotic surveys, wildlife trade, and conservation genetics of various wildlife groups in

Indochina. He is currently characterizing the genetic diversity of highly threatened reptiles and mammals in

the region and has pioneered the application of molecular tools in surveying critically endangered species in

Vietnam. Minh has long been involved in studying the impact of wildlife trade on biodiversity conservation

in Vietnam, and is developing a multidisciplinary framework to address the issue in that country.

Truong Quang Nguyen is a researcher at the Institute of Ecology and Biological Resources (IEBR),

Vietnam Academy of Science and Technology (VAST) and is key member of the Biodiversity and Nature

Conservation projects of the Cologne Zoo. He finished his Ph.D. in 2011 at the Zoological Research

Museum Alexander Koenig (ZFMK) and the University of Bonn, Germany (DAAD Fellow). From 2011 to

2014 he was a postdoc in the Institute of Zoology at the University of Cologne. He has conducted numerous

field surveys and is the co-author of seven books and more than 150 papers relevant to biodiversity and

conservation in Southeast Asia. His research interests include the systematics, ecology, and phylogeny of

reptiles and amphibians from Southeast Asia.

Amphib. Reptile Conserv. 102 February 2019 | Volume 13 | Number 1 | e170

Amphib. Reptile Conserv.

Pham et al.

Thomas Ziegler has been the Curator of the Aquarium/Terrartum Department at the Cologne Zoo

since 2003 and is the coordinator of the Cologne Zoo’s Biodiversity and Nature Conservation Projects

in Vietnam and Laos. Thomas studied biology at the University Bonn (Germany), and conducted

his diploma and doctoral thesis at the Zoological Research Museum Alexander Koenig in Bonn,

with a focus on zoological systematics and amphibian and reptile diversity. He has been engaged

with herpetodiversity research and conservation in Vietnam since 1997. As a zoo curator and project

coordinator he tries to combine in situ and ex situ approaches, viz., to link zoo biological aspects with

diversity research and conservation, in the Cologne Zoo, in rescue stations and breeding facilities in

Vietnam, and in Indochina’s last remaining forests. He is a Professor at the Institute of Zoology at

Cologne University. Since 1994, Thomas has published 422 papers and books, mainly dealing with

herpetodiversity.

103 February 2019 | Volume 13 | Number 1 | e170

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

13(1) [General Section]: 104-110 (e171).

Genetic diversity of Egyptian populations of the African

Common Toad (Sclerophrys regularis, Reuss 1833)

Lamiaa Elsayed Mokhtar Deef

Department of Zoology, Faculty of Science, Damietta University, New Damietta, Damietta, EGYPT

Abstract.—Genomic data have become invaluable for answering questions in biological conservation and for

gaining high resolution in population genetic studies. A molecular dataset has been integrated to provide

genetic variation and baseline genetic information, using mitochondrial cytochrome oxidase subunit | (COl)

gene analysis of the African Common Toad (Sclerophrys regularis), order Anura, family Bufonidae. In the

present study, mitochondrial DNA sequence data were analyzed for Sclerophrys regularis from many localities

in Egypt. Based on COI sequences, the phylogenetic tree was constructed using the Maximum Likelihood (ML)

method. Results show that Egyptian Sclerophrys regularis populations have very high genetic diversity and

gene flow among them. The haplotype diversity was 1.000 for all studied regions, except for Gharbiya and Beni

Suef populations which were0.900 and 0.833, respectively. The low haplotype diversity values in these two

regions could indicate a possible genetic barrier between the South and North River Nile that is restricting gene

flow, such as water sources, climatic conditions or distances between habitats. At present, there is insufficient

data to determine the evolutionary significant units (ESU) for the conservation of Sclerophrys regularis. More

exhaustive studies should examine the more variable genetic markers and the ecology of this species to

establish a conservation strategy.

Keywords. Amphibian, Anura, Bufonidae, COI gene, DNA sequence, haplotype

Citation: Deef LEM. 2019. Genetic diversity of Egyptian populations of the African Common Toad (Sclerophrys regularis, Reuss 1833). Amphibian &

Reptile Conservation 13(1) [General Section]: 104-110 (e171).

ternational (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in any medium,

provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced, are as follows:

Official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org.

Received: 13 June 2018; Accepted: 27 June 2018; Published: 10 March 2019

Introduction

Amphibians play a pivotal role in properly functioning

ecosystems, sharing in nutrient cycling, bioturbation,

energy flow, food webs, and other ecosystem dynamics

(Hocking and Babbitt 2014; Cortés-Gomez et al. 2015).

Indeed, these animals provide additional ecosystem ser-

vices valuable to humans, such as regulating pests, serv-

ing as a food source, functioning as models for medical

research, and giving enjoyment and intangible contribu-

tions that vary across cultures (Warkentin et al. 2009).

Because of their critical importance in functional eco-

systems, anthropogenic efforts toward the maintenance

of amphibian diversity are essential. As a prerequisite,

amphibian diversity first needs to be measured accurately

so that improvement scan be documented and restoration

measures implemented in disturbed systems.

Stuart et al. (2004) and the International Union for

Conservation of Nature (IUCN) (2015) reported that am-

phibians are the most threatened group of vertebrates as-

sessed to date. In spite of this fact, amphibian species are

among the most poorly known vertebrate groups in many

geographic areas (Pino-Del-Carpio et al. 2014).

Correspondence. /amiaadeef@yahoo.com, lamiaadeef@du.edu.eg

Amphib. Reptile Conserv.

Recently, declines of native amphibian populations

have received a great deal of attention. Reports indicate

that diseases, pollution, habitat destruction, predation,

and competition with exotic species may be related to

amphibian declines (Houlahan et al. 2000; Kiesecker et

al. 2001; Stuart et al. 2004; Whiles et al. 2013). One of

the most important families of the class Amphibia is the

Bufonidae, which is distributed in all parts of the world

except for Antarctica. Due to their wide distribution,

bufonids are frequently used as model organisms in ex-

perimental biology studies. The genus Sclerophrys (pre-

viously part of the genus Bufo) is a widespread and well-

known bufonid genus consisting of 17 species (Saad et al.

2009), including the African Common Toad, Sc/erophrys

regularis Reuss, 1833 (Borkin 1999: 338; Ibrahim 2001;

Baha El Din 2006). Recently, the name Amietophrynus

regularis (Reuss, 1833) was applied (Borkin and Litvin-

chuk 2013; Ibrahim 2013a, b), however, the generic name

should be replaced by the senior valid name Sclerophrys

Tschudi, 1838 (Ohler and Dubois 2016) providing the

combination Sclerophrys regularis (Reuss, 1833). The

African Common Toad ts a large, strong toad with warty

skin. The dorsal surface is dark olive-brown in color with

March 2019 | Volume 13 | Number 1 | e171

Deef

dark, often symmetrically arranged patches on the back.

Smaller dark blotches occur on the upper lip and the eye-

lids and dark markings separate the warts on the flanks.

The undersides of both sexes are white to beige and the

throats of males are black (Rodel 2000).

The African Common Toad is often found near rivers,

where it also breeds. Furthermore, it is distributed across

a wide geographic range that makes this species an ideal

candidate for a geographic analysis of its genetic vari-

ability. This toad is abundant, found in both moist and

dry savanna, forest margins, montane grassland, and ag-

ricultural habitats. It is distributed widely in Sub-Saharan

Africa, with its range extending to the oases in Algeria

and Libya and into northern Nilotic Egypt (Frost 2007).

In Egypt, this toad is adaptable, but the molecular

characterization of this species remains unclear (Sakr et

al. 2014). Presently, pollution, habitat destruction, preda-

tion, and competition with exotic species may be contrib-

uting to the decline of this species, drawing attention to

the study of its genetic variation. Therefore, the purpose

of this study was to elucidate the genetic variability of

Sclerophrys regularis populations in Egypt and to pro-

vide baseline genetic information, using mitochondrial

cytochrome oxidase subunit I (COI) gene analysis.

Material and Methods

Study area and sample collection

Egypt covers an area of about one million km? (Fig. 1) in

the central part of the great Palearctic desert belt which

extends from the Atlas Mountains in the west to the Gobi

Desert in the east. Most of Egypt is occupied by some

of the driest deserts of the world, only interrupted by

the Nile Valley and Delta, and a few small oases. Saleh

(1997) divided Egypt into four habitats for amphibians:

The Western Desert, the Eastern Desert, the Sinai Penin-

sula, and the Nile Valley and Delta.

In the present study, fourteen locations were selected as

the study areas, representing each of the different habitats

of Egypt that are suitable for this toad:1. North Coast of

Egypt, Matrouh, Alexandria, Damietta, and Ismailia (Ma-

rine and Coastal Habitat, toads were captured from parks

and around buildings); 2. Arish and Sharm El-Shaikh

(Marine and Coastal Habitat of the Sinai Peninsula, toads

were captured from agricultural lands); 3. Gharbrya, Cai-

ro, Bani Sweif, Menia, Sohag, Qena, and Aswan (the Nile

Valley, Delta and the Eastern Desert, toads were captured

from slow-flowing pools along streams and swamps, and

around building and roads); 4. Sitwa Oasis (the Western

Desert, toads were captured from farms).

Sixty-nine African Common Toads were obtained from

the fourteen locations in Egypt, with between four and

seven toads obtained per locality (Fig. 1). These locations

were numbered from 1 to 14, with the number of toads

from each shown in parentheses: 1. Sharm El-Shaikh (4);

2. Arish (4); 3. Ismailia (5); 4. Damietta (6); 5. Alexandria

Amphib. Reptile Conserv.

Fig. 1. Map of the sampled localities.

(5); 6. Matrouh (4); 7. Gharbrya (5); 8. Cairo (4); 9. Stwa

Oasis (7); 10. Beni Suef (4); 11. Menia (5); 12. Sohag (5);

13. Qena (6), and 14. Aswan (5).Toads were sampled from

April 2015 to March 2016. The specimens were caught in

special nets or picked up by volunteers during nocturnal

and diurnal surveys.

DNA extraction, PCR amplification, and mito-

chondrial DNA sequencing

Samples of muscle tissues from the toads were taken 1m-

mediately and frozen at -80 °C. DNA was extracted using

a GeneJET™ kit Genomic DNA Kit#K0721. COI gene

was amplified using primers FE] (5'- GGT CAA CAA

ATC ATA AAG ATA TTG G -3’) and RE] [(5'- TAA ACT

TCA GGG TGA CCA AAG AAT CA -3')]. The poly-

merase chain reactions (PCR) consisting of ~50 ng of

template DNA were carried out in volumes of 15 wl with

1x PCR Buffer: 2 mM MgCl, 0.5 uM each of FEI] and

RE1, 0.2 mM dNTP, and 0.6 U Taq DNA Polymerase.

The thermocycling conditions to amplify DNA using the

polymerase chain reaction (PCR) were: 1 cycle 96 °C/3

min; 35 cycles 95 °C/30s, 55 °C/45 s, 72 °C/1.5 min; and

lucycles/22C/ min:

The PCR products were separated on 1.0% agarose

gels, bands were visualized by ethidium bromide staining

and viewed with an ultraviolet light source. The amplified

products were purified using a GeneJET™ kit (Thermo

K0701) according to the manufacturer’s instructions. Se-

quencing was performed using an ABI 3730xl DNA se-

quencer.

Data Analysis

For sequence alignment, MEGA version 7.0 (Kumar et

al. 2016) and Geneious version 5.3 (Drummond et al.

2010) were used, followed by visual editing. Numbers

of haplotypes, haplotype diversity (h) and nucleotide di-

March 2019 | Volume 13 | Number 1 | e171

Genetic diversity of Egyptian populations of Sclerophrys regularis

Baie je}SOO pue sulle

7 Nile Delta

s Cairo

9° Western Desert

| 4

a 1

7 @

m Zz

orm

12 a

13 Uo ©

14 30

Fig. 2. Phylogenetic tree of African Common Toad, using COI haplotypes based on the Maximum Likelihood method. Numbers

refer to localities mentioned in the text: 1. Sharm El-Shaikh; 2. Arish; 3. Ismailia; 4. Damietta; 5. Alexandria; 6. Matrouh; 7.

Gharbtya; 8. Cairo; 9. SiwaOasis; 10. Bani Sweif; 11. Menia; 12. Sohag; 13. Qena; 14. Aswan.

versity (7) were calculated from aligned DNA sequences

by DnaSP version 6.0 (Rozas et al. 2017). Within and

among population genetic diversities were estimated by

calculating Nei’s nucleotide diversity (Pi) indices us-

ing DnaSP, version 6.0. Within-population gene diver-

sity (H), gene diversity in total populations [H, = H,+

D,, D,, gene diversity between populations and Hs,

the average intrapopulation diversity (Nei 1973)], and a

measure of population differentiation (G,,), were calcu-

lated according to the methods described by Pons (1996).

MEGA version 7.0 was used to construct a Maximum

Likelihood tree.

Results

Partial COI fragments of 654 bp were obtained from the

69 individual Egyptian toads from the 14 populations

and some of the sequences were deposited in GenBank

(Accession numbers KF665552, KF665569, KF665599,

KF665651, | KF665716, | KF665756, KF665824,

KY079472, KY079473, KY079476, KY079477,

KY079478, KY079479, KY079480, KY079483,

KY079484, KY079487, KY079488).

Haplotype information is shown in Table 1. For COI

data of Sclerophrys regularis, 67 haplotypes were recov-

Amphib. Reptile Conserv.

ered. Of the 14 populations, the Siwa Oasis population

had seven haplotypes; Damietta and Qena had six; Ma-

trouh, Arish, Sharm El-Shaikh, Cairo, and Gharbitya each

had four; and Beni Suef three.

The average values of haplotype diversity (h) and

nucleotide diversity (7) of the toads are shown in Table

2. The haplotype diversity was 1.000 for each of the

studied regions except for the Gharbiya and Beni Suef

populations, which were 0.900 and 0.833, respectively,

indicating lower haplotype diversity values in these two

localities. However, nucleotide diversity was the high-

est in Cairo at 0.71177. Damietta and Beni Suef had the

lowest nucleotide diversities at 0.27339 and 0.23012, re-

spectively.

Total genetic diversity (H, = 0.999) was higher than

the average intrapopulation diversity (H, = 0.00001) re-

sulting in high levels of genetic differentiation (G,, =

0.99899) as shown in Table 3. These results indicate that

this toad has high levels of genetic variation among its

Egyptian populations and distinct population structures

at all but two of the locations, Beni Suef and Gharbiya.

Based on COI sequences, a phylogenetic tree was con-

structed using the Maximum Likelihood method (Fig. 2).

The tree shows two distinct clades: the first clade com-

prises the Sharm El-Sheikh and Arish populations and the

March 2019 | Volume 13 | Number 1 | e171

Deef

Table 1. Haplotype numbers of the study localities of African

Common Toad (Sclerophrys regularis).

Population

Sharm E]-Shaikh

Arish

Ismailia

Haplotype numbers

Damietta

Alexandria

Matrouh

Gharbiya

Cairo

Siwa Oasis

Bani Sweif

Menia

Sohag

N”nn fFN FA fA HW Nn F&

Qena

Aswan

second clade comprises individuals from all other studied

localities. In the constructed phylogenetic trees of this spe-

cies based on COI sequences, haplotypes of the second

clade consist of two main sub-groups. The first sub-group

represents the haplotypes of Ismailia, Damietta, Alexan-

dria, Matrouh, and Gharbiya populations, which includes

one cluster of the Ismailia and Damietta populations and

a second cluster of the Alexandria, Matrouh and Gharbiya

populations. The second sub-group consists of haplotypes

of Cairo, Siwa Oasis, Bani Sweef, Menia, Sohag, Qena,

and Aswan populations, which includes one cluster of Cai-

ro, Stwa Oasis and Beni Suef populations, a second cluster

of the Menia and Sohag populations, and a third cluster of

the Qena and Aswan populations.

In addition, the phylogenetic tree generated in the

Table 2. The average values of haplotype diversity (h)

and nucleotide diversity (2) of the African Common Toad

(Sclerophrys regularis).

Population Haplotype Nucleotide

diversity (h) diversity (7)

Sharm El-Shaikh 1.000 0.53517

Arish 1.000 0.56702

Ismailia 1.000 0.47080

Damietta 1.000 0.27339

Alexandria 1.000 0.50122

Matrouh 1.000 0.42355

Gharbiya 0.900 0.38746

Cairo 1.000 0.71177

Siwa Oasis 1.000 0.47306

Bani Sweif 0.833 0.23012

Menia 1.000 0.41116

Sohag 1.000 0.55214

Qena 1.000 0.56381

Aswan 1.000 0.47523

current study using the Maximum Likelihood method

revealed that individuals of Sharm El-Shaikh and Arish

populations have the greatest genetic distances from the

other studied populations. The other individuals exam-

ined are included in a single branch. On the other hand,

the tree showed that haplotypes of Ismailia and Damietta

populations are grouped in one branch while haplotypes

of Alexandria, Matrouh and Gharbiya populations are in

a different branch next to each other in one clade and are

more closely related to each other. In contrast, haplotypes

of Cairo, Siwa Oasis and Beni Suef populations are situat-

ed in another branch close to the haplotypes of the Menia,

Sohag, Qena, and Aswan populations (Fig. 2).

Table 3. Genetic differentiation (G,,.) and diversity parameters (H.,, H,) for the combined mtDNA sequences in all studied populations

of the African Common Toad (Sclerophrys regularis).

Zoon H,

Sharm El-Shaikh 0.031

Arish 0.031

Ismailia 0.016

Damietta 0.009

Alexandria 0.016

Matrouh 0.031

Gharbiya 0.026

Cairo 0.031

Siwa Oasis 0.005

Bani Sweif 0.050

Menia 0.016

Sohag 0.016

Qena 0.009

Aswan 0.016

Total 0.00001

Amphib. Reptile Conserv.

107

H, Gy,

1.000 0.969

1.000 0.984

1.000 0.991

1.000 0.984

1.000 0.969

0.900 0.874

1.000 0.969

1.000 0993

0.833 0.783

1.000 0.984

1.000 0.991

1.000 0.984

0.999 0.99899

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Genetic diversity of Egyptian populations of Sclerophrys regularis

Discussion

It is well-established that the long-term evolution of a spe-

cies produces its genetic variation and this genetic varia-

tion represents its evolutionary potential for survival and

development (Soltis et al. 1992; Gitzendanner and Soltis

2000). During a long evolutionary history, high levels of

genetic variation are expected to accumulate. At present,

there is insufficient data to determine the evolutionary

significant units (ESU) for the conservation of Scleroph-

rys regularis. However, it seems prudent to protect and

preserve all of the habitats of Sclerophrys regularis due

to its high genetic variation and wide distribution. More

exhaustive studies should focus on more highly variable

genetic markers and on the ecology of Sclerophrys regu-

laris to establish an effective conservation strategy. As

expected, these toads were found to have high genetic

diversity and genetic differentiation at the species level.

The results provide insight into the genetic variabil-

ity of Sclerophrys regularis using mitochondrial cyto-

chrome oxidase subunit I (COI) gene analysis. Avise et

al. (1987) and Moritz (1994) reported that mitochondrial

DNA genes have been used extensively in evolutionary

biology to measure genetic variation within populations,

especially in those that diverged recently, and to assess

the conservation value of specific populations or areas.

In particular, mtDNA is useful for phylogenetic studies

because it has maternal inheritance, a mutation rate 10

times faster than nuclear DNA, and a low recombination

rate (Brown et al. 1979; Masuda and Yoshida 1994). As

a barcoding gene, COI is also widely used for many taxa

(including amphibians) for species identification and

taxonomic discovery (Marshall 2005; Salvolainen et al.

2005; Little and Stevenson 2007) because it often yields

deeper phylogeny than other genes, such as cytochrome

b (Lynch and Jarrell 1993; Simmons and Weller 2001).

The results indicate that the Siwa Oasis population

and the Damietta and Qena populations had the largest

numbers of haplotypes. On the other hand, the Matrouh,

Arish, Sharm El-Sheikh, Cairo, and Beni Suef popula-

tions had the lowest numbers of haplotypes, suggesting a

possible genetic barrier between South and North River

Nile which restricts gene flow, such as water sources, cli-

matic conditions or distances between habitats. In addi-

tion, a high haplotype diversity was found in all studied

locations, except for the Gharbiya and Beni Suef popu-

lations which had low haplotype diversity values. This

may be related to dry climate and limited water sources

in the two locations.

The nucleotide diversity was highest in the Cairo

population, while the Damietta and Beni Suef popula-

tions had the lowest nucleotide diversity. High haplotype

diversity and low nucleotide diversity in a species can

often be explained by many singular haplotypes with few

base substitutions.

Samples from the Sharm El-Shaikh and Arish locali-

ties had the greatest genetic distances to the other stud-

Amphib. Reptile Conserv.

ied populations, which may be related to differences in

habitat and climate specifically, the Sharm El-Shaikh and

Arish regions are particularly arid (Borkin et al. 2016).

In Egypt, this toad mainly inhabits savannas. Borkin

et al. (2016) reported that western Africa, the northern

part of eastern Africa, and the Nile River valley in Egypt

were the most suitable environments for Sclerophrys reg-

ularis. Distribution of this toad is restricted by the Sahara

Desert in the north and, perhaps, by concurrent closely

related species in the south, such as Sclerophrys guttura-

lis (Power 1927). The population at Sharm El-Sheikh is

highly isolated from the native range of the species by

the desert of Sinai. However, the coastal strip along the

Red Sea in the southernmost part of the Sinai Peninsula

has quite suitable environmental conditions for the sur-

vival of this species (Borkin et al. 2016).

Large agricultural reclamation projects using Nile wa-

ter are being established in northwest Sinai. Some have

suggested that Sclerophrys regularis will spread into this

region (Baha El Din 2006). Recently, at the eastern side

of the Suez Canal, Sclerophrys regularis was found in

six localities in green fields east of the Bitter Lakes up

to 9-10 km into Sinai, including the vicinities of villages

Al-Tagaddom, Al-Abtal and Meet Abul Kum Al-Jadeeda

(Ibrahim 2013a, b). As in the western bank of the Suez

Canal, Sclerophrys regularis introduced from the River

Nile is becoming quite common around freshwater trri-

gation canals.

Conclusions

The results from COI analysis show that Scleroph-

rys regularis has very high genetic diversity and gene

flow among different Egyptian populations. Therefore,

it seems reasonable to perform many additional studies

on Sclerophrys regularis due to its high genetic variation

and wide distribution. These data may indicate that the

ecological niche of this species is somewhat broader than

was revealed for populations within its native distribu-

tional range.

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Lamiaa Elsayed Mokhtar Deef received her Master’s degree in 2008 from the

Faculty of Science, Mansuora University and then a Ph.D. degree in 2014 from

the Faculty of Science, Damietta University. She is currently a lecturer at the same

faculty at Damietta University. Her research interests include genetics, taxonomy,

and phylogenetic studies.

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Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

13(1) [General Section]: 111-121 (e172).

Analysis of diet composition and morphological characters

of the little-known Peruvian bush anole Polychrus peruvianus

(Noble, 1924) in a northern Peruvian dry forest

‘Antonia Beuttner and 2>Claudia Koch

‘Universitat Tiibingen, Geschwister-Scholl-Platz, 72074 Tiibingen, GERMANY *Zoologisches Forschungsmuseum Alexander Koenig (ZF.MK),

Adenauerallee 160, 53113 Bonn, GERMANY

Abstract.—Analyses of diet composition are an important element of studies focusing on biological diversity,

ecology, and behavior of animals. Polychrus peruvianus was found to be a quite abundant lizard species in the

northern Peruvian dry forest. However, our knowledge of the ecology of this species remains limited. Herein,

we analyze the species dietary composition and the morphological features that may be related to the feeding

behavior. Our results show that P. peruvianus is a semi-herbivorous food generalist, which also consumes

faunistic prey. All age groups prefer mobile prey as sit-and-wait predators. However, during ontogenesis, plant

material becomes the main component in the diet of adult specimens.

Keywords. Iguania, bite force, ontogenetic change, foraging strategy, stomach contents, food niche, feeding behavior,

Marafion river

Citation: Beuttner A, Koch C. 2019. Analysis of diet composition and morphological characters of the little-known Peruvian bush anole Polychrus

peruvianus (Noble, 1924) in a northern Peruvian dry forest. Amphibian & Reptile Conservation 13(1) [General Section]: 111-121 (e172).

tribution 4.0 International (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in

any medium, provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced,

are as follows: official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org.

Received: 07 December 2017; Accepted: 26 July 2018; Published: 14 March 2019

Introduction

The study of diet and trophic relationships between a

species and its environment is a central theme in ecology.

Field and museum-based studies provide insights about

feeding behavior, foraging strategies, and sexual or onto-

genetic variation within species (Futuyama 1998; Clark

2002). According to their type of prey, lizards can be di-

vided into different trophic categories: insectivorous, her-

bivorous, carnivorous, or omnivorous (Vitt and Pianka

2007). Some lizard species even show an ontogenetic

shift in diet from carnivory to omnivory, or even to her-

bivory (Cooper and Vitt 2002; van Leeuwen et al. 2011).

They can be food generalists that ingest a random sample

of prey available (and also plant material), or specialists

that select specific food items (Vitt and Caldwell 2013).

Depending on a species’ behavior to locate and capture

prey, two basic foraging strategies are recognized: “sit-

and-wait” and active foraging (Vitt and Caldwell 2013).

“Sit-and-wait” foragers remain motionless and wait for

mobile prey (e.g., spiders, beetles, dipterans, and but-

terflies) to pass through their field of vision (Anderson

and Karasov 1988; Bergallo and Rocha 1994; Huey and

Pianka 1981; Magnusson et al. 1985; Nagy et al. 1984;

Pianka 1970; Pianka and Parker 1975; Toft 1985; Vitt

Correspondence. * c.koch@leibniz-zfmk.de

Amphib. Reptile Conserv.

and Caldwell 2013). In contrast active foraging species

normally prey on sedentary (e.g., insect larvae), clustered

(e.g., termites), or hidden prey (e.g., scorpions) (Bergallo

and Rocha 1994; Huey and Pianka 1981; Magnusson et

al. 1985; Nagy et al. 1984; Vitt and Caldwell 2013). Es-

pecially based on morphological variation, there may ex-

ist differences in size or type of preferred prey between

males and females (species with a distinct sexual size di-

morphism) or between adults and juveniles (Pough 1973;

Verrastro and Ely 2015; Vitt and Caldwell 2013). It is

generally accepted that lizards with higher and broader

heads are capable of producing stronger bite forces

(Campos 2016; McBrayer and Corbin 2007; Miles et al.

2007) and are thus able to consume harder dietary items

(Herrel et al. 2001a, b; Herrel 2007; Huyghe et al. 2009;

Verwaijen et al. 2002). Campos (2016) and Herrel et al.

(2006) subdivided food items in “hard,” “intermediate,”

and “soft” prey, the latter requiring the lowest bite forces.

Knowledge of a species’ diet as part of its natural history

may be crucial for the implementation of conservation

actions for a species and its environment (Greene 1994;

Verrastro and Eli 2015).

Until a few years ago the polychrotid lizard Poly-

chrus peruvianus (Squamata: Iguania) was only known

from a few specimens, and sufficient data was lacking

March 2019 | Volume 13 | Number 1 | e172

Beuttner and Koch

for a proper assessment of its conservation status. Conse-

quently, it was listed as Data Deficient by the IUCN Red

List of Threatened Species (Cisneros-Heredia 2010).

Recent fieldwork (Koch et al. 2011) provided new in-

sights into the distribution and ecology of the species and

contributed to an updated assessment of its conservation

status. Hence, the species is now listed as Vulnerable by

the IUCN due to its relatively small extent of occurrence

(~12.000 km7?), the low number of known locations (<5),

and a decline in the extent and quality of its habitat, due

to logging, agricultural expansion, and hydroelectric dam

constructions (Venegas et al. 2017). Nevertheless, dur-

ing our intensive fieldwork in different inter-Andean dry

forest regions in northern Peru, we could lately provide

records of P. peruvianus for 12 of 28 surveyed localities

in the departments of Amazonas and Cajamarca. As per

the research permits, we collected up to six specimens

per locality for preservation and examination, to further

contribute to the knowledge of this scarcely known spe-

cies. Surprisingly, the species seemed to be quite abun-

dant in most surveyed regions, as we often found many

more individuals than just the ones we were allowed to

collect (Koch et al. 2011, 2018). The species is further

known from the Zamora-Chinchipe province in south-

ern Ecuador and Peruvian department of Piura at eleva-

tions of 400—-1,750 m a.s.l. (Carrillo and Icochea 1995;

Duellman 1979; Gorman et al. 1969; Koch et al. 2011;

Noble 1924; Peters and Donoso-Barros 1970; Schliiter

2010; Torres-Carvajal et al. 2017; Yafiez-Mufioz et al.

2006). Polychrus peruvianus is diurnal and highly arbo-

real, being almost exclusively found on trees or shrubs

(Koch et al. 2011). Males and females show a conspicu-

ous sexual dimorphism in coloration; females have lime

green heads and a straight white stripe laterally between

the axilla and the insertion of the hind limbs, whereas the

heads of males are brownish and a white lateral stripe is

lacking (Koch et al. 2011). The congeners P. marmora-

tus and P. acutirostris are known to feed on arthropods

as well as plant material (Garda et al. 2012; Koski et al.

2016). However, the only available information regard-

ing the diet of P. peruvianus derives from a single indi-

vidual from Tingo in the Peruvian department of Amazo-

nas, whose stomach contained several hymenoptera and

a leaf fragment (Gorman et al. 1969).

In the present study, we sought to describe the diet

of populations of Polychrus peruvianus from the inter-

Andean dry forest valley in northern Peru to provide

information that may be relevant for the establishment

of conservation criteria for this species. Therefore, we

analyzed the trophic niche to reveal sexual differences

and ontogenetic changes in diet composition, to define

the foraging strategy, and to determine whether members

of the species are food generalists or specialists. Further-

more, we performed morphological analyses in order to

detect sexual dimorphism that may be related to differ-

ences in feeding behavior.

Amphib. Reptile Conserv.

Material and Methods

In the framework of an intensive herpetological inven-

tory in the inter-Andean dry forest valleys of the Mara-

fion river and its tributaries we collected a total number

of 64 individuals of Polychrus peruvianus (Koch et al.

2018). Most of these specimens were deposited in the

collections of the Centro de Ornitologia y Biodiversidad

(CORBIDI), in Lima, Pert, and the Museo de Histo-

ria Natural de la Universidad Nacional de San Agustin

(MUSA), in Arequipa, Pert. However, we were allowed

to export 24 individuals (9 males, 10 females, 5 juveniles)

for further examination and to add them to the collection

of the Zoologisches Forschungsmuseum Alexander Koe-

nig (ZFMK), in Bonn, Germany. The specimens housed

in the ZFMK built the basis of the dietary analysis, pre-

sented herein. We detected these specimens between

May 2008 and December 2009 at different localities dur-

ing visual encounter surveys after nightfall, when they

were sleeping on branches of trees and shrubs in heights

between 1.5 m and 7.0 m above the ground (Koch et al.

2018). We collected one individual in May 2008 (ZFMK

88707), two in June 2008 (ZFMK 88708-09), four in July

2008 (ZFMK 88710-13), four in March 2009 (ZFMK

90817-20), two in April 2009 (ZFMK 90821-22), and

eleven in December 2009 (ZFMK 90823-33). Exact lo-

calities of each specimen are given in the Appendix. We

either captured the specimens by hand or by use of a fish-

ing net and euthanized them by an injection of the veteri-

nary anesthetic T61 the following morning, no later than

10 hours after their capture, to guarantee that the stomach

contents were little digested and in good condition for

identification. After fixation in 10% formalin for 24 to

48 hours, the specimens were preserved in 70% ethanol.

Sexes of the specimens were determined by coloration

and/or by internal reproductive organs (testicles or ova-

ries) or by the presence of everted hemipenes. We catego-

rized individuals with a snout-vent-length (SVL) of less

than 100 mm as juveniles, since none of the dissected liz-

ards up to this size had mature gonads.

Definition of head length, head width, and head height

follows Meyers et al. (2002). Measurements of head

(length, width, height), body (SVL, width) and tail length

of the lizards and dimensions of prey items (length, width)

were taken with a dial caliper (0 — 150 mm; to the nearest

0.01 mm) and weights of bodies, fat bodies, and stomachs

were calculated with a digital weighing machine (Alma-

sa® MT 7; max. 200 g; to the nearest 0.01 g). The bodies

of the preserved specimens were opened ventrally and fat

bodies and stomachs were removed and the latter were dis-

sected.

Stomach contents were placed on a Petri dish and

prey items were analyzed under a stereomicroscope and

were verified to at least the level of order according to

Bahrmann (2011). Bergallo and Rocha (1994) and Cooper

(1994) defined Formicidae, Isoptera, and larvae of Diptera

March 2019 | Volume 13 | Number 1 | e172

Diet of Polychrus peruvianus

and Lepidoptera as “sedentary prey” and Araneae, Cole-

optera, Diptera (adult), Hemiptera, Hymenoptera (exclud-

ing Formicidae), Lepidoptera (adult), Odonata, Orthoptera

and Lithobiomorpha as “mobile prey.” Thus, in order to

determine whether the lizard species is a sit-and-wait or

active forager, we subclassified the orders Hemiptera into

“Heteroptera” and “Other Hemiptera,” Hymenoptera into

“Formicidae” and “Other Hymenoptera,’ Lepidoptera

into “adult Lepidoptera” and “Lepidoptera Larvae,” and

likewise we subclassified Diptera into “adult Diptera”

and “Diptera larvae.” Leaves, fibers, seeds and fruits were

grouped together as “Plant material.” To identify the bite

force food items were divided into three groups: hard prey,

medium hard prey, and soft prey according to Campos

(2016) and Herrel et al. (2006). Coleoptera, Hymenop-

tera, and plant material were considered as “hard prey,”

Hemiptera, Formicidae, Isoptera, Odonata, Orthoptera,

and Lithobiomorpha as “medium hard prey,” and Araneae,

Diptera, Lepidoptera, and larvae as “soft prey.” All food

items were quantified and measured.

The number of consumed items, the percentage, the

frequency of occurrence (number of stomachs in which a

given prey item was found), the percent by frequency, the

volume, and the percent by volume for each prey category

were estimated.

All statistical tests were executed with the software

OriginPro version 8.0724 (OriginLab, Northampton, Mas-

sachusetts, USA). Before comparisons, the Shapiro-Wilk-

Test was used (W < 0.966, p < 0.86) to check if data were

normally distributed and depending on the result paramet-

ric (t-test) or non-parametric (Mann-Whitney U-test) tests

were used for data analysis.

For comparison of the head dimensions, regressions

were performed for every dimension against SVL and a

residual analysis was completed to avoid influence of dif-

ferent SVL values within the species. Similarly, for com-

parison of the weights, regressions were conducted on the

weights of fat bodies and stomachs against body weight

and residual analyses were completed to avoid influence

of body weight values.

Prey items were subdivided according to the mean val-

ues of their size (very large, large, medium-sized, small,

very small) to estimate the length and width of incomplete

prey items for each order (see Supplement Table S1).

The length (L) and width (W) of the individual prey

items were calculated for prey volume (V) with the formu-

la for an ellipsoid according to Colli and Zamboni (1999):

To quantify the diversity of prey used by the animals,

niche breadth (B) was calculated using the Simpson-Index

B (Simpson 1949):

_ 4 ys ni(ni-1)

B=1 i=1 n(n-1)

Amphib. Reptile Conserv.

where n, is the number of prey items in each category i

(prey items of different orders and plant material) and n is

the total number of prey items. Niche breadth values vary

from 0 (no diversity, exclusive use of a single prey type,

specialist) to 1 (highest diversity, prey items of all catego-

ries, generalist).

Additionally, the inverse of Simpson-Index B’ was

used to compare the values with other published data:

nj(ni-1)

ri S

B= 1/ Qi=1 PEt

where niche breadth values (B’) vary from 1 (no diversity,

exclusive use of a single prey type, specialist) to n (highest

diversity, prey items of all categories, generalist).

The Index of Relative Importance (IRI) was calculated

for each prey category (1) in relation to the total food spec-

trum with the following formula:

where PO, is the percent by frequency F% (100 x number

of stomachs which contain the prey items i/total number of

stomachs), PI, is the percent by number N% (100 x num-

ber of prey items of each category i/total number of prey

items), and PV, is the percent by volume V% (100 x vol-

ume of prey items of each category i/total volume of prey

items) of each prey category.

The Pianka-Index (Pianka 1974) was estimated to cal-

culate niche overlap in the food spectrum of the different

groups of examined animals (females, males, juveniles,

adults):

On Li PijPix

Di PF DT Pi

where P. and P.. represent the number of prey categories

i used in the groups j and k to be compared. The value for

niche overlap (O,,) varies from 0 (no overlap, the com-

pared groups have a completely different food spectrum)

to 1 (complete overlap, the compared groups have the

same food spectrum).

Results

Morphological analysis. There was no difference in

SVL and weight of males and females (length: U-test,

U =41, Z = -0.2858, p = 0.78: weight: t-test, t= -0.07 p

= 0.94; see Supplement Table S2). All head dimensions

were positively correlated with the SVL (head length

against SVL: y = 0.262x + 1.783, R? = 0.854; head width

against SVL: y = 0.157x — 0.076, R? = 0.831; head height

against SVL: y = 0.130x + 2.204; R* = 0.684). There

were no differences between adults and juveniles for all

the head dimensions’ residuals (head length: test: t =

-0.44, p=0.66; head width: ¢-test: t= 0.68, p = 0.50; head

height: ¢-test: t = -1, p = 0.30), but males showed lon-

ger, wider and higher heads than females (head length:

March 2019 | Volume 13 | Number 1 | e172

Beuttner and Koch

t-test: t= -3.99, p = 0.0009; head width: r-test: t= -3, p=

0.006; head height: t-test: t = -4, p = 0.001; see Supple-

ment Table S2).

Fat bodies were found in 21 individuals (87.5%) of P.

peruvianus (8 males, 10 females, 3 juveniles). Although

there was no difference between adults and juveniles for

fat body weight residuals (U-test: U =41, Z = 1.3568, p=

0.17), females had heavier fat bodies than males (U-test:

U=69, Z = 2.53229, p< 0.05). Further, there was no dif-

ference between males and females for stomach weight

residuals (t-test, t= 0.79380, p = 0.44), but juveniles had

heavier stomachs than adults (t-test, t = 4.46367, p =

0.03) (see Supplement Table S2).

Stomach contents. Table 1 shows the composition of the

stomach content of Polychrus peruvianus for all speci-

mens together and by sex/age classes. Altogether, 226

items were identified, representing 195 faunistic prey

items (N% = 86.3 %) and 31 pieces of plant material (N%

= 13.7 %), which were found in 21 stomachs (F% = 87.5

%). The percent by volume of the plant material (V% =

75.25 %) was three times higher than that of the faunistic

prey items (V% = 24.75 %). However, with respect to the

IRI the latter (IRI = 9715.4) played a slightly more im-

portant role in the species’ diet than plant material (IRI =

7784.6). Inorganic material (e.g., stones) was not found.

In 19 stomachs of adult individuals 105 food items were

found with a total volume of V = 31651.8 mm°, whereas

121 food items were found in five stomachs of juveniles,

that had a total volume of V = 5630.4 mm‘%. In adults

plant material was more important with an IRI = 9830.7

than faunistic prey items with an IRI = 7589.7, whereas

in juveniles the latter played a far more important role in

the diet with an IRI = 16989.9 than plant material with

an IRI = 2408.1. In nine stomachs of male individuals

34 food items were found with a total volume of V =

10994.0 mm?, whereas 71 food items were found in 10

stomachs of female specimens with a total volume of V

= 20657.9 mm*. In males plant material was more impor-

tant with an IRI = 11387.0 than faunistic prey items with

an IRI = 4793.1, whereas in females the latter played a

slightly more important role in the diet with an IRI =

9900.6 than plant material with an IRI = 9089.5.

The most dominant faunistic prey category was the

order Coleoptera with respect to the number of consumed

items (N = 76, N% = 33.6 %), the frequency in how many

stomachs the category was found (F = 14, F% = 58.3 %),

the volume (V = 2400.7 mm?; V% = 6.4 %), and the IRI

(IRI = 2336.1). The second most important category was

the suborder Heteroptera with respect to the frequency

(F = 9, F% = 37.5), the volume (V = 1938.6 mm?, V% =

5.2 %), and the IRI (IRI = 642.4). However, the number

of Heteroptera items (N = 27, N% = 12.0 %) was slightly

lower than that of the order Diptera (N = 30, N% = 13.3

%) which ranked third in volume (V = 1135.2 mm?, V%

= 3.0 %) and fourth in IRI (IRI = 203.9) but only played

an underpart with respect to the frequency (F = 3, F% =

12.5 %). The order Orthoptera ranked third in frequency

(F = 7, F% = 29.2 %, together with Hymenoptera) and

IRI (IRI = 219.4) and fourth with respect to the volume

(V = 993.2 mm?, V% = 2.7 %). On closer inspection of

the IRI, the Coleoptera played a major role in the diet of

adults (IRI = 1247.7), juveniles (IRI = 4656.6), and fe-

males (IRI = 2889.7), but only ranked fifth in the diet of

males (IRI = 74.4). The second most important category

in adults (IRI = 962.3) and females (IRI =1648.7) was

Heteroptera, which ranked third in males (IRI = 265.1)

and fifth in juveniles (IRI = 475.7). The Lepidoptera lar-

vae ranked third in adults (IRI = 270.6) and was even

the most important prey category in males (IRI = 637.4),

however it ranked only fifth in females (IRI = 121.3)

and played an underpart in juveniles (IRI = 59.3). Im-

portant in the diet of juveniles were also the categories

Diptera (rank 2, IRI = 1729.2), Hymenoptera (rank 3, IRI

= 967.8), and Hemiptera (rank 6, IRI = 348.3), which

only played an underpart in males and females. Orthop-

tera ranked third in females (IRI = 199.4) and fourth in

adults (IRI = 134.4) as well as in juveniles (IRI = 808.4)

(Table 2).

With regard to the food niche breadth, the Simpson-

Index for the species was B = 0.82 and the inverse Simp-

son-Index was B’ = 5.65. Values did not differ greatly

between sexes (males: B = 0.76, B’ = 4.19; females: B =

0.82, B’ = 5.63), whereas juveniles had noticeably lower

Table 1. Composition of the stomach content (faunistic prey items (FP), plant material (PM)) of Polychrus peruvianus. Data were

obtained by pooling stomachs of all specimens and by separating them according to sex/age classes. For each category the number

(N), the percent by number (N%), the frequency (F), the percent by frequency (F%), the calculated volume (V [mm*]), the percent

by volume (V%), and the Index of Relative Importance (IRI) are shown.

Adults (n = 19) Juveniles (n = 5)

Males (n = 9)

Females (n = 10) Total (n = 24)

N 78 vi ales; 4 20 14 58 Is 195 31

N% 74.3 DES alt 96.7 ou) 58.8 41.2 81.7 18.3 86.3 L387

F 16 17 5 4 6 8 10 9 P| 21

F% 84.2 89.5 100.0 80.0 66.7 88.9 100.0 90.0 87.5 87.5

V[mm*} 5014.2 26637.6 4121.7 1508.7 1437.2 9556.8 soda) 17080.9 9257.2 28146.3

V% 15.8 84.2 Foe 26.8 3a] 86.9 LS S24 24.75 te,

IRI 7589.7 9830.7 16989.9 2408.1 4793.1 11387.0 9900.6 9089.5 9715.4 7784.6

Amphib. Reptile Conserv. 114 March 2019 | Volume 13 | Number 1 | e172

Diet of Polychrus peruvianus

Table 2. Faunistic prey items of Polychrus peruvianus. For each category the number (N), the percent by number (N%), the

frequency (F), the percent by frequency (F%), the calculated volume (V [mm/°]), and the percent by volume (V%) are shown for all

specimens of the species together. The Index of Relative Importance (IRI) of each prey category is given for adults, juveniles, males,

and females separately and for all specimens together, respectively.

Araneae 8 BAD 2 20.8 90.3

Coleoptera 76 336 14 58.3 2400.7

Diptera 30.0) 13.3 3 1S 1135.2

Formicidae 5 22 4 16.7 8.3

Hemiptera 1] 49 3 a3 751.8

Heteroptera Dis) ZO 9 375 1938.6

Hymenoptera 10 4.4 7 Dore. 633.4

Lepidoptera adult 1 0.4 1 4.2 191.7

Lepidoptera larvae 10 44 6 25.0 661.6

Lithobiomorpha l 0.4 l 4.2 3.6

Odonata 1 0.4 l 4.2 63.5

Orthoptera 1] 49 7 29.2 993.2

Undetermined 4 1.8 S) 1255 385.3

values than adults (adults: B = 0.84, B’ = 6.43; juveniles:

B = 0.74, B’ =3.78; Table 3).

The Pianka-Index (O;,) showed highest niche overlap

value between females and juveniles (O;, = 0.91), where-

as males and juveniles had the lowest overlap value (0,

= 0.74). Adults and juveniles (Oo = 0.88) showed almost

a similar value for the overlap as males and females (O;,

= 0.87).

Foraging strategy. In 21 stomachs of Polychrus peru-

vianus, 191 faunistic prey items were determined with

a total volume of V = 8871.8 mm*, of which 176 were

assigned to the category mobile prey (N% = 92.15 %)

with a volume of V = 8202.0 mm? (V% = 92.45 %) and

15 (N% = 7.85 %) belonged to sedentary prey with a

volume of V = 669.8 mm? (V% = 7.55 %). Mobile prey

was found in 19 stomachs (F% = 90.5 %) and sedentary

prey in 10 (F% = 47.6 %). The IRI of mobile prey (IRI =

16701.6) was almost 23 times higher than the IRI of sed-

entary prey (IRI = 733.5). With respect to the different

specimen groups (adults, juveniles, males, females) con-

siderably higher values were examined for mobile prey

than for sedentary prey (except for the same frequency

with which mobile and sedentary prey were found in

males). In adults the IRI for mobile prey was about 11

times higher than for sedentary prey and in juveniles it

was even about 103 times higher. In adults the IRI value

for mobile prey was about three times higher than for

sedentary prey and in females it was even about 21 times

higher (Table 4).

Bite force. Of the 226 food items found in the 24 stom-

achs of Polychrus peruvianus four faunistic prey items

were not determined (Table 2). Hence, for the bite force

Amphib. Reptile Conserv.

0.2 hu 525 32.8 144.1 78.8

6.4 1247.7 4656.6 74.4 2889.7 = 2336.1

3.0 Shy, 1729.2 0 16.5 203.9

0.02 60.3 18.3 32.8 84.8 oz

2.0 34.9 348.3 74.8 16.5 86.0

5.2 962.3 475.7 265.1 1648.7 = 642.4

Ly. 71.7 967.8 40.4 99:5 178.5

0.5 0 84.6 0 0 40

1.8 270.6 593 637.4 eS 154.8

0.01 0 17.8 0 0 1.9

0.2 0) ey hl 0 0 2.6

2: 134.4 808.4 63.4 199.4 219.4

1.0 79.4 0) 270.4 14.9 35.0

analyses 222 items remained, of which 117 were as-

signed to the category hard prey (N% = 52.7 %) with a

volume of V = 31180.3 mm? (V% = 84.2 %), 56 (N% =

25.2 %) belonged to medium hard prey with a volume

of V = 3759.0 mm? (V% = 10.2 %) and 49 (N% = 22.1 %)

belonged to soft prey with a volume of V = 2078.8 (V% =

10.2 %). Hard prey items were found in 23 stomachs (F%

= 95.8 %), medium hard prey in 15 (F% = 62.5 %) and

soft prey in 13 stomachs (F% = 54.2 %). The IRI of hard

prey (IRI = 13122.7) was the highest, followed by the IRI

of medium hard prey (IRI = 2211.2). With respect to the

different specimen groups all examined values were con-

siderably higher for hard prey than for medium hard prey

or soft prey (except for the same frequency with which hard

and medium hard prey was found in juveniles). In adults,

the IRI for hard prey was about six times higher than for

medium hard prey and 14 times higher than for soft prey.

In juveniles, the IRI of hard prey was two times higher than

for medium hard prey and three times higher than for soft

prey. In females, the IRI for hard prey was four times higher

than for medium hard prey and 15 times higher than for soft

prey. In males, the IRI of hard prey was 12 times higher

than the IRI for medium hard prey or soft prey (Table 5).

Table 3. Diversity in the dietary spectrum of Polychrus

peruvianus. The Simpson-Index (B), the inverse Simpson-

Index (B’), and the number of consumed prey categories

(S, including plant material) are given for adults, juveniles,

males, and females separately and for all specimens together,

respectively.

B 0.84 0.74 0.76 0.82 0.82

Bs 6.43 3.78 4.19 5.63 5.65

S 10 1k, ] 10 ike,

March 2019 | Volume 13 | Number 1 | e172

Beuttner and Koch

Table 4. Mobile (MP) and sedentary prey (SP) items in the stomach content of Polychrus peruvianus for all specimens together and

by sex/age classes. The number of items (N), the percent by number (N%), the frequency (F), the percent by frequency (F%), the

calculated volume (V [mm?]), the percent by volume (V%) and the Index of Relative Importance (IRI) are shown.

Adults (n = 16)

Juveniles (n = 5) Males (n = 6) Females (n = 10) Total (n = 21)

N 65 13 115 2 13 7 Sy 6 176 15

N% 83.3 16.7 98.3 | ies 65.0 35.0 89.7 10.3 OAS 7.85

F 14 8 5 2 4 4 10 4 19 10

F% 87.5 50.0 100.0 40.0 66.7 66.7 100.0 40.0 90.5 47.6

V [mm] 4469.8 544.4 3996.3 1235 1274.4 162.8 3195.5 381.6 8202.0 669.8

V% 89.1 ee 97.0 3.0 88.7 elves 89.3 10.7 92.45 eS

IRI L509 bie. 1376.2 19524.7 190.1 10244.7 3088.6 17898.9 840.5 16701.6 Too

Discussion cies as compared to adults. Additionally, in juveniles the

This is the first detailed study of the diet of Polychrus pe-

ruvianus, based on specimens collected in the dry forest

of northern Peru. Findings suggest that P peruvianus is a

semi-herbivorous food generalist, which also consumes

faunistic prey. All age groups prefer mobile prey as sit-

and-wait predators. However, during ontogenesis, plant

material becomes the main component in the diet of adult

specimens.

Although in Polychrus acutirostris, P. gutturosus, P.

Jacquelinae, P. liogaster, and P. marmoratus a sexual

size difference with females growing larger than males

is stated (Avila-Pires 1995; Garda et al. 2012; Koch et

al. 2011), this was not observed in our specimens of P.

peruvianus examined herein. Nevertheless, a relatively

small number of specimens was analyzed and a larger

sample size would allow a more reliable estimation

whether a sexual size dimorphism exists in this spe-

cies. No significant differences between most specimen

groups in fat body weight and stomach weight in propor-

tion to body weight were observed. It can be assumed

that the examined specimens had a similar nutritional

state, except for females of P. peruvianus, who were in a

better nutritional state with heavier fat bodies in propor-

tion to body weight. Juveniles of P. peruvianus proved to

have heavier stomachs than adults in proportion to body

weight. This coincides with the fact that distinctly more

prey items were found in juvenile stomachs of this spe-

percent by volume of faunistic prey items (V% = 73.2

%), Which are usually heavier, was much higher as com-

pared to plant material (V% = 26.8 %), whereas stomachs

of both adult sexes contained a significantly higher per-

cent by volume of plant material as compared to faunistic

prey items (86.9 % vs. 13.1 % in males, 82.7 % vs. 17.3

% in females). This observation further suggests that P.

peruvianus has a semi-herbivorous dietary spectrum, as

was also observed in the congeners P. acutirostris and P.

marmoratus (Avila-Pires 1995; Garda et al. 2012; Van-

zolini 1983). In adult specimens of P. peruvianus, plant

material plays a greater role (IRI = 9830.7) in the di-

etary spectrum than faunistic prey items (IRI = 7589.7).

Nevertheless, the importance of plant material is much

lower in the diet of juveniles (IRI = 2408.1) compared

to adults (IRI = 9830.7). Many lizard species undergo an

ontogenetic diet shift from eating insects as a juvenile to

an herbivorous or semi-herbivorous diet as an adult (..e.,

Cooper and Vitt 2002; Dessem 1985; Estes and Williams

1984; van Leeuwen et al. 2011). This is explained by the

fact that juvenile lizards need more animal protein for

their growth and development (Johnson and Lillywhite

1979; Mayhew 1963). Furthermore, juveniles are less ef-

ficient at digesting plant material than older lizards, until

they acquire the intestinal flora that degrades cellulose

(Troyer 1982).

With a relatively high value for the Simpson-Index (B

= 0.82) Polychrus peruvianus can be considered as a food

Table 5. Hard (H), medium hard (M) and soft prey (S) items in the stomach content of Polychrus peruvianus for all specimens

together and by sex/age classes. The number of items (N), the percent by number (N%), the frequency (F), the percent by frequency

(F%), the calculated volume (V [mm*]), the percent by volume (V%) and the Index of Relative Importance (IRI) are shown.

Adults (n = 19) Juveniles (n = 5)

34.7 54.8

11 8

94.7 57.9 52.6 100.0 100.0 60.0 88.9

TPO,

91.6 6.4 2.0 45.0 29.2 25.8 11S)

13007.6 984.8

28644.8 1994.4 627.3 | 2535.6 1643.4 1451.5

13369.3. 2375.6 939.4 | 10040.9 46543 3183.1

Amphib. Reptile Conserv.

Males (n = 9)

Females (n = 10) Total (n = 24)

7 28 g 117

22.6 40.0 12.9 Sw DIS\ 2) 22.1

3 8 6 23 15 13

33.3 100.0 80.0 60.0 95.8 62.5 54.2

740 18922 1254.5 463.8 | 311803 3759 2078.8

7.0 ; 91.7 6.1 Das) 84.2 10.2 5.6

13881.8 3686.2 906.2 | 13122.7 2211.2 1499.8

March 2019 | Volume 13 | Number 1 | e172

Diet of Polychrus peruvianus

generalist with a great diversity in consumed food items.

The congener P. acutirostris has a lower value (B = 0.67;

calculated with the data of Garda et al. 2012) and thus

shows a lower diversity in the dietary spectrum. Adult

P. peruvianus exhibit the greatest diversity (B = 0.84),

whereas juveniles show the lowest diversity (B = 0.74).

The niche overlap in the dietary spectrum of the dif-

ferent subgroups of P. peruvianus 1s relatively high with

all values for 0; 2074. The highest overlap in the food

niches exists between juveniles and females (O7= 0.91),

whereas the niches of the juveniles and males show the

lowest overlap values Or = 0.74). One reason for this

could be that male specimens have higher and wider

heads, which enable them to consume larger prey items

(Pough 1973). Additionally, their head dimensions al-

low them to produce higher bite forces (Campos 2016;

McBrayer and Corbin 2007; Miles et al. 2007), whereby

they are able to feed on harder dietary items (Herrel et

al. 2001a, b; Herrel 2007; Huyghe et al. 2009; Verwaijen

et al. 2002). Thus, male specimens are able to consume

a greater diversity of food items. In fact, females have

as well larger heads than juveniles, but the difference 1s

smaller than between males and juveniles. “Hard prey”

items are the preferred category in the spectrum of con-

sumed food in all subgroups of P. peruvianus, although

the percent by volume in juveniles is only half as high as

in adults, indicating that juveniles are less able to con-

sume a broad spectrum of hard prey items. Plant material

also belongs to hard prey and as already mentioned 1s

consumed to a lesser amount by the juveniles.

The most important faunistic prey categories in the

diet of all subgroups represent mainly mobile insects

which are capable of flying (Coleoptera, Heteroptera,

Diptera, and Orthoptera) and are thus easily accessible

for arboreal animals (Hogue 1993) such as P. peruvia-

nus. Almost no sedentary prey items were found (e.g.

larvae, ants, and termites) in the stomachs of this species.

A similar preference for mobile, volant, or arboreal prey

was observed for the congener P. acutirostris (Garda et

al. 2012). Mobile prey is preferred by sedentary spe-

cies with a sit-and-wait foraging strategy (Anderson

and Karasov 1988; Bergallo and Rocha 1994; Huey and

Pianka 1981; Magnusson et al. 1985; Nagy et al. 1984;

Pianka 1970; Pianka and Parker 1975; Toft 1985). Thus,

it can be assumed that P peruvianus is a sit-and-wait

predator, which lives up in trees and mainly preys on

mobile species that are able to fly or climb trees. This

coincides with the general assumption that sedentary or

hidden prey types are relatively unavailable to iguanian

lizards (Vitt and Pianka 2005). Our observed great diver-

sity in the dietary spectrum coincides with the general

observation that sit-and-wait predators consume a wide

variety of prey types as they spend a larger amount of

time motionless, whereas actively foraging species can

be more selective (Bergallo and Rocha 1994).

Although Polychrus peruvianus seems to be quite

abundant in many localities studied herein the popula-

Amphib. Reptile Conserv.

tion size and exact distribution range of the species has

never been determined. Most surveyed localities are fac-

ing multiple and complex threats due to logging, agri-

culture and narcotics plantations, mining, and several

planned hydroelectric projects that will lead to flooding

of vast portions of the species’ habitat (Koch et al. 2013).

These threats, together with the enormous lack of knowl-

edge that still exists about the biodiversity of the region,

and no protected area has so far been designated in the

inter-Andean dry forest, highlight the urgent priority

for conservation and research activities in this area. It is

likely that populations of P. peruvianus and many other

endemic species will decline in the near future without

further knowledge on their natural history and activities

to protect their habitat.

Acknowledgements.—CK_ thanks the Deutscher

Akademischer Austauschdienst (DAAD), the Alexander

Koenig Stiftung (AKS) and the Alexander Koenig Gesell-

schaft (AKG) for financial support. The Ministerio de Ag-

ricultura, Peru kindly provided collecting (071-2008-IN-

RENA-IFFS-DCB, 0020-2009-AG-DGFFS-DGEFFS,

0424-2010-AG-DGFFS-DGEFFS) and export _ per-

mits (0017799-AG-INRENA, 001829-AG-DGFFS,

003983-AG-DGFFS). For assistance during fieldwork

we are indebted to Alfredo Beraun, Sibylle Duran Zo-

pazo, Marco Enciso, Antonio Garcia Bravo, Erick Hoyos

Granda, Jorge Novoa Cova, Napoleon Monsalve, and

Manuel Palacios Panta.

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Antonia Beuttner is currently writing her doctoral thesis at the institute of microbiology in Stuttgart

Wolfgang Wagele.

Amphib. Reptile Conserv.

(Germany). She performed her master thesis “Identification of serum resistance genes in Pseudomonas

aeruginosa by Transposon Directed Insertion Sequencing (TraDIS)’ at the University Tubingen

(Germany) in October 2018. Her bachelor thesis “Vergleichende Nahrungsanalysen der sympatrischen

Echsenarten Microlophus stolzmanni und Polychrus peruvianus” (=Comparative nutritional analyzes

of the sympatric lizard species Microlophus stolzmanni and Polychrus peruvianus) was submitted in

September 2016 at the University of Bonn, under the supervision of Dr. Claudia Koch and Prof. Dr.

Claudia Koch is currently curator of herpetology at the Zoological Research Museum Alexander

Koenig in Bonn (ZFMK). She studied biology at the University Bonn (Germany), and conducted her

diploma and doctoral thesis at the ZFMK, with focus on the diversity of amphibians and reptiles from

Peru. Claudia discovered and described Polychrus jacquelinae and provided information on several

congeners. This is her third published contribution to the genus Polychrus.

March 2019 | Volume 13 | Number 1 | e172

Beuttner and Koch

APPENDIX

Examined specimens

Polychrus peruvianus. AMAZONAS: Bagua: Bagua Grande: ZFMK 88712, 88713 (05°477°33.”S, 78°23’04.9" W,

570 m a.s.l.); Utcubamba: Cumba: ZFMK 90830-90833 (05°56’S,78°39’ W, 450-500 m a.s.l.); Zapatalgo/Chiiu-

fia: ZFMK 90823-90825 (06°04’S, 78°29’ W, 900—1,030 m a.s.l.); Puerto Malleta: ZFMK 90826-90828 (06°03’S,

78°36’ W, 480-510 m a.s.l.); CAJAMARCA: Jaén: Santa Rosa de la Yunga: ZFMK 88710 (05°26’S, 78°33’W,

1,250-1,300 m a.s.l.); Bellavista: ZFMK 88707 (05°39’49.8”S, 78°40’713.9” W, 411 ma.s.l.); ZFMK 90818, 90819

(05°38’06.6”S, 78°39°36.2” W, 405 m a.s.l.); ZFMK 90820 (05°34’35.7”S, 78°38’ 10.8” W, 700 m a.s.l.); Gotas de

Agua: ZFMK 88708 (05°41’S, 78°46’ W); Perico: ZFMK 88709 (05°21716.5”S, 78°47’30.6”W, 443 ma.s.1.); ZFMK

90822, 90821 (05°21’S, 78°47’ W, 460-720 m a.s.l.); Pucara: ZFMK 88711, 90817, (06°02’S, 79°07’ W, 900-930 m

a.s.l.); Cutervo: Across from Puerto Malleta: ZFMK 90829 (06°04’24.5”S, 78°36°47.8” W, 535 maz.s.l.).

Supplementary material

Supplementary Table S1. Mean values of prey item sizes to estimate the length and width of incomplete prey items for each order.

Orders of which only complete prey items were found are not listed.

large 5:0) 2.6

medium 3.6 1.9

Araneae

small Le Lee

very small ne 0.5

very large 16.4 10.9

large 12.8 O27.

Coleoptera

medium 4.7 Da

small 28 0.9

very large 89 27

large 6.8 25s)

Formicidae medium 40 0.9

small 2.3 0.6

very small 1.6 0.4

large 11.6 6.3

Heteroptera medium 8.9 5.1

small 48 23

medium es ae)

Hymenoptera

small 8.1 27

large 29.9 49

medium 14.2 32

Lepidoptera Larve

small 10.0 ES

very small Ont 0.9

large 17.4 4.5

Orthoptera

medium 13.1 Bue

Amphib. Reptile Conserv. 120 March 2019 | Volume 13 | Number 1 | e172

Diet of Polychrus peruvianus

Supplementary Table S2. Measurements of Polychrus peruvianus: Snout-vent-length [mm], body width [mm], body weight [g],

head length [mm], head width [mm], head height [mm], fat body weight [g], percental fat body weight [%], stomach weight [g],

and percental stomach weight [%] of females, males, juveniles and all animals together (total). Shown is the range, the mean value

(x) and the standard deviation (SD).

Females Males Juveniles Total

eee 9 10/10 9/8 5/3 24/21

Snout-Vent-Length [mm] Range 106-138 116-135 74-96 74-138

x+SD 123.44 11.6 127.0+6.4 90.4+ 8.4 117.9+ 17.0

Body weight [g] Range 24-68 30-57 9-20 9-68

x+SD 40.1+12.1 404+ 8.9 14.7+3.6 34.94 14.2

Head length [mm] Range 29-37 34-40 21-28 21-37

x+SD 32.8+2.9 36.6+1.8 2S ene 32.7448

Head width [mm] Range 16-21 18-24 12-15 12-24

+ SD 18.74 1.5 20.8 + 1.6 13:9-41.0 18. 542,9

Head height [mm] Range 15-19 18-23 12-16 12-23

x+SD ve calls} 20.0 + 1.5 14.0+1.5 eG eean

Fat body weight [g] Range 0.01-1.3 0.01-0.3 0.01-0.1 0.01-1.3

x+SD 0.53 +0.38 0.114+ 0.09 0.03 + 0.03 0.30 + 0.35

Fat body weight [%] Range 0.04-2.6 0.03-1.0 0.08-0.4 0.03-2.6

x+SD 2s Ore 0.30 + 0.28 O20 O15 0.74 + 0.75

Stomach weight [g] Range 0.9-1.9 0.6-2.6 0.5-1.7 0.5-2.6

x+SD 1.37+40.24 1.35 40.57 1.01 40.45 1.29 + 0.46

Stomach weight [%| Range 2.1-5.7 2.0-4.9 4.3-9.9 2.0-9.9

x+SD 3.66 + 1.01 3.28 + 0.95 6.60 + 1.97 2A be ee al eae)

Amphib. Reptile Conserv. 121 March 2019 | Volume 13 | Number 1 | e172

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

13(1) [General Section]: 122-142 (e173).

Distribution of the Neotropical water snakes

Hydrops caesurus, H. marti, and H. triangularis in

South America, with new records from Peru and Brazil

'*Rudolf von May, *Nelson Rufino de Albuquerque, *Henrique Bartolomeu Braz,

“Roy Santa-Cruz, ‘Emanuele Biggi, °Francesco Tomasinelli, and ‘Daniel L. Rabosky

'Museum of Zoology and Department of Ecology and Evolutionary Biology, University of Michigan, 2220 Biological Sciences Building, 1105 North

University Avenue, Ann Arbor, Michgan 48109, USA *Universidade Federal de Mato Grosso do Sul/Campus do Pantanal, Laboratorio de Zoologia,

Corumbd, Mato Grosso do Sul, BRAZIL *Laboratory of Ecology and Evolution, Butantan Institute, Av. Vital BRAZIL, 1500, SGo Paulo-SP, BRAZIL

‘Area de Herpetologia, Museo de Historia Natural de la Universidad Nacional de San Agustin (MUSA), Av. Alcides Carrion s/n, Arequipa, PERU

°International League of Conservation Photographers °Milan, ITALY

Keywords. Amazonia, Colubridae, Dipsadinae, Hydropsini, map, ecoregions

Citation: von May R, de Albuquerque NR, Braz HB, Santa-Cruz R, Biggi E, Tomasinelli F, Rabosky DL. 2019. Distribution of the Neotropical water

snakes Hydrops caesurus, H. marti, and H. triangularis in South America, with new records from Peru and Brazil. Amphibian & Reptile Conservation

13(1) [General Section]: 122-142 (e173).

4.0 International (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in any

medium, provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced, are

as follows: official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org.

Received: 2 February 2018; Accepted: 6 September 2018; Published: 18 April 2019

Neotropical water snakes in the genus Hydrops Wagler,

1830 (Serpentes, Colubridae, Dipsadinae, Hidropsin1)

are distributed across the northern two-thirds of South

America and have been recorded in 12 countries: Colom-

bia, Venezuela, Trinidad and Tobago, Guyana, Suriname,

French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay,

and Argentina (Albuquerque 2000; Scrocchi et al. 2005;

Albuquerque and De Lema 2008). Three species are

currently recognized: Hydrops caesurus Scrocchi, Fer-

reira, Giraudo, Avila and Motte, 2005, Hydrops martii

(Wagler 1824), and Hydrops triangularis (Wagler 1824).

The most recent geographical distribution maps for these

species have been provided by Scrocchi et al. 2005 (4.

caesurus), Entiauspe-Neto et al. 2017 (H. martii), and

Albuquerque and De Lema 2008 (#7. triangularis). Here,

we present an updated map and a list of individual geo-

referenced records for these species. Our database also

includes new records of H. triangularis from southern

Peru (extending the known geographic distribution of

this species) and northern Brazil, and a new record of H.

caesurus from Brazil.

We reviewed the literature, museum specimens, and

several publicly available biodiversity databases to

compile a list of geo-referenced locality records for the

three species of Hydrops (Appendix I). In cases where

a specimen could not be physically inspected, we com-

municated with researchers who confirmed the identity

and location of a particular record and/or the curators in

charge of specific herpetological collections. Addition-

ally, all coauthors carried out a critical review of the

literature, paying special attention to publications from

the country or region where they conduct their research.

Most records were supported by voucher specimens

listed in VertNet (http://vertnet.org) and/or deposited

in one of the following natural history museum collec-

tions: Colecéo Herpetologica Alphonse Richard Hoge,

Instituto Butantan (IBSP), Cole¢ao Herpetolégica das

Faculdades Integradas do Tapajés (LPHA), Colecao

Zoologica de Vertebrados da Universidade Federal de

Mato Grosso (UFMT), Instituto Nacional de Pesquisas

da Amazonia (INPA), Museo de Historia Natural de la

Universidad Nacional Mayor de San Marcos (MUSM),

Museo de Historia Natural de la Universidad Nacional

de San Agustin (MUSA), Museu de Ciéncias e Tecnolo-

gia, Pontificia Universidade Catolica do Rio Grande do

Sul (MCTPUCRS), Museu de Historia Natural Capao da

Imbuia (MHNCI), Museu de Zoologia da Universidade

de Sao Paulo (MZUSP), Museu Nacional, Universidade

Federal do Rio de Janeiro (MNRJ), Museu Paraense

Emilio Goeldi (MPEG), Universidade Federal do Acre

(UFAC), and University of Michigan Museum of Zool-

ogy (UMMZ). The dataset also includes 12 geo-refer-

enced records available in GBIF (https://www.gbif.org)

that we were able to verify; all other records from GBIF

Correspondence. ! rvonmay@umich.edu (corresponding author); * ne/son.rufino@ujfms. br; > h.braz@hotmail.com;

* chara53@hotmail.com; ° ebiggi@anura. it; ° ftomasinelli@gmail.com, ' drabosky@umich.edu

Amphib. Reptile Conserv.

April 2019 | Volume 13 | Number 1 | e173

von May et al.

were not included in our analysis either because accurate

voucher specimen data were not available or they lacked

geo-referenced data.

The database contains 337 geo-referenced records,

most of which were supported by voucher specimens de-

posited in a museum collection (Appendix I). We also

included one observation from southern Peru lacking a

voucher specimen, but in this case a high-resolution pho-

tograph was available. Altogether, the dataset includes 17

records of H. caesurus, 98 of H. martii, and 222 of H.

triangularis.

We used the R package maptools (Bivand and Lewin-

Koh 2014) to produce a map depicting the distribution of

the three species of Hydrops in South America. We also

consulted a layer depicting the Global Biomes according

to the World Wildlife Fund (WWF) classification, ob-

tained from the Terrestrial Ecoregions of the World da-

taset (WWE 2008), to determine the primary ecoregions

used by each species.

We report four individual records of H. triangularis

from Madre de Dios region, Peru. The first two individu-

als were found at Pampas del Heath National Sanctuary

on 10 November 2015. This site is located at the tran-

sition between lowland rainforest and tropical savanna

grasslands. Both individuals were collected in a flooded

area, at a site dominated by seasonally-flooded grassland

(12°56’42.25”S, 68°55’4.62”W; 212 m elev.) and that

had limited canopy cover. Both specimens were deposit-

ed at the Herpetological Collection at the Museo de His-

toria Natural de la Universidad Nacional de San Agustin

(MUSA), Arequipa, Peru. One was an adult female

(MUSA-4749, Fig. 1) with a total length of 774 mm, and

the other an adult male (MUSA-4750) with a total length

of 565 mm (male H. triangularis are sexually mature

at body sizes > 415 mm; Scartozzoni 2009). The third

individual of H. triangularis was found at Los Amigos

Biological Station on 26 July 2016. This site is primar-

ily covered by continuous, undisturbed lowland rainfor-

est and includes several aquatic and terrestrial habitats

typical to western Amazonia (von May et al. 2009).

The snake was encountered on a humid night following

heavy afternoon rainfall. This individual (Fig. 2) was

moving on the ground in a mature floodplain forest, and

had an estimated total length of ~700 mm. A GPS point

(12°34’07’S, 70°05’57°W, 250 m elev.) and photographs

were taken (photographic voucher PERU2016_ 5732),

but the specimen was not collected. A fourth individual

of H. triangularis was found at Los Amigos Biological

Station on 23 November 2017. This individual was cap-

tured in a funnel trap placed in a wetland area in a swamp

dominated by Mauritia flexuosa trees. This habitat is lo-

cally known as Aguajal and is one of the main forest

types at the site. This individual had a total length of 313

mm and was deposited in the herpetological collection

at the Museo de Historia Natural Universidad Nacional

Mayor de San Marcos (MUSM), Lima, Peru (MUSM-H

38992; field number RAB2664). These four specimens

Amphib. Reptile Conserv.

represent the first records of the genus Hydrops in Madre

de Dios region.

The four specimens reported here represent an exten-

sion of the known geographic range of H. triangularis,

this species was not included in the most up-to-date rep-

tile species lists available for Madre de Dios (Catenazzi

et al. 2013; Whitworth et al. 2016). The closest known

record of H. triangularis was based on a specimen col-

lected at Cachoeira Chapacura, Bolivia, and deposited in

the Colecao Herpetologica Alphonse Richard Hoge at In-

stituto Butantan (IBSP 41351), which was lost when the

institute’s collection was destroyed by fire (T. Guedes,

pers. comm.). Specifically, the records from Los Amigos

Biological Station extend the known geographic range

163 km to the southwest, and the records from Pampas

del Heath National Sanctuary extend the known geo-

graphic range 167 km to the south. It is worth noting

that these records of H. triangularis were obtained in a

region that has been the focus of numerous herpetologi-

cal inventories spanning several decades (Rodriguez and

Cadle 1990; Morales and McDiarmid 1996; Duellman

2005; von May and Donnelly 2009; von May et al. 2006;

von May et al. 2009, 2010; Catenazzi et al. 2013). This

finding demonstrates the need for further field research

even at sites that have been the focus of multi-year sur-

veys (e.g., Los Amigos Biological Station).

Fig. 1. Hydrops triangularis (MUSA 4749) collected at Pampas

del Heath National Sanctuary, Madre de Dios region, Peru.

Photo by Roy Santa Cruz.

Fig. 2. Hydrops triangularis (photographic voucher Nbr.

PERU2016_ 5732) recorded at Los Amigos Biological Station,

Madre de Dios, Peru. Photo by Emanuele Biggi.

April 2019 | Volume 13 | Number 1 | e173

New distributional records for Hydrops in Peru and Brazil

H. triangularis

(new records)

H. martii

H. caesurus

(new records)

Fig. 3. Distribution of Hydrops triangularis, Hydrops martii, and Hydrops caesurus in South America. Triangles indicate new

records.

Our database also includes new geo-referenced data

from specimens of H. caesurus (UFMT-R 8684) in the

State of Mato Grosso, Brazil, and the first record of H.

triangularis (MPEG 16697) in the State of Roraima,

Brazil. We also note that one record of H. triangularis

from Guyana was based on a voucher specimen de-

posited in the Muséum National d’Histoire Naturelle

(MNHN), Paris, France (MNHN 1996.4586). This speci-

men was subsequently identified as H. caesurus because

it matched the diagnostic features for this species (AI-

buquerque and De Lema 2008). However, given that it

was collected far from the previously known geographic

range of H. caesurus (Scrocchi et al. 2005), further stud-

ies including genetic data are needed to assess the status

of this population.

The updated map (Fig. 3) shows that H. martii and

H. triangularis have largely overlapping geographic

distributions, and are often found in syntopy - a pat-

tern suggested by previous research (Albuquerque and

Camargo 2004) - especially along the central portion of

the Amazon basin, from multiple sites around the Iquitos

region to sites located east and south of Belém, Brazil. In

contrast, H. caesurus has a separate geographic distribu-

tion outside of the Amazon basin. Furthermore, the dis-

tribution of Hydrops in South America is strongly linked

Amphib. Reptile Conserv.

to several major biomes. The distribution of H. martii

is restricted to tropical and subtropical moist broadleaf

forest. We also note that H. martii does not occur at sites

located west of the Andes in Ecuador and northern Peru,

nor in southeastern Peru, as was claimed previously (Al-

buquerque 2000). The distribution of H. triangularis is

primarily associated with tropical and subtropical moist

broadleaf forest, though it also occurs in tropical and

subtropical grasslands, savannas and shrublands (e.g., in

Venezuela and Bolivia). The distribution of H. caesurus

is strongly associated with flooded grasslands and savan-

nas, tropical and subtropical grasslands, and savannas

and shrublands (1.e., in Brazil, Paraguay, and Argentina).

Given that the geographic distribution of H. caesurus

does not overlap with that of the other two species, it

is likely that strong ecological and historical (e.g., river

basins) barriers drive the distributions of these species. It

is worth noting that one biome, tropical and subtropical

dry broadleaf forest, is present between the northernmost

localities of H. caesurus and the southernmost localities

of H. triangularis, and that neither species has been re-

corded in this biome.

In addition to the genus Hydrops, two other genera

of water snakes in the tribe Hydropsini - Helicops and

Pseudoeryx - are distributed in South America. Recent

April 2019 | Volume 13 | Number 1 | e173

von May et al.

studies focusing of the reproductive mode of Neotropical

water snakes have shown that all three species of Hy-

drops, in addition to Pseudoeryx plicatilis and two spe-

cies Helicops (H. gomesi and H. hagmanni) are ovipa-

rous (Albuquerque and Camargo 2004; Braz et al. 2016).

In contrast, most other species of Helicops are viviparous

and only one (H. angulatus) exhibits both reproductive

modes (Braz et al. 2016). Documented records of the re-

productive mode of H. angulatus indicate that vivipar-

ity is more prevalent in western Amazonian populations

(e.g., Madre de Dios region, Peru), while oviparity is

prevalent in the eastern portion of the range of this spe-

cies (Braz et al. 2016).

Acknowledgements.—We thank Ana Lucia da Costa

Prudente for kindly providing geo-referenced data avail-

able at the Museu Paraense Emilio Goeldi, Brazil, and

Thais Guedes for providing a geo-referenced data for a

specimen from Bolivia. We are thankful to Paulo Pas-

sos (MNRJ), Richard Vogt (INPA), Francisco Franco

(IBSP), Hussam Zaher (MZUSP), Hipocrates Chalkidis

(LPHA), Julio Moura-Leite (MHNCI), Glaucia Funk-

Pontes (MCTPUCRS), Marcos Carvalho (UFMT), Moi-

sés Souza (UFAC), César Aguilar (MUSM), Evaristo

Lopez (MUSA), and Greg Schneider (UMMZ) for al-

lowing access to specimens under their care. We thank

the Amazon Conservation Association and the staff at

Los Amigos Biological Station for facilitating our work

at the station. This research was supported with grants

from the National Science Foundation (Postdoctoral Re-

search Fellowship DBI-1103087), the Amazon Conser-

vation Association, and the National Geographic Society

(Grant # 9191-12) to Rudolf von May, by a fellowship

from the David and Lucile Packard Foundation to Daniel

L. Rabosky, and by a doctoral fellowship provided by the

Fundacao de Amparo a Pesquisa do Estado de Sao Paulo

(FAPESP) to Henrique B. Braz (2009/54478-3). Roy

Santa Cruz thanks the Asociacion para la Investigacion

y Desarrollo Integral (AIDER), for supporting the Rapid

Assessment of the biodiversity of Pampas of Heath and

Bahuaja-Sonene National Park; Roberto Gutiérrez, for

coordinating expeditions to Pampas del Heath; Deyvis

C. Huaman, for obtaining collection and research permits

(Resolucion del Jefe del Parque Nacional Bahuaja Sonene

N° 012-2015-SERNANP-DGANP-PNBS/J); and Evaristo

Lopez Tejeda, for providing access to the specimen collec-

tion at Museo de Historia Natural de la Universidad de San

Agustin de Arequipa (MUSA). Additional research and

collecting permits were issued by the Direccién General

Forestal y de Fauna Silvestre (DGFFS) and the Servicio

Nacional Forestal y de Fauna Silvestre (SERFOR), Peru

(permits 292-2014-AG-DGFFS-DGEFFS, R.D.G. No

029-2016-SERFOR-DGGSPFFS, R.D.G. 405-2016-SER-

FOR-DGGSPFFS). We thank Vanda Lucia Ferreira and

two anonymous reviewers for providing constructive

comments on the manuscript.

Amphib. Reptile Conserv.

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Rudolf von May is a Postdoctoral Research Fellow at the Museum of Zoology and the Department

of Ecology and Evolutionary Biology at the University of Michigan. His current research seeks to

understand how biological communities are structured across habitats and elevations, with a special

focus on amphibians and reptiles living in the Andes-Amazon region.

Nelson Rufino de Albuquerque is an Associate Professor at the Universidade Federal de Mato Grosso

do Sul (UFMS), in Corumba, Brazil. He studies systematics, taxonomy, anatomy, and natural history

of snakes in the Neotropics. One of his current projects focuses on the taxonomy and systematics of

the Green Parrotsnake, Leptophis ahaetulla complex, a morphologically diverse assemblage of species

widely distributed in Latin America. He joined the UFMS in 2009, where he teaches Chordates, Animal

Physiology, and Comparative Vertebrate Anatomy, and advises graduate and undergraduate students.

Henrique Bartolomeu Braz is a Research Associate at the Laboratory of Ecology and Evolution,

Butantan Institute, Sao Paulo, Brazil. He has a broad interest in natural history, reproductive biology,

and evolution of squamate reptiles. His current research projects include the evolution of viviparity

(live-birth) in squamates, reproductive phenology, nest-site selection, and developmental plasticity in

April 2019 | Volume 13 | Number 1 | e173

New distributional records for Hydrops in Peru and Brazil

Roy Santa-Cruz is a Research Associate at Area de Herpetologia del Museo de Historia Natural

(MUSA) de la Universidad Nacional de San Agustin de Arequipa, Peru. His current research interests

include taxonomy, natural history, and conservation of amphibian and reptiles. He currently coordinates

several research projects focusing on threatened species of Andean frogs.

Emanuele Biggi is an Italian naturalist with a Ph.D. in Environmental Sciences. He is an Associate

Fellow of the International League of Conservation Photographers. He is mainly focused on the smaller

creatures, and conservation of nature and science-at-work in photography. He is the author and curator

of scientific expositions in which he tries to bring nature to people, to raise the awareness about all the

beauty and issues of the natural world. Emanuele is also winner and finalist of various international

photography prizes and speaker of international photography symposia and festivals. Website: www.

anura. it

Francesco Tomasinelli has a degree in Environmental Science and works as freelance biologist,

journalist, and photographer. As an ecologist, he mostly does surveys and assessments on local fauna

and flora in agricultural and semi-urbanized areas. As a photojournalist, he has joined several scientific

expeditions in the tropics and has covered the activity of conservation and environmental programs

in Italy and abroad. He specializes in unusual stories about nature, and has a growing collection of

geographic, travel and science images. He writes books and creates exhibits for museums, involving

live invertebrates, photos and activities for visitors. Website: www. isopoda.net

Daniel L. Rabosky is an Assistant Professor and Curator of Herpetology in Museum of Zoology at

the University of Michigan. He studies macroevolution, global diversity gradients, and the ecology

and evolution of squamate reptile communities. He conducts fieldwork on reptiles and amphibians in

Australia and the Neotropics.

Appendix 1. Geo-referenced data for individual species records included in this study.

caesurus Corrientes Couturier, 1984 | Couturier, 1984

foo | “| eaters” | [LONE | a

caesurus Miguel, Corrientes 2005

Hydrops Puerto Carambola, Scrocchi et al., Scrocchi et al.,

caesurus Departamento San Miguel, 2004 2005

Provincia de Corrientes

foam | “| Memwme | ve | ee | | | ae

caesurus 2006

Hydrops Puerto Tala, Isla Apipé -27.55000 -56.80000 UNNEC 11436 | Etchapare et Etchapare et al.,

caesurus Grande, Provincia de al., 2012 2012

Corrientes

Hydrops AR Isla Apipé Grande, -27.50 -56.90 Zaracho et al.,

caesurus Corrientes 2014

Hydrops Complejo Isla Yacyreta, -27.48278 -56.73583 MNHNP 4951 | Scrocchi et al., Scrocchi et al.,

caesurus Departamento Itapua 2005 2005

Hydrops Canal de los Jesuitas, Isla -27.46670 -56.41670 MNHNP 06698 | Scrocchi et al., Scrocchi et al.,

caesurus Paloma, Departamento 2005 2005

Itapua

Hydrops Isla Yacyreta and -27.43000 -56.77000 MNHNP 06697 | Scrocchi et al., Scrocchi et al.,

caesurus surroundings, 2005

Departamento Itapua

Amphib. Reptile Conserv. 128 April 2019 | Volume 13 | Number 1 | e173

von May et al.

Paraguay River; 14 km -25.53944 -57.17083 MNHNP 06462 | Scrocchi et al., Scrocchi et al.,

S from Puerto Rosario, 2005

Departamento Presidente

Hayes

Rio Paraguay ca. 14 km by -24.53944 -57.17083 MNHNP 06462 | P. Buongermini Herp Review

river from Puerto Rosario & T. Waller 1998

Hydrops

caesurus

Hydrops

caesurus

Hydrops B Miranda, Mato Grosso -20.23333 -56.36667 CEUCH 3061 | Scrocchi et al., Scrocchi et al.,

caesurus do Sul 2005 2005

Hydrops B Ladario, Mato Grosso do -19.00500 -57.53348 CEUCH 27 Scrocchi et al., Scrocchi et al.,

caesurus Sul 2005 2005

B > Ps

Hydrops Aquidauana Municipality, -20.46667 -55.80000 IBSP 29171 Nelson R. de Herp Review

caesurus Mato Gross do Sul Albuquerque 2001

Hydrops lagoa Negra, Ladario, Mato | -18.97083 -57.56250 CEUCH 208 Scrocchi et al. Scrocchi et al.

caesurus Grosso do Sul 2005 2005

Hydrops BR Fazenda Acurizal, Serra -17.83083 -57.55167 UFMT-R 1188 | Scrocchi et al., Scrocchi et al.,

caesurus do Amolar, Mato Grosso 2005 2005

do Sul

Hydrops Poconé, Mato Grosso - 16.7845 -56.9485 UFMT-R 8684 R.A. This study

caesurus Kawashita-

Ribeiro

Hydrops Senador Guiomard, Acre -10.151 -67.736 UFAC 294 Henrique Braz This study

martit

Hydrops Roaboya (Rio Ucayal1), -7.78330 -74.91660 Roze (1957)

martii Ucayali

Hydrops PE Roaboya, Rio Ucayali, -7.760556 -74.9275 AMNH 53086 Roze, 1957b

martii Loreto

Hydrops PE Cashiboya, Rio Ucayali, -7.658321 -74.928645 AMNH 53130 Roze, 1957b

martii Loreto

Hydrops BR (Rio Purus), Amazonas -6.83330 -66.11660 Albuquerque

martit (2000)

Hydrops PE Pampa Hermosa (Rio -6.11660 -76.26660 Roze (1957)

martii Cushabatay) , Loreto

Hydrops PE Pampa Hermosa, Rio -5.766667 -74.931389 | AMNH 55429 Roze 1957b;

martii Cuchabatay, Loreto Albuquerque

2000

Hydrops PE Challavitas, Alto -5.45 -76.8 BMNH 1946 Roze, 1957b

martii Amazonas, Loreto

Hydrops BR Fazenda Sao Gabriel, -5.36889 -49.11778 MPEG 24073 | [no agent data] na

martii Maraba, PA

Hydrops PE Monte Carmelo, Requena, -5.07000 -73.91000 AMNH 55492 Nelson R. de Albuquerque

martii Loreto, PE Albuquerque 2000

Hydrops Requena, Uresti, Requena, -5.068389 -73.861994 AMNH Roze 1957b;

martii Loreto R55494 Albuquerque

2000

martii

Hydrops BR Base Operacional Gedlogo -4.885369 -65.349997 MPEG 22225 Prudente et al.,

martii Pedro de Moura, Provincia 2010

Petrolifera de Urucu,

Igarapé Tartaruga, Coari,

Amazonas

Hydrops BR Pimental, Itaituba, PA -4.57694 -56.26083 MPEG 24940 | [no agent data] na

martii

Hydrops BR Barreirinha, Rio Tapajos, -4.416667 -56.216667 MZUSP 5136 | Henrique Braz This study

mattii proximo a Sao Luis do

Tapajos, Para

Amphib. Reptile Conserv. 129 April 2019 | Volume 13 | Number 1 | e173

New distributional records for Hydrops in Peru and Brazil

Hydrops BR Rio Madeira, Borba, -4.388 -59.594 MNRJ 2985 Henrique Braz This study

martii Amazonas

Hydrops Rio Itacoai, 30 km do Rio -4.383 -70.031 MNRJ 3013 Henrique Braz This study

martii Javari, Benjamim Constant,

Amazonas

Hydrops Leticia, Amazonas -4.205428 -69.932808 MCTPUCRS | Henrique Braz This study

martii 14100

Hydrops Hoogmoed &

martii Gruber, 1983;

Albuquerque,

2000; Franzen &

Glaw, 2007

Hydrops Reserva de Waldez et al.

martii Desenvolvimento (2013)

Sustentavel Piagacgu-Purus

(Rio Purus) , Amazonas

Albuquerque,

2000

Hydrops

martii

Hydrops

martii

Hydrops

martii

Hydrops Moropon, Iquitos, Maynas, -3.766667 -73.416667 TCWC 45572 Albuquerque,

martii Loreto 2000

Roze 1957b;

Albuquerque

Rio Itaya, Iquitos, Maynas, | -3.7658333 | -73.2316667 | AMNH 55299

Loreto

2000

Hydrops Iquitos, Loreto, PE -3.75000 -73.25000 TCWC 45571 Nelson R. de Albuquerque

martii Albuquerque 2000

Hydrops Iquitos, Maynas, Loreto -3.75 -73.25 AMNH 53408 Roze, 1957b

martit

Hydrops Nanay, Loreto, PE -3.74068 -73.24299 AMNH 52031 Nelson R. de Albuquerque

martii Albuquerque 2000

Hydrops Nanay, Loreto -3.66660 -73.23330 Roze (1957)

martit

Hydrops Mazan, Maynas, Loreto -3.496494 -73.089961 Monge &

martii Cabanillas, 2009

Hydrops Sao Paulo de Oliven¢a -3.46660 -68.93330 Roze (1957)

martii (Sao Joao, Rio SolimG6es),

Amazonas

Hydrops S40 Joao, Rio Solimées, -3.39836 -68.91610 AMNH 25194 Roze, 1957b

martii Sao Paulo de Olivenga,

Amazonas

Hydrops BR Tefé, Amazonas -3.38330 -64.70000 Roze (1957)

martil

Hydrops BR Boca de Tefé, confluéncia -3.37644 -64.561592 MP 461 Roze, 1957b

martii dos rios Tefé e Solim6es,

Tefé, Amazonas

Amphib. Reptile Conserv. 130 April 2019 | Volume 13 | Number 1 | e173

von May et al.

Hydrops Pévas, Ampiyacu River, -3,32257 -71.86264 CAS-SU 8724

marti E. Peru

Hydrops Pebas, Mariscal Ramon -3,322222 -71.816667 CASSU 8724

martii Castilla, Loreto

Nelson R. de Albuquerque

; Albuquerque 2000

Hydrops Rio Xingu, 6h de barco de -3.203 -52.206 IBSP 56498

martii Altamira, Para

Albuquerque,

2000

; Albuquerque,

2000

Hydrops Manaus, Amazonas -3.13330 -60.01660 Roze (1957)

martil

Hydrops Rio Madeira, Manaus, -3.102 -60.025 INPA 12031 Henrique Braz This study

martii Amazonas

Hydrops BR Lago Tapaiuna, margem -3.10194 -60.02500 IBSP 32832 Nelson R. de Albuquerque

martii esquerda do rio Preto da Albuquerque 2000

Eva, Manaus, AM, Brasil

Hydrops Manaus, AM -3.10000 -60.01667 AMNH 36161 Nelson R. de Albuquerque

martii Albuquerque 2000

Hydrops Mezza (Boca de Tefé) , Amazonas | -3.01660 -64.80000 Roze (1957)

martil

Albuquerque &

Camargo, 2004;

Frota et al., 2005

CisnerosHeredia,

2005

Hydrops BR UHE CuruaUna, Santarém, -2.812492 -54.298675 MCTPUCRS

martii Para T916

Hydrops

martii

Mashumarentsa, divisa -2.75 -77.216667 FHGO 1266

com Peru, 600 m de

altitude, MoronaSantiago

R Estacao Ecologica -2.75 -60.75

Anavilhanas, Arquipélago

Anavilhanas, Amazonas

R Novo Airao, Amazonas -2.621 -60.944 MM J26

martit

Hydrops R Lago Amana, Rio Japura -2.59674 -64.66291 MPEG 16775

martii Maraa, AM

Hydrops R Santa Luzia do Parua, BR- -2.59278 -45.74667 MPEG 13662

316, MA

Silveira &

Magnusson, 1999

Henrique Braz This study

[no agent data]

Albuquerque

(2000)

Henrique Braz This study

Nelson R. de Albuquerque

Albuquerque 2000

Henrique Braz This study

Nelson R. de Albuquerque

Albuquerque 2000

Hydrops

martii

Hydrops

matrtii

Hydrops

martii

Hydrops

martii

Ipixuna do Para, PA -2.55778 -47 49389 MPEG 21344

Fazenda Santa Monica, -2. 443 -54.708 LPHA 1301

martii Santarém, Para

Hydrops BR U.HLE. Curua-Una, -2.44278 -54.70778 MCP 7916

martii Santarém, PA

Hydrops ToméAcu, Para -2.419 -48.152 MHNCI 8306

B Ipitinga, Estrada Moju- -2.41889 -48.15194 MPEG 12610

Acara, Tomé-Acu, PA

B Curumucuri, Juruti, PA -2.15194 -56.09194 MPEG 22545

martii

Hydrops B Km 11 da Estrada do -2.02548 -47.72209 MPEG 15500

martii Acara, PA

| 7

Hydrops

martii

Hydrops

martii

Hydrops

martii

Hydrops km 16 da estrada do Acara -2.02361 -47.68750 MPEG 15500 | [no agent data]

Monte Alegre, Para -2.008 -54.069 LPHA 2840 Henrique Braz This study

martil

Hydrops BR ECFPn/MPEG/ Floresta - 1.90667 -51.38222 MPEG 21871 | [no agent data] na

martii Nacional de Caxiuaa,

Itaperu, Melgaco, PA

Amphib. Reptile Conserv. 131 April 2019 | Volume 13 | Number 1 | e173

Hydrops

New distributional records for Hydrops in Peru and Brazil

Rio Uba. Povoac¢ao do -1.8839 -48.7689 MPEG 13297 | Henrique Braz This study

Luso. Km 36 da estrada

Moju-Acara, Moju, Para

Hydrops Santa Luzia, Capitéo Poco, - 1.74500 -47.06500 MPEG 4173 Nelson R. de Albuquerque

martii PA Albuquerque 2000

Hydrops Santa Luzia, Capitéo Poco, -1.745 -47.065 MPEG 6042 Henrique Braz This study

martii Para

Hydrops Fea) Sao Pedro, Capitio Poco, | -1.6189670 | -47.1678830 | MPEG 10471 This study

martii Para

Hydrops

martii

Hydrops

martii

Hydrops

martii

Hydrops

Hydrops Limao Grande, Ourém, PA | -1.51528 -47.00211 | MPEG 4241 | [no agent data]

Belém, PA -1.43730 -48.47060 | MPEG 18893 | [no agent data]

Santa Maria, Rio Santa -1.398125 -74.644075 TCWC 38227 Dixon & Soini,

Maria, Loreto 1977

Macapazinho, Castanhal, -1.389119 -47.984161 MPEG 11800 Albuquerque

Para 2000

matrtii

Hydrops

martii

Hydrops Santa Maria, Rio Santa - 1.38333 -74.65000 Nelson R. de

martii Maria, Loreto Albuquerque

Hydrops BR Pratinha, Estrada de - 1.36083 -48.24500 MPEG 8618 Nelson R. de Albuquerque

martii Genipauba, Benevides, PA Albuquerque 2000

Hydrops

martii

| Boa Vista, Castanhal, Para | -1.360117 | -47.985536 | MPEG 2701 This study

€ Puerto Charapa, La -1.338264 -69.558428 PérezSantos &

martii Pedrera, Amazonas Moreno, 1988

Hydrops Curupaiti, Viseu, Para -1.298289 -46.322239 MPEG 12327 Albuquerque

martii 2000

Hydrops Castanhal, PA - 1.29694 -47.92194 MPEG 11800 Nelson R. de Albuquerque

martii Albuquerque 2000

Hydrops [Racial Bom Jesus, Braganca, PA | -1.26472 -46.68944 | MPEG 2228 | [no agent data]

martil

Hydrops

Albuquerque

2000

This study

Hydrops Viseu, PA -1.19694 -46.14000 MPEG 12327 Nelson R. de

marttii Albuquerque

Hydrops BR Lago Jacaré, Rio -1.188611 -56.67083 MZUSP 3827 | Henrique Braz

martii Trombetas Reserva

Bioldgica do Rio

Trombetas, Para

Hydrops SolimGes, Tefé, AM, Brasil -1.10444 -47.51194 IBSP 15086 Nelson R. de

marttii Albuquerque

Hydrops Bela Vista, Viseu, Para -1.0700 -46.7900 MPEG 13192 | Henrique Braz

martit

Hydrops Ilha de Marajo, Para -0.966667 -49_566667

martit

Albuquerque

2000

This study

Hoge & Nina,

1969

Roze (1957)

matrtii

Hydrops Ilha de Marajo, Para -0.85000 -49.40000

Auca Via Cononaco, km -0.708403 -76.887861 DHMECN 139

martii 135, Orellana

Hydrops EC Estacion de Biodiversidad -0.616667 -76.166667 DFCHUSFQ

martii Tiputini, 215 m de altitude, OH15

Orellana

Hydrops B Santa Isabel do Rio Negro, -0.414 -65.019 INPA 12624 Henrique Braz

martii Amazonas

Hydrops B Rio Preto, Afua, Ilha do -0.15694 -50.38694 MPEG 25324 | [no agent data]

martii Marajo, PA

Amphib. Reptile Conserv. 132 April 2019 | Volume 13 | Number 1 | e173

Hydrops CisnerosHeredia,

2005

CisnerosHeredia,

2005

This study

von May et al.

Cuyabeno, Sucumbios -0.1167 -75.8333

Macapa, Amapa 0.03330 -51.06660

Hydrops Rio Cairary, proximo 1.073611 -69.845 AMNH 4459

martii a Cachoeira Jurupary,

proximo fronteira entre

BRACOL, Vaupés

Hydrops Rio Branco, Igarapé do 1.919069 -61.001122 MPEG 16696 Albuquerque,

martii BotaPanela, perto da 2000

Cachoeira de Bem Querer,

Cararau, margem do rio,

Caracarai, Roraima

DHMECN 138

CisnerosHeredia,

2005

Entiauspe-Neto et

al. (2017)

Roze 1957b;

Albuquerque

2000

(Rio Cairary, near Jurupary 1.93330 -68.25000 Roze (1957)

Waterfall) , Guainia

Hydrops conjun¢ao do Canal del 2.119158 -66.462072 Rivas et al., 2012

mattii Casiquiare e Rio Siapa,

Amazonas

= pa

BurroBurro, 83 m, 4.731 -58.850667

PotaroSiparuni

km 23 da estrada de -3.35417 -64.71140

Maracana, PA

Hydrops Comunidad Oromomo, -15.833 -66.367

triangularis Moxos, Beni

confluéncia dos rios Ichoa -15.733 -65.25

e Aguas Claras, Moxos,

Beni

Ibiato, Provincia de -14.82998 -64.44855

triangularis Cercado, Departamento de

Cochabamba

El Refugio, Provincia de -14.76700 -61.03300

triangularis San Ignacio de Velasco,

Departamento de Santa

Cruz

Hydrops Estacion Biologica Beni, -14.38000

triangularis Provincia de Yacuma,

Departamento del Beni

Hydrops Cachuela Chapacura, -14.283

triangularis margem do Rio Blanco,

Iténez, Beni

Hydrops Lago Rogoagua -13.03500 -65.942222

triangularis (=Roguaguado),

Departamento del Beni

Hydrops PE Pampas del Heath National | -12.94507 -68.91795 MUSA - 4749 | Roy Santa Cruz This study

triangularis Sanctuary, Madre de Dios

Hydrops PE Pampas del Heath National | -12.94507 -68.91795 MUSA - 4750 | Roy Santa Cruz This study

triangularis Sanctuary, Madre de Dios

Hydrops Lago Rogoaguado, -12.867 -65.717 AMNH 22449 Roze, 1957b;

triangularis Yacuma, Beni Albuquerque

& Lema, 2008;

Scrocchi et al.,

2005

Hydrops Curichi, Trinidad -12.61667 -63.51667 MNKR 3698 Nelson R. de Albuquerque &

triangularis Albuquerque Lema, 2008

Amphib. Reptile Conserv. 133 April 2019 | Volume 13 | Number 1 | e173

FA 184-185

MZUSP 10121 This study

Donnelly et al.,

2005

MPEG 2110 [no agent data]

CBF 1010 Albuquerque &

Lema, 2008

CBF 1005 Albuquerque &

Lema, 2008

CBF 1038 Nelson R. de Albuquerque &

Albuquerque Lema, 2008

MNKR 2588 Nelson R. de Albuquerque &

Albuquerque Lema, 2008

CBF 393 Nelson R. de Albuquerque &

Albuquerque Lema, 2008

IBSP 41351 Albuquerque &

Lema, 2008

AMNH 22449 Nelson R. de Albuquerque &

Albuquerque Lema, 2008

New distributional records for Hydrops in Peru and Brazil

Hydrops Estacion Biologica Los -12.56861 -70.09917 Emanuele This study

triangularis Amigos Biggi &

Francesco

Tomasinelli

Estacion Biologica Los -12.55970 -70.11030 Rudolf von This study

Amigos May & Daniel

Rabosky

Rio Guaporé, Costa -12.445 -64.227 MCT-PUCRS Albuquerque &

Marques, Rondénia 6300 Lema, 2008

BR Praia Alta, Costa Marques, -12.44500 -64.227222 MCP 6300 Nelson R. de Albuquerque &

RO Albuquerque Lema, 2008

Hydrops Rio Itenez, Beni -11.891 -65.011 AMNH 101863 Albuquerque &

triangularis

Lema, 2008

Cachoeira de Tchapa Cura -11.43365 -69.13108 IBSP 41351 Thais Guedes Thais Guedes,

& Fausto personal

Barbo communication

Rio Mamoré, -10.82667 -65.356667 | AMNH 101862} Nelson R. de Albuquerque &

triangularis Guayaramerin, Albuquerque Lema, 2008

Departamento del Beni

Guayaramerin, Rio -10.8 -65.383 UMMZ 56896

triangularis Mamoré, Antonio Vaca

Diez, Beni

Hydrops Guajara-Mirim, Rond6énia -10.783 -65.339

triangularis

Roze, 1957b;

Albuquerque

& Lema, 2008;

Scrocchi et al.,

2005

Bernarde et al.,

2012

Hydrops U.H.E. Lajeado, Porto -10.70778 -48.416944 IBSP 66641 Nelson R. de Albuquerque &

triangularis Nacional, TO, Brasil Albuquerque Lema, 2008

Hydrops UHE Luis Eduardo -10.167 -48 333 IBSP 65686 Henrique Braz This study

triangularis Magalhaes, Palmas,

Tocantins

Hydrops BR Rio Branco, Acre -9.975 -67.8] UFAC 162 Henrique Braz This study

triangularis

Hydrops UHE Luis Eduardo -9.751 -48.358 IBSP 65822 Henrique Braz This study

triangularis Magalhaes, Lajeado,

Tocantins

Hydrops BR UHE Luis E Magalhaes, -9.75083 -48.35778 IBSP 65362 Nelson R. de Albuquerque &

triangularis Lajeado, TO Albuquerque Lema, 2008

Hydrops Rio Pacaya, Rio Ucayali, -9.627 -74.133 BMNH Roze, 1957b;

triangularis Loreto O13 F285 Albuquerque &

Lema, 2008

UHE Jirau, Rond6nia -9.33] -64.734 NATURAE,

2010a, b

UHE de Jirau, Jaci-Parana, -9.26444 -64.64194 MPEG 23968 | [no agent data] na

Porto Velho, RO

Hydrops Santa Barbara, Rond6énia -9.167 -63.067 MZUSP 8780 | Henrique Braz This study

triangularis

Hydrops UHE Santo Antonio, Porto -8.806 -63.937 Marcal et al.,

triangularis Velho, Rond6énia 2011

Hydrops Concei¢ao do Araguaia, PA -8.25778 -49.26500 IBSP 24038 Nelson R. de Albuquerque &

triangularis Albuquerque Lema, 2008

Hydrops Urucuiuna, Ribeiro -7.558 -45.242 MHNCI 2566 Yuki, 1997

triangularis Gongalves, Piaui

Hydrops Humaita, Amazonas -7.506 -63.021 MNRJ 19353 | Henrique Braz This study

triangularis

Teles Pires River, Mato -7.35000 -58.05000 IBSP 30868 Nelson R. de Herp Review

Grosso Albuquerque 2001

Amphib. Reptile Conserv. 134 April 2019 | Volume 13 | Number 1 | e173

von May et al.

Hydrops PE Rean Rean, Suhaya, -7.334 -75.204 AMNH 53579 Albuquerque &

triangularis Ucayali, Loreto Camargo, 2004

Hydrops BR Rio Purus, Labrea, AM -7.27000 -64.78000 RMNH 27784 Nelson R. de Albuquerque &

triangularis Albuquerque Lema, 2008

Hydrops ca. 170 m, Carolina, -7.196 -47.422 aT 2007

triangularis Maranhao

Hydrops Rio Madeira - Col. -6.995 -62.681 IBSP 15034 Henrique Braz Bestia study

triangularis Harold Sioli, Trés Casas,

Amazonas

Hydrops Projeto Cristalino, Posto -6.091 -49 54] MPEG 20746 | Henrique Braz This study

triangularis de coleta, Alojamento,

Curionopolis, Para

Hydrops Projeto Cristalino, Lagoa -6.09083 -49 54083 MPEG 20742 | [no agent cl

triangularis do Plat6, Curiondpolis, PA

Hydrops Rio Araguaia, Porto Jarbas -5.70667 -48.17944 MPEG 12752 Nelson R. de a &

triangularis Passarinho, Rodovia Albuquerque Lema, 2008

Transamazonica, Palestina

do Para, PA

Hydrops Barra do Corda, Maranhao -5.533 -45.267 MZUSP 3139 Hoge & Nina,

triangularis 1969

Hydrops Fazenda Sao Gabriel, -5.36889 -49.11778 MPEG 24104 | [no agent data]

triangularis Maraba, PA

Hydrops km 11 da PA-222 -5.29440 -49.07533 MPEG 9483 Nelson R. de Albuquerque &

triangularis Albuquerque Lema, 2008

Hydrops Fazenda Santa Maria -5.09417 -42.83670 IBSP 44149 Nelson R. de Albuquerque &

triangularis - km 602 - BR 316 Albuquerque Lema, 2008

- rodovia Sao Luiz a

Terezinha, Timon, MA

Hydrops Monte Carmelo, -5.07000 -73.91000 AMNH 55934 Nelson R. de Albuquerque &

triangularis Departamento de Loreto Albuquerque Lema, 2008

Hydrops Base Operacional Gedlogo -4.885 -65.35 MPEG 22226 | Henrique Braz This study

triangularis Pedro de Moura, Provincia

Petrolifera de Urucu, Coari,

Amazonas

Hydrops Pimental, Itaituba, PA -4.57694 -56.26083 MPEG 24942 | [no agent ol | ae

triangularis

Hydrops Itinga, Km 337 Belém- -4.45 -47.526 MPEG 1121 Henrique Braz This eile

triangularis Brasilia, Itinga do

Maranhao, Maranhao

Hydrops BR Sitio Bela Vista, PA-222, -4.43806 -47.53917 MPEG 15532 Nelson R. de Albuquerque &

triangularis Dom Eliseu, PA Albuquerque Lema, 2008

Hydrops BR Sitio Bela Vista, PA-222, -4.285 -47.505 MPEG 12135 | Henrique Braz This study

triangularis Dom Eliseu, Para

Hydrops BR Itaituba, Para -4.276 -55.984 LPHA 1345 Henrique Braz This study

triangularis

Hydrops CO Leticia, Amazonas -4.205 -69.933 MCT-PUCRS | Henrique Braz This study

triangularis 14099

Hydrops BR Boa Vista ???, Maranhao -4.017 -43.283 MZUSP 1298 Hoge & Nina,

triangularis 1969

Hydrops Igarapé Manjuru, AMNH 114261 Albuquerque &

triangularis Amazonas Lema, 2008

Hydrops Moropon, Loreto -3.88673 -73.73534 Nelson R. de

triangularis Albuquerque

Hydrops Mishana (=Minchana), -3.88080 -73.49170 TCWC 39095 Nelson R. de Albuquerque &

triangularis Departamento de Loreto Albuquerque Lema, 2008

Hydrops Codajas, AM -3.83694 -62.05694 BMNH Nelson R. de Albuquerque &

triangularis 1965.1325 Albuquerque Lema, 2008

Amphib. Reptile Conserv. 135 April 2019 | Volume 13 | Number 1 | e173

New distributional records for Hydrops in Peru and Brazil

Hydrops Mishana, Rio Nanay, -3.8 -73.533 TCWC 39096 Albuquerque &

triangularis Maynas, Loreto Lema, 2008

Hydrops Cachoeira do Espelho, Rio -3.8

triangularis Xingu, Altamira, Para

Hydrops PE Monte Carmelo, Maynas, -3.796 -73.046 AMNH 52354 Scrocchi et

triangularis Loreto al., 2005;

Albuquerque &

Lema, 2008

Hydrops Iquitos, Maynas, Loreto -3.75 -73.25 AMNH 52017 Roze, 1957b;

Albuquerque &

Lema, 2008

-3.74810 -73.24720 | MCZ R-56054 | Harvey Bassler GBIF

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

Alto Alegre do Pindaré, -3.70089 -44.91761 MPEG 25454 | [no agent data] na

Santa Luzia do Parua, BR- -3.60778 -45.34278 MPEG 11187 | [no agent data] na

triangularis 316, MA

Hydrops BR NE de Santa Inés, SW. -3.55556 -44.92028 MPEG 21278 | [no agent data] na

triangularis de Vitoria do Mearim,

Povoado de Tirirical, Sao

Luis, MA

Hydrops Gancho do Arari, BR-222 -3.517 -44.767 MPEG 13484 | Henrique Braz This study

triangularis entre Miranda e Arari,

Arari, Maranhao

Hydrops Arari, MA -3.46667 -44.78330 MNRJ 4497 Hussam Zaher Herp Review

& Ulisses 1996

Caramaschi

Hydrops BR Gancho do Arari, BR-222 -3.45389 -44.78000 MPEG 14619 Nelson R. de Albuquerque &

triangularis entre Miranda e Arari, MA Albuquerque Lema, 2008

Hydrops B Ega, lago Tefé, confluéncia -3.376 -64.562 ZSMH 1846/0 Roze, 1957b;

triangularis com rio Solimées, Tefé, Hoogmoed &

Amazonas Gruber, 1983

Hydrops BR Tefé, Amazonas -3.354 -64.711 MZUSP 8375 | Henrique Braz This study

triangularis

Hydrops BR Pedras, margem do Rio -3.349 -58.3 IBSP 32831 Henrique Braz This study

triangularis Preto da Eva, Manaus,

Amazonas

Hydrops BR Fazenda Santo Amaro, -3.20778 -43.40389 MPEG 20558 | [no agent data] na

triangularis Urbano Santos, MA

Hydrops BR Manaus, AM -3.10000 -60.01667 MZUSP 7679 Nelson R. de Albuquerque &

triangularis Albuquerque Lema, 2008

Hydrops BR Sao José, Jacaré, Rio -2.933 -66.133 MZUSP 5478 Albuquerque &

triangularis Solim6es, Amazonas Lema, 2008

Hydrops BR Estagao Ecologica -2.75 -60.75 Silveira &

triangularis Anavilhanas, Arquipélago Magnusson, 1999

Anavilhanas, Amazonas

Hydrops Reserva do Alto do Rio -2.616 -46.511 MZUSP 4220 Albuquerque &

triangularis Guama, Aldeia Canindé, Lema, 2008

Rio Gurupi, Para

Hydrops BR Pindaré-Mirim, Puraqueu, -2.59278 -45.74667 MPEG 14703 | [no agent data] na

triangularis BR-222, MA

Hydrops BR Vila Nova, Rio Xingu, -2.591 -51.954 IBSP 14939 Henrique Braz This study

triangularis Col. Bach, Senador José

Porfirio, Para

Amphib. Reptile Conserv. 136 April 2019 | Volume 13 | Number 1 | e173

Hydrops Compagnie Creek -3.73333 -73.25000 RMNH 13609 Nelson R. de Albuquerque &

triangularis Brokopondo District Albuquerque Lema, 2008

triangularis

von May et al.

Povoado de Ponta do -2.579 -42.745

Mangue, Barreirinhas,

Maranhao

Hydrops

triangularis

Souza, 2007

barragem entre Fazenda -2.578 -44 854 IBSP 21768 Henrique Braz This study

Sao José de Canaa e

Fazenda Sao Luiz, durante

a limpeza da represa, Peri

Mirim, Maranhao

Canindé, Rio Gurupi -2.57000 -46.52000 MZUSP 4420 Nelson R. de Albuquerque &

Albuquerque Lema, 2008

Hydrops

triangularis

Hydrops

triangularis

Hydrops

Maica, Santarém, Para -2.55 -54.4 MCT-PUCRS Frota et al., 2005

triangularis 10608

Hydrops Taparinha -2.53333 -54.28333 [no agent data] GBIF

triangularis [Taperinha],Brazil 177374

triangularis

Hydrops

triangularis

MCZ

,Brazi R-

Hydrops Taparinha -2.53333 -54.28333 R-177373 [no agent data] GBIF

[Taperinha],Brazil

Taparinha -2.53333 -54.28333 MCZ [no agent data] GBIF

[Taperinha],Brazil R-177371

Taparinha -2.53333 -54.28333 R-177372 [no agent data] GBIF

[Taperinha],Brazil

Hydrops

triangularis

Hydrops Taperinha, AM -2.53333 -54.28333 MCZ 177371 Nelson R. de Albuquerque &

Albuquerque Lema, 2008

Sao Luiz, MA -2.53000 -44 30278 IBSP 55162 Nelson R. de Albuquerque &

Albuquerque Lema, 2008

Santarém, Para -2.443 -54.708 LPHA 2433 Frota et al., 2005

triangularis

Hydrops BR Bairro da Esperanga, area -2.433 -54.7 LPHA 1205 Henrique Braz This study

triangularis da COSAMPA, Santarém,

Para

Hydrops Tomé-Acu, Para -2.419 -48.152 IBSP 14829 Henrique Braz This study

triangularis

Hydrops Reserva INPA-WWF, -2.417 -59.717 MZUSP 8432 Albuquerque &

triangularis Manaus, Amazonas

triangularis

Hydrops

triangularis

Hydrops

Lema, 2008

Hydrops -2.15194 -56.09194 | MPEG 22682 | [no agent data]

Montalvo, Pastaza -2.067 -76.967 DHMECN 252

triangularis

Hydrops BR Estrada do Acara, km 16, -2.058 -48.715 MPEG 15507

triangularis Sao Domingos do Capim,

Para

Hydrops Igarapé Pirajuara, Estrada -2.001 -47.893 MPEG 9417 Henrique Braz This study

do Acara, Para

triangularis

Cisneros-

Heredia, 2005

Hydrops

Albuquerque &

Lema, 2008

triangularis

Hydrops

Obidos, Para -1.918 -55.518 IBSP 14938 Henrique Braz This study

triangularis

Hydrops Obidos, PA -1.91778 -55.51778 MCZ 3671 Nelson R. de Albuquerque &

triangularis Albuquerque Lema, 2008

Hydrops UHE Balbina, Presidente -1.916 -59.473 IBSP 51496 Henrique Braz This study

Figueiredo, Amazonas

Obydos - 1.90833 -55.51889 R-3671 Louis Agassiz GBIF

triangularis

Hydrops BR ECFPn/MPEG/ Floresta - 1.90667 -51.38222 MPEG 19431 | [no agent data] na

triangularis Nacional de Caxiuaa,

Itaperu, Melgaco, PA

Hydrops B Nova Vida, 25 km distante -1.81389 -46.10750 MPEG 10341 Nelson R. de Albuquerque &

triangularis do rio Gurupi, Junco do Albuquerque Lema, 2008

Maranhao, BR-316, MA

Amphib. Reptile Conserv. 137 April 2019 | Volume 13 | Number 1 | e173

triangularis

Hydrops

New distributional records for Hydrops in Peru and Brazil

Hydrops Colénia Nova, proxima do - 1.80889 -46.40389 MPEG 10299 Nelson R. de Albuquerque &

triangularis Rio Gurupi, PA Albuquerque Lema, 2008

Hydrops ECFPn/MPEG/ Floresta -1.792 -51.434 MPEG 21870 | Henrique Braz This study

triangularis Nacional de Caxiuana,

Melgaco, Para

Rio Gurupi, Nova Vida, -1.774 -46.529 MPEG 12254 | Henrique Braz This study

25 km distante do rio, BR-

316, Para

BR Sao Pedro, Capitaéo Pogo, - 1.74500 -47.06500 MPEG 1627 Nelson R. de Albuquerque &

PA Albuquerque Lema, 2008

Capit&o Poco, Para -1.745 -47.065 MPEG 776 This study

Hydrops Santa Luzia, Capitéo Poco, -1.745 -47.065 Cunha &

triangularis Para Nascimento,

1978

Rio Gurupi, Col6nia Nova, -1.727 -46.318 MPEG 10300 | Henrique Braz This study

triangularis proximo do rio, BR-316,

Viseu, Para

km 220 da BR-316, antigo -1.721 -46.622 MPEG 2989 Henrique Braz This study

km 74 de Capanema,

Viseu, Para

This study

Albuquerque &

Lema, 2008

Albuquerque &

Lema, 2008

Hydrops BR Limao Grande, Ourém, PA -1.51528 -47.00211 MPEG 1272 [no agent data] na

triangularis

Hydrops BR Instituto Agrondémico do -1.454 -48.446 IBSP 14677 Henrique Braz This study

triangularis Norte, Belém, Para

GBIF

Hydrops BR Belém, PA -1.43730 -48.47060 MPEG 18678 Nelson R. de Albuquerque &

triangularis Albuquerque Lema, 2008

Hydrops BR Gurupa, PA - 1.40500 -51.64000 MPEG 15158 Nelson R. de Albuquerque &

triangularis Albuquerque Lema, 2008

Hydrops Macapazinho, Castanhal, -1.389 -47.984 MPEG 8648 Albuquerque &

triangularis Para

Lema, 2008

Hydrops BR-316, km 6, -1.38 -48 393 MPEG 16704 Albuquerque &

triangularis Transportadora Elo Ltda., Lema, 2008

Ananindeua, Para

Albuquerque &

Lema, 2008

This study

Amaral, 1935

Albuquerque &

Lema, 2008

Albuquerque &

Lema, 2008

Hydrops BR Ilha de Outeiro, Belém, PA -1.25100 -48.45610 MPEG 13070 Nelson R. de Albuquerque &

triangularis Albuquerque Lema, 2008

Amphib. Reptile Conserv. 138 April 2019 | Volume 13 | Number 1 | e173

von May et al.

[rca Bela Vista, Viseu, Pard -1.197 -46.14 MPEG 15924 This study

Bela Vista, Viseu, PA -1.19694 -46.14000 MPEG 15925 Nelson R. de Albuquerque &

Albuquerque Lema, 2008

Igarapé Rio das Gatas, -1.19583 -47.18083 MPEG 18552 Nelson R. de Albuquerque &

Capanema, PA Albuquerque Lema, 2008

Hydrops Peixe-Boi, Para -1.192 -47.314 MPEG 683 Albuquerque &

triangularis Lema, 2008

Hydrops

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops Fazenda Real, Viseu, Para -1.192 -46.218 MPEG 5284 Albuquerque &

Camargo, 2004

Peixe-Boi, PA -1.19194 -47.31389 MPEG 1403 Nelson R. de Albuquerque &

Albuquerque Lema, 2008

triangularis

Hydrops

triangularis

Hydrops

triangularis

B Cacoal, Augusto Corréa -1.167 -46.8 MPEG 6650 Henrique Braz This study

Para

Hydrops BR Ilha do Mosqueiro, Baia -1.162 -48.471 Cunha &

triangularis de Guajara, proximo de Nascimento,

Belém, Para 1978

Hydrops B Santo Antdnio do Taua, -1.152 -48.129 MPEG 5699 Albuquerque &

triangularis Para Lema, 2008

Hydrops B Santo Anténio do Taua, PA -1.15194 -48.12889 MPEG 4719 Nelson R. de Albuquerque &

triangularis Albuquerque Lema, 2008

Hydrops B Igarapé-Acu, Para -1.127 -47.618 MPEG 908 Henrique Braz This study

triangularis

B Igarapé-Agu, PA -1.12694 -47.61778 MPEG 906 Nelson R. de Albuquerque &

triangularis Albuquerque Lema, 2008

Hydrops BR Baia do Sol, ilha de - 1.10000 -48.40000 MPEG 22831 | [no agent data] na

triangularis Mosqueiro, Belém, PA

B Bom Jesus, Braganga, PA - 1.06278 -46.77278 MPEG 5118 Nelson R. de Albuquerque &

triangularis Albuquerque Lema, 2008

Hydrops B Cacoal, Augusto Correa, -1.02194 -46.64500 MPEG 5366 Nelson R. de Albuquerque &

triangularis PA Albuquerque Lema, 2008

B UHE Luis E Magalhaes, -1.01844 -48 333611 IBSP 65594 Nelson R. de Albuquerque &

triangularis Palmas, TO Albuquerque Lema, 2008

EC,

Hydrops EC Parque Nacional Yasuni, -1 -76 FHGO 2468 Cisneros-

triangularis km 28 Estrada Pompeya Heredia, 2005

Sur-Iru, ca. 300 m de

altitude, Orellana

Santa Rosa, Estrada da -0.95556 -48 08611 MPEG 4613 Nelson R. de Albuquerque &

Vigia, Vigia Albuquerque Lema, 2008

Hydrops

triangularis

Hydrops

triangularis

Hydrops

aed) Colares, PA -0.93694 -48.28194 | MPEG 18937 | [no agent data]

Trombetinha, Santarém -0.92889 -47.396944 MPEG 3243 Nelson R. de Albuquerque &

triangularis Novo, PA Albuquerque Lema, 2008

Hydrops Serra do Navio, AP -0.89583 -52.001944 IBSP 27395 Nelson R. de Albuquerque &

triangularis Albuquerque Lema, 2008

triangularis

Hydrops

Igarapé Parijo, Salindpolis, -0.62889 -47.35583 MPEG 19711 | [no agent data] na

Limoncocha, Provincia de -0.41000 -76.62 KU 183515 Nelson R. de Albuquerque &

triangularis Sucumbios Albuquerque Lema, 2008

Hydrops B Ilha de Santana, AP -0.03500 -51.175 IBSP 14826 Nelson R. de Albuquerque &

triangularis Albuquerque Lema, 2008

Hydrops CO Rio Vaupés, Mitu, Vaupés 1.183 -70.167 ANPS 25733 Roze, 1957b;

triangularis Albuquerque &

Lema, 2008

Amphib. Reptile Conserv. 139 April 2019 | Volume 13 | Number 1 | e173

triangularis

Hydrops Auca Via Cononaco, km -0.708 -76.888 DHMECN 82 Cisneros-

135, Orellana Heredia, 2005

Hydrops

New distributional records for Hydrops in Peru and Brazil

Hydrops CO Rio Vaupés, Mitu, 1.26000 -70.23 ANPS 25733 Nelson R. de Albuquerque &

triangularis Departamento de Vaupés Albuquerque Lema, 2008

Hydrops B Parque Nacional 22193: -54.588 Lima, 2008

triangularis Montanhas do

Tumucumaque, proximo

a triplice fronteira com

o Suriname e Guiana

Francesa, Amapa

Hydrops SR OELEMARIE 3.10000 -54.51667 84673 GROEN, GBIF

triangularis JANEA

Hydrops SU Oelemari, Sipaliwini ou 3.105 -54.54] CM 84673 Albuquerque &

triangularis Marowijne? Lema, 2008

Hydrops BR Rio Branco, em frente a 3.417 -61.667 MPEG 16697 | Henrique Braz This study

triangularis Ilha de Maraca, Igarapé do

Cojubim, Roraima

Hydrops BR Anapu, UHE de Belo 3.47194 -51.19778 MPEG 22363 | [no agent data] na

triangularis Monte, PA

Hydrops GU Maripasoula, Guyane 3.64443 -54.03377 MNHN Nelson R. de Albuquerque &

triangularis 1989.3052 Albuquerque Lema, 2008

Hydrops BR BR-156, Km 90, Aldeia 3.83300 -51.83330 MPEG 21725 | [no agent data] na

triangularis Tukai, Oiapoque, AP

Hydrops Finca Las Mercedes, 14 4.02730 -69.20444 UTA 32076 Nelson R. de Albuquerque &

Km South of Villavicencio, Albuquerque Lema, 2008

Departamento del Meta

Hydrops Villavicencio, 4.14000 -73.63 MZUSP 5994 Nelson R. de Albuquerque &

triangularis Departamento del Meta Albuquerque Lema, 2008

Hydrops G Kabocali, 101 m, Potaro- 4.285 -58.748 Donnelly et al.,

triangularis Siparuni 2005

Hydrops Rio Manacacias, Hacienda 4.383 -72.067 MZUSP 6103 Albuquerque &

triangularis La Esperanza, Meta Lema, 2008

Hydrops GF Matoury, Cayenne 4.85 -52.333 MNHN Gasc &

triangularis 1978.25 Rodrigues, 1980;

Albuquerque &

Lema, 2008

triangularis

Hydrops

Hydrops Matoury, Guyane Francaise 4.85000 -52.33000 MNHN Nelson R. de Albuquerque &

1978.2500 Albuquerque Lema, 2008

Camp de Saint Eugene, 4.85056 -53.05583 MNHN Nelson R. de Albuquerque &

Guyane 1996.4586 Albuquerque Lema, 2008

triangularis

Hydrops

triangularis

floresta, 2 km ao Leste, San -61.133 MHNLS 10953 Gorzula &

Ignacio de Yuruani, Bolivar Sefiaris, 1998

Iquitos, Departamento de 5.11670 -54.98330 AMNH 52017 Nelson R. de Albuquerque &

triangularis Loreto Albuquerque Lema, 2008

Hydrops CO vereda Ulere, finca 5.186047222 | -70.98163889 ICN 235

triangularis Altamira, San Luis de

Palenque, Casanare

Hydrops G Rio Essequibo, Maripa, del -58.928 AMNH Roze, 1957b;

triangularis Potaro-Siparuni R-18162 Scrocchi et al.,

2005

Hydrops Rio Suriname, proximo a 5.343 -54.995 RMNH 13610 Albuquerque &

triangularis Kadjoe, Para

Hydrops

Hydrops La Madeleine, Guyane 4.92000 -52.32000 MNHN Nelson R. de Albuquerque &

triangularis Francaise 1988.167 Albuquerque Lema, 2008

Angarita-Sierra,

2014

Lema, 2008

Hydrops

Corregimiento de Santa 5.36300 -67.86000 ITAvH- Juan M Renjifo GBIF

triangularis Rita CT-16167

Hydrops Cgto. Santa Rita. 5.36333 -67.86000 TAvH-R-5133 Renjifo, Juan GBIF

triangularis

Amphib. Reptile Conserv. 140 April 2019 | Volume 13 | Number 1 | e173

Hydrops GC

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Hydrops

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Hydrops G

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Hydrops

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Hydrops

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Hydrops

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Hydrops GU

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Hydrops

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops VE

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

triangularis

Amphib. Reptile Conserv.

Hydrops

triangularis

von May et al.

Corregimiento Santa Rita 5.36333 -67.86000 TAvH-R-5133 | Renjifio, Juan- GBIF

Manuel

Sinnamary, Cayenne 5.372 -52.952 IBSP 13760 This study

Berbice, Upper Demerara- 5.45 -57.95 BMNH Roze, 1957b

Berbice 53.4.6.12

Kamakusa, Guyana 5.95000 -59.9 AMNH 25035 Nelson R. de Albuquerque &

Albuquerque Lema, 2008

Mazaruni-Potaro, Kartabo, 6.35 -58.683 AMNH Scrocchi et

Cuyuni-Mazaruni R-14141 al., 2005;

Albuquerque &

Lema, 2008; Cole

et al., 2013

Rio Dunoon-Dumerara 6.43333 -58.30000 UMMZ 47737 Nelson R. de Albuquerque &

Albuquerque Lema, 2008

Kurupung, Upper Mazaruni 6.46667 -59.166667 | UMMZ 83642 | Nelson R. de Albuquerque &

District Albuquerque Lema, 2008

Lama Creek, Rio 55 -57.95 AMNH 36100 Roze, 1957b;

Demerara, Demerara- Albuquerque &

Mahaica Lema, 2008

El Dorado, Bolivar 6.715 -61.638 | MBUCV 1998 | Roze, 1957b

iTS

Roze, 1957b;

Albuquerque &

Onverwacht, Para District 5.59000 -55.19000 RMNH 27371 Nelson R. de Albuquerque &

Albuquerque Lema, 2008

Kartabo, Mazaruni-Potaro 6.35000 -58.68333 AMNH 14134 Nelson R. de Albuquerque &

District Albuquerque Lema, 2008

Kurupung, Upper Mazaruni 6.467 -59.167 UMMZ 83642

district, Cuyuni-Mazaruni

Lema, 2008

6.75000 -58.500 AMNH 18163 Nelson R. de Albuquerque &

Albuquerque Lema, 2008

Georgetown, Demerara- -58.167 AMNH 36141 Roze, 1957b;

Mahaica Albuquerque &

Camargo, 2004

Cafio Leon, 3 km de El 7.4555556 -71.9166667 FA Roze, 1957b

Jordan, Amazonas Tachira

Cafio Guaritico, Mantecal, 7.80000 -69.03333 MHNLS 8008 Nelson R. de Albuquerque &

Estado Apure Albuquerque Lema, 2008

San Fernando de Apure 7.893 -67.472 MHNLS 755 Roze, 1957b

Apure

Bolivar, Estado Bolivar 8.11667 -63.55000 USNM 56235 Nelson R. de Albuquerque &

Albuquerque Lema, 2008

Ciudad Bolivar, Bolivar 8 -63.55 USNM 56235 Roze 1957b;

Albuquerque &

Lema, 2008

Curiapo, Estado Delta 8.56667 -68.59167 MHNLS 12973 | Nelson R. de Albuquerque &

Amacuro Albuquerque Lema, 2008

Sacupana del Cerro, Delta 8.573 -61.652 MCNC 5859 Molina et al.,

Amacuro 2004

Curiapo, Delta Amacuro 8.576 MHNLS Lf Albuquerque &

12973-4

Lema, 2008

Hato Pifiero, Estado 8.93263 -68.08149 EBRG 3665 Nelson R. de Albuquerque &

Cojedes Albuquerque Lema, 2008

141 April 2019 | Volume 13 | Number 1 | e173

New distributional records for Hydrops in Peru and Brazil

Estacion Pisciola, 8.95944 -69.47278 MCNG 1602 Allen Herp Review

Municipio Papelon Markezich 2001

Guanare, on road to 9.044 -69.75 MCNG 1006 Markezich, 2001

Barinas, Guanare,

Portuguesa

cafio Jarisiduina, isla 9.167 -60.85 MHNLS 10940 Gorzula &

Barril, Delta Amacuro Sefiaris, 1998;

Molina et al.,

2004

Hydrops

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

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Estado Portuguesa 9.55389 -61.11667 MHNLS 7622 Nelson R. de Albuquerque &

Albuquerque Lema, 2008

Hydrops SW San Carlos City, 9.57167 -68.62056 MBUCV 7117 G.R. Rivas Herp Review

triangularis Rincon Moreno Fuenmayor & 2000

O. Fuentes

Hydrops San Carlos, Estado Cojedes 9.65722 -68.59167 MHNLS 6444 Nelson R. de Albuquerque &

triangularis Albuquerque Lema, 2008

Hydrops

triangularis

Cafio Manamo, Isla 9.91667 -61.11667 EBRG 2617 Nelson R. de Albuquerque &

del Tigre, Estado Delta Albuquerque Lema, 2008

Amacuro

Hydrops

Mole, 1924

Thais Guedes;

OBS. PESS.

Nariva Swamp 10.417 -61.083 Ford & Ford,

2002

North Manzanilla Beach 10.473 -61.049 SS Boos, 2001

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

10.487 61.216 Mole, 1924

Talparo, TR 10.50000 -61.266667 ANSP 23178 Nelson R. de Albuquerque &

Albuquerque Lema, 2008

Talparo 10.51 -61.265 ANSP 23178 Roze 1957b;

Albuquerque &

it ‘ee

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

triangularis

Hydrops

Mole, 1924

10.631 -61.275 Mole, 1924

Orange Grove Estate, 10.637 -61.368 RMNH 10177 Brongersma,

Tacarigua 1956

Albuquerque &

Tucker Valley 10.714 -61.609 AMNH

64463a, b Lema, 2008

km 23 da estrada de -3.35417 -64.71140 MPEG 1887 [no agent data] na

Maracana, PA

triangularis

triangularis Museet,

Goteborg 4754

Amphib. Reptile Conserv. 142 April 2019 | Volume 13 | Number 1 | e173

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

13(1) [General Section]: 143-161 (e174).

New reptile records from Lekedi Park and Haut-Ogooue

Province, southeastern Gabon

"Olivier S. G. Pauwels, Stephan Morelle, *Jean-Louis Albert, *Piero Carlino,

SNil Rahola, and °Jean-Frangois Trape

‘Département des Vertébrés Récents, Institut Royal des Sciences naturelles de Belgique, Rue Vautier 29, B-1000 Brussels, BELGIUM *Parc de

la Lékédi, Bakoumba BP 52, GABON ?BP 5423, Libreville, GABON *Museo di Storia naturale del Salento, Sp. Calimera-Borgagne km 1, 73021

Calimera, ITALY *Institut de Recherche pour le Développement (IRD), UMR MIVEGEC (University of Montpellier, CNRS 5290 IRD 224), Centre

IRD de Montpellier, Montpellier, FRANCE & Centre International de Recherches Meédicales de Franceville, Franceville, GABON Institut de

Recherche pour le Développement (IRD), Laboratoire de Paludologie et Zoologie médicale, UMR MIVEGEC, B.P. 1386, Dakar, SENEGAL

Abstract.—Presented is a compilation of new locality records with natural history data for 49 reptile species

in Haut-Ogooué Province, southeastern Gabon, Equatorial Africa. One new snake species record is added to

Gabon (Colubridae: Philothamnus hughesi), 14 new reptile species records to Haut-Ogooueé Province, three

new reptile records for Léconi Park, and 28 new reptile records for Lekédi Park. A predation case is reported of

Naja melanoleuca preying on Dasypeltis fasciata and of Boaedon perisilvestris on a micromammal.

Resumé.—Nous presentons une compilation de nouvelles mentions de localiteés accompagnées de notes

d’histoire naturelle pour 49 espeéces de reptiles de la Province du Haut-Ogooue, sud-est du Gabon, Afrique

équatoriale. Nous ajoutons une espeéce de serpent (Colubridae: Philothamnus hughesi) a la faune du Gabon,

14 espéces de reptiles a la Province du Haut-Ogooue, 3 espéces de reptiles au Parc de Léconi, et 28 especes

de reptiles au Parc de la Lekédi. Nous rapportons un cas de predation par Naja melanoleuca sur Dasypeltis

fasciata, et par Boaedon perisilvestris sur un micromammifeére.

Keywords. Biodiversity, herpetofauna, Testudines, Crocodylia, Squamata, mandrill, safari park, conservation, Chaillu

Massif

Citation: Pauwels OSG, Morelle S, Albert J-L, Carlino P, Rahola N, Trape J-F. 2019. New reptile records from Lékédi Park and Haut-Ogooué Province,

southeastern Gabon. Amphibian & Reptile Conservation 13(1) [General Section]: 143-161 (e174).

4.0 International (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in any

medium, provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced, are

as follows: official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org.

Received: 27 May 2018; Accepted: 22 October 2018; Published: 11 April 2019

Introduction Additions to the species list for the province were

since presented by Pauwels et al. (2010, 2016b, 2018c)

The herpetofauna of the Equatorial African country Ga-

bon is still poorly known. Among its nine provinces, one

of the least surveyed is Haut-Ogooué Province, in the

southeastern corner of the country, along the Republic

of Congo. The first herpetological collection from Haut-

Ogooué Province, studied by Mocquard (1887), was

made by the members of an expedition led by the explor-

er Savorgnan de Brazza. Only 17 reptile species were re-

corded from the province until the beginning of the 21st

Century, when a number of additions were documented

by Pauwels et al. (2007), Pauwels and David (2008a,b)

and Pauwels and Sallé (2009), bringing the total number

of species in Haut-Ogooué Province to 39 in the herpeto-

logical synthesis presented by Pauwels and Vande weghe

(2008).

Correspondence. * osgpauwels@yahoo,.fr

Amphib. Reptile Conserv.

and Ineich and Le Garff (2015). Additional locality re-

cords within the province were added by Pauwels et al.

(2016a, 2017a,d,e, 2018a). Although numerous records

were made during the last decade, it is obvious that the

actual number of species inhabiting the province and its

protected areas must be much higher, in view of the di-

versity of savanna and forest habitats represented and the

herpetofauna known from neighbouring areas in Gabon

and the Republic of Congo.

The present contribution 1s mostly based on opportu-

nistic observations made by one of us (Stephan Morelle ~

SM) while working from March 2016 to February 2018,

i.e., during two full calendar years, as a veterinarian for

the mammals of the Parc de la Lékédi (Lékédi Park, ca.

1°45'32"S, 13°03'16"E) in Lékoko Department, a 14,000

April 2019 | Volume 13 | Number 1 | e174

Pauwels et al.

ha fenced safari park managed by the private company

Sodepal (Société d’Exploitation du Parc de la Lékédi;

see Auzias and Labourdette 2011; Vande weghe 2008).

Many of our new records were also made in Bakoumba

(ca. 1°49°43.5”S, 13°00’08.7”E; elevation 609 m above

sea level), a small town located at about seven km S of

Lékédi Park, in the same administrative department.

Materials and Methods

The new voucher material was identified using the keys

and morphological information provided by Brygoo and

Roux-Esteve (1983), Trape and Roux-Esteve (1990),

Chippaux (2006), Trape and Mané (2006), Pauwels and

Vande weghe (2008), Wagner et al. (2009), Pauwels et al.

(2010), Ineich and Le Garff (2015), and Trape and Medi-

annikov (2016). Geographic coordinates for new locality

records are provided in Table 1. Most snake vouchers de-

rive from dead-on-road individuals. Snake ventral scales

were counted according to Dowling’s (1951) method.

Snake dorsal scale rows were counted at one head length

behind head, at midbody (above the ventral correspond-

ing to half of the total number of ventrals), and at one

head length before vent; subcaudal counts exclude the

terminal pointed scale. Paired meristic characters are giv-

en left/right. The sex of preserved snakes was determined

by dissection of the tail base. Specimens’ main diagnos-

tic morphological characters are provided in Appendix 1

and within the species accounts.

Abbreviations: CIRMF = Centre International de Re-

cherches Médicales de Franceville, Franceville, Gabon;

Dept. = Department; Prov. = Province; MNHN = Na-

tional Museum of Natural History, Paris, France, MSNS

= Natural History Museum of Salento, Calimera, Italy;

NP = National Park; RBINS = Royal Belgian Institute

of Natural Sciences, Brussels, Belgium; RMCA = Royal

Museum for Central Africa, Tervuren, Belgium.

Results

TESTUDINES

PELOMEDUSIDAE

Pelusios gabonensis (Dumeril, 1856)

On 4 October 2016 SM photographed a subadult indi-

vidual in a forest stream in Lékédi Park. It showed the

typical brown carapace with a black vertebral line, a

poorly marked median keel on the carapace, a brown

head without vermiculations but with a black dorsal

triangle. This constitutes the first record for the park.

Within the same Dept., the species had already been re-

corded from Bakoumba by Maran and Pauwels (2005).

On 22 January 2017 at 17h25 Nil Rahola (~ NR) photo-

graphed a subadult individual in the northwestern sub-

urbs (1°36’58.3”S, 13°34’48.7”E) of Franceville in Pas-

sa Dept. (Fig. 1). This represents a new locality record

(see the compilation on the distribution of this species in

Gabon by Maran and Pauwels 2005). Widely distributed

in Gabon, this terrapin is heavily hunted for food, and is

so far known from only three protected areas in the coun-

try (Pauwels 2016).

Fig. 1. Subadult Pe/usios gabonensis in Franceville, Haut-Ogo-

oué Prov., southeastern Gabon. Photograph by N. Rahola.

TESTUDINIDAE

Kinixys erosa (Schweigger, 1812)

On 9 October 2016 SM photographed a young individual

in a savanna area in Lékédi Park (Fig. 2). This constitutes

the first record for the park. Within the same Dept., SM

also photographed on 18 January 2018 an individual in

Bakoumba, from where the species had already been re-

corded by Maran and Pauwels (2005). This tortoise, eval-

uated as Data Deficient by the International Union for the

Conservation of Nature IUCN; see Tortoise & Freshwa-

ter Turtle Specialist Group 1996), is a very popular food

item in all parts of Gabon, but it seems to cope with it and

is the most common and widely distributed chelonian in

the country, reported from nearly all national protected

areas (Pauwels 2016).

Table 1. Main localities for new herpetological records in Haut-Ogooué Province, Gabon.

Locality name

Administrative department

Bakoumba Lékoko

Franceville Passa

Lékédi Park Lékoko

Lemanassa Lékoko

Moanda Lébombi-Léyou

Mounana Lébombi-Léyou

Amphib. Reptile Conserv.

144

Geographic coordinates

ca. 1°49°43.5”S, 13°00°08.7”E

ca. 1°36758.3”S, 13°34’48.7°E

ca. 1°45'32”S, 13°03'16” E

2°00'07.9"S, 12°56'05.6"E

1°32)13-2"S. 13° 1435.97 E

1°24'05.0"S, 13°09'41.0"E

April 2019 | Volume 13 | Number 1 | e174

New reptile records from southeastern Gabon

ci ei ee . ya .

Fig. 2. Young Kinixys erosa in Lékédi Park, Haut-Ogooué

Prov., southeastern Gabon. Photograph by S. Morelle.

CROCODYLIA

CROCODYLIDAE

Mecistops cataphractus (Cuvier, 1824)

Pearson et al. (2007) presented the picture of an indi-

vidual in Plateaux Batéké NP, clearly showing the whole

body and the long snout. Fig. 3 shows an adult individual

photographed on 17 October 2016 in Lékédi Lake (Lac

Lékédi) in Lékédi Park. This individual is regularly ob-

served on this dead tree when one navigates on the lake,

and it jumps forward when it is too closely approached.

It climbs on this tree on sunny days as well as on cloudy

days, and often keeps its mouth widely open. No census

of the Mecistops population of the 120 ha Lékédi Lake

was ever done, but a45 minutes canoe trip by night on the

lake in June 2017 by SM revealed ten distinct individu-

als, mostly juveniles and subadults. A second canoe tour

on the lake on 21 July 2017 between 18h30 and 21h30

revealed 25 distinct individuals, indicating that further

search efforts might demonstrate the existence of a vi-

able population. Our observations represent a new Dept.

record and a new record for the park (not listed from this

Dept. by Pauwels and Vande weghe 2008; Pauwels et al.

2016b, 2017e).

: ( f f ‘ :

= 4 eS dG 's — = = _ =

= CO —y—————

Fig. 3. Adult Mecistops cataphractus on a dead tree in Lékédi

Park, Haut-Ogooué Prov., southeastern Gabon. Photograph by

S. Morelle.

Amphib. Reptile Conserv.

Osteolaemus tetraspis Cope, 1861

On 13 November 2017 at 17h00 SM photographed an

adult individual in a ditch in Lékédi Park (Fig. 4). New

record for the Dept. and for the park (not listed from

this Dept. by Pauwels and Vande weghe 2008; Pauwels

2016). The Dwarf crocodile is often sold as food, but

it still ubiquitous in Gabon, even in degraded areas; it

has been recorded from nearly all protected areas in the

country (Pauwels 2016).

Fig. 4. Adult Osteolaemus tetraspis in Lékédi Park, Haut-Ogo-

oué Prov., southeastern Gabon. Photograph by S. Morelle

SQUAMATA

AGAMIDAE

Agama agama (Linnaeus, 1758)

This highly anthropophilic species was already recorded

by Pauwels et al. (2016b) from Bakoumba, where both

Agama species probably co-exist, like in an increasing

number of localities in Gabon. The propagation of Aga-

ma agama within Gabon is facilitated by human activi-

ties and the development of roads (Pauwels et al. 2004).

TT) liye ir aed

€

Fig. 5. Ventral view of an adult dead-on-road male Agama leb-

retoni found on the road between Lékédi Park and Bakoumba,

Haut-Ogooué Prov., southeastern Gabon. Photograph by S.

Morelle.

Agama lebretoni Wagner, Barej & Schmitz, 2009

Fig. 5 illustrates the ventral surface of a dead-on-road

individual found in June 2016 by SM on the road be-

tween Lékédi Park and Bakoumba. Other individuals

were observed by SM in Lékédi Park where the species

is common. The male RBINS 18497 was caught by SM

April 2019 | Volume 13 | Number 1 | e174

Pauwels et al.

in Léconi Park, Plateaux Dept., on 25 December 2017.

It shows a pale vertebral stripe, a reticulated throat color

pattern, 71 scale rows at midbody, a snout-vent length

of 93 mm, and a tail length of 144 mm. It feigned death

while handled. New records for both Depts. and for the

parks (not recorded from these Depts. by Pauwels et al.

2016a—b, 2017b).

CHAMAELEONIDAE

Trioceros cristatus (Stutchbury, 1837)

In a booklet on the animals of Gabon, Le Garff (2015)

presented the photograph of an adult male Trioceros cris-

tatus with a well-developed dorsal crest, without a pre-

cise locality. This photograph was actually taken in the

vivarium of Lékédi Park (Le Garff, pers. comm. to OSGP

2015). The exact locality where the individual was cap-

tured is unknown, but it was certainly caught within the

Dept. in the vicinity of Lékédi Park. New Prov. record.

The species is uncommon in Gabon and known from

only two protected areas in the country (Pauwels et al.

2016a; Pauwels 2017).

Trioceros owenii (Gray, 1831)

On 28 March 2017 at 14h00 SM encountered a young

male individual in Lékédi Park, showing three short

horns (Fig. 6). On 9, 12, and 15 January SM encoun-

tered and photographed three distinct adult males in Bak-

oumba, all on bare ground between houses in the village.

They had distinctly more developed horns than the young

individual shown on Fig. 6. Put in presence of each other

before being released, two males immediately began a

fight. New record for the park and new Dept. record. In

Haut-Ogooué Prov., this species had been recorded so

far only from Franceville in Passa Dept. (Pauwels et al.

2007) and from Akiéni in Lékoni-Lékori Dept. (Pauwels

et al. 2016a).

Mucrhel es ol

Fig. 6. Young male Trioceros owenii in Lékédi Park, Haut-

Ogooué Prov., southeastern Gabon. Photograph by S. Morelle.

Rhampholeon spectrum (Buchholz, 1874)

On 8 September 2016 SM photographed an adult male

individual of this relatively common forest leaf litter cha-

meleon in Lemanassa (ca. 23 airline km S-SW of Bak-

oumba), Lékoko Dept. (Fig. 7). New Prov. record (not

Amphib. Reptile Conserv.

listed from this Prov. by Pauwels and Vande weghe 2008;

Pauwels et al. 2008). This village is located at about five

km from the border with the Republic of Congo. Like the

two chameleons above, this species has been evaluated

as Least Concern by the IUCN (Mariaux and LeBreton

2010).

Fig. 7. Adult male Rhampholeon spectrum in Lemanassa, Haut-

Fig. 8. Adult Hemidactylus mabouia in Léconi Park, Haut-

Ogooué Prov., southeastern Gabon. Photograph by S. Morelle.

“3

GEKKONIDAE

Hemidactylus mabouia (Moreau de Jonneés,

1818)

This anthropophilic gecko was regularly observed by SM

on the buildings of Lékédi Park. We examined the pho-

tograph of a hatchling and its egg taken by A. Willaume

on 3 March 2016 at 17h41 in a house in Bakoumba. The

hatching date of this gecko falls within the hatching peri-

od (December to April) known for the species in Gabon.

On 28 March 2016 SM photographed an adult individual

on a tree in Léconi Park (Parc de la Léconi), Plateaux

Dept. (Fig. 8). We examined the photograph of an adult

individual taken in July 2017 (exact date unknown) by

N. Longin inside a house in Moanda in Lébombi-Léyou

Dept. New Dept. records and new records for Léconi and

Lékédi parks (not listed from these Depts. by Pauwels

and Vande weghe 2008; Pauwels et al. 2016b, 2017b;

Pauwels 2017). Olivier S.G. Pauwels (~ OSGP) exam-

April 2019 | Volume 13 | Number 1 | e174

New reptile records from southeastern Gabon

ined three adult individuals (RMCA 28230-28232, not

individually numbered) collected on 20-22 November

1964 in “Franceville, Riv. [=River] Passa.’’ One has a

snout-vent length of 62 mm, a tail length of 59 mm (last

39 mm regenerated), 35 femoro-precloacal pores, and 16

dorsal tubercle rows at midbody. Another has a snout-

vent length of 58 mm, a tail length of 54 mm (only first 8

mm original), 32 femoro-precloacal pores, and 16 dorsal

tubercle rows at midbody. The last one has a snout-vent

length of 57 mm, a tail length of >30 mm (tail original,

but tip missing, healed), 35 femoro-precloacal pores, and

15 dorsal tubercle rows at midbody. The species was al-

ready known from Franceville (Pauwels and Vande we-

ghe 2008).

GERRHOSAURIDAE

Gerrhosaurus nigrolineatus Hallowell, 1857

A juvenile (RBINS 18472) was found dead-on-road on

2 November 2016 by SM near the camp of the Projet

Protection des Gorilles (PPG) of the Aspinall Foundation

in the northern part of the Plateaux Batéké NP. It has a

snout-vent length of 47 mm and a tail length of 105 mm.

Its dorsal scales are keeled, its ventral scales smooth; its

caudal scales are all keeled, except the ventral ones at the

base of the tail. New locality record. This lizard is com-

mon in the savannas and coastal grasslands of Gabon and

has already been recorded in nine national protected ar-

eas (Christy et al. 2008; Pauwels and Vande weghe 2008;

Vande weghe 2008: 156-157).

Fig. 9. Adult /chnotropis bivittata in Bakoumba, Haut-Ogooué

Prov., southeastern Gabon. Photograph by S. Morelle.

LACERTIDAE

Ichnotropis bivittata Bocage, 1866

On 22 November 2016 at midday SM encountered an

adult individual in Bakoumba (Fig. 9). It shows strongly

keeled scales on the dorsal surface of its head, smooth

and imbricate cycloid ventrals; its dorsals and supracau-

dals are lanceolate and strongly keeled. A short fold in

front of each arm houses red mites, and mites are also

gathered in a pocket above the posterior insertion of the

hind limbs. This represents only the second locality for

the whole country. The species was recently reported for

Amphib. Reptile Conserv.

the first time from Gabon based on a single specimen

from Lékédi Park (Ineich and Le Garff 2015). This lac-

ertid species is most probably widespread in the savan-

nas of southeastern Gabon and is expected to occur in

Plateaux Batéké NP.

SCINCIDAE

Feylinia currori Gray, 1845

On 10 April 2016 at 17h00 SM found an adult individual

(RBINS 18473) active along a road in Lékédi Park. It

has a snout-vent length of 156 mm, a tail length of 60

mm, 25 scale rows at midbody, two supranasal scales,

one loreal on each side separating the supranasal from

the preocular; on each side the ocular scale is in contact

with the 3" supralabial. Another adult individual (RBINS

18474) was found at 17h00 by SM on 11 October 2016 in

a natural ditch in the park; it was then active and quickly

moved in the soft soil to hide under a rock. It had a snout-

vent length of 164 mm, a tail length of 67 mm, two supra-

nasal scales, 26 scale rows at midbody; its ocular scale

is in contact with the 3“ supralabial. A third individual

(RBINS 18498) was found in the park by SM on 7 July

2017 at 17h30; it has a snout-vent length of 128 mm,

a tail length of 49 mm, 26 scale rows at midbody, two

supranasal scales, and on each side the ocular scale is in

contact with the 3“ supralabial. On 14 November 2017

Jean-Louis Albert (~JLA) found a young individual (to-

tal length 120 mm), freshly dead-on-road in Franceville

in Passa Dept. (Fig. 10). New record for the park and new

Prov. record; Haut-Ogooué Province was the last Gabo-

nese province from which this common species had not

yet been recorded (see distribution given by Pauwels and

Vande weghe 2008).

Fig. 10. Young Feylinia currori in Franceville, Haut-Ogooué

Prov., southeastern Gabon. Photograph by J.-L. Albert.

Trachylepis affinis (Gray, 1839)

This species was regularly observed in open areas in

Lékédi Park by SM in 2016 and 2017, and is locally com-

mon, especially near human settlements. It had already

been recorded and documented from the park by Ineich

and Le Garff (2015).

April 2019 | Volume 13 | Number 1 | e174

Pauwels et al.

Trachylepis albilabris (Hallowell, 1857)

A clutch of five eggs, half-buried in the ground, was

found in Lékédi Park by SM on 10 October 2016. They

were carefully collected and kept in a box in the park.

They all safely hatched three days later. All were released

except two (RBINS 18499-18500), which show a trans-

parent disk in each lower eyelid; separated supranasals;

prefrontals in contact; 4/4 supraoculars; on each side one

scale separating the last supraocular and the anterior su-

pratemporal; three keels per dorsal scale; and a tail length

of 31 mm. One has 30 scale rows at midbody and a snout-

vent length of 23 mm; while the other has 32 scale rows

at midbody and a snout-vent length of 24 mm. So far

clutches of only two or three eggs were reported for this

species in Gabon (Pauwels and Vande weghe 2008; Pau-

wels et al. 2017c); however it cannot be excluded that

more than one clutch was here involved. New record for

the park and for the Dept.; within Haut-Ogooué Prov.,

this skink was known so far only from Franceville in

Passa Dept. (Pauwels et al. 2016a).

Trachylepis maculilabris (Gray, 1845)

The individual RBINS 18501 was encountered by SM

on 4 January 2017 in Lékédi Park. It shows a transparent

disk in each lower eyelid, 4/4 supraoculars, 5/5 supracili-

aries; 6/6 supralabials; supranasals separated by a narrow

gap, prefrontals separated by a narrow gap; 54 scales on

a line between the nuchals and a point above the base of

the tail; a snout-vent length of 71 mm, a tail length of

125 mm; and 30 scale rows at midbody, each dorsal scale

with 5, sometimes 6 or 7, keels. RBINS 18502 was found

by SM in the park on 19 August 2017. It shows a trans-

parent disk in each lower eyelid, 4/4 supraoculars, 5/5

supraciliaries; 7/7 supralabials; supranasals separated,

prefrontals in contact; 54 scales on a line between the nu-

chals and a point above the base of the tail; a snout-vent

length of 64 mm, a tail length of 132 mm; and 30 scale

rows at midbody; each dorsal scale with 5 keels. This

species 1s locally common in open areas in the park, and

was regularly observed by SM in 2016 and 2017. It had

already been recorded and documented from the park by

Ineich and Le Garff (2015).

Fig. 11. Young Trachylepis polytropis polytropis in Lékédi

Park, Haut-Ogooué Prov., southeastern Gabon. Photograph by

S. Morelle.

Amphib. Reptile Conserv.

Trachylepis polytropis polytropis (Boulenger,

1903)

A young individual was photographed on 28 December

2017 at 17h00 by SM in Lékédi Park (Fig. 11). It shows

the species’ typical dorsal black zig-zag and green belly.

New record for the park and new Prov. record (not list-

ed from the Prov. by Pauwels and Vande weghe 2008;

Pauwels et al. 2016b). This species is found in pristine

as well as in degraded environments, but is still poorly

documented in Gabon, probably due to the fact that it 1s

more shy and arboreal than its congeners.

VARANIDAE

Varanus ornatus (Daudin, 1803)

Pearson et al. (2007) presented a picture of a moni-

tor taken in Plateaux Batéké NP that they identified as

“Varanus sp. (probably V. niloticus).” The photo shows a

juvenile individual with five transversal rows of ocellae

on the back between the fore and hindlimbs, as is typi-

cal for V. ornatus. A photograph of another individual

from Plateaux Batéké NP was presented by Pauwels and

Vande weghe (2008:127). On 25 March 2016 SM photo-

graphed an adult individual in Bakoumba (Fig. 12). On

22 October 2016 at midday SM photographed a subadult

individual (total length about 70 cm) inside a dead tree

standing above the surface of Lékédi Lake in Lékédi

Park. New Dept. record and new record for the park (not

listed from the Dept. by Pauwels et al. 2007, 2017d,e).

Although heavily hunted, this monitor is still abundant in

all parts of the country and in all kinds of environments,

from pristine forests and savannas to urban areas; it was

recorded from nearly all protected areas of Gabon (Pau-

wels, 2016). Dowell et al. (2015) did not fully resolve

the systematics within the Varanus niloticus complex and

called for “urgent taxonomic revisions in this group;”

until more conclusive results are obtained, we prefer to

maintain here the use of the name Varanus ornatus for

the Varanus populations found in Gabon.

Fig. 12. Adult Varanus ornatus in Bakoumba, Haut-Ogooué

Prov., southeastern Gabon. Photograph by S. Morelle.

BOIDAE

Calabaria reinhardtii (Schlegel, 1851)

An adult individual was photographed by day on 16 May

April 2019 | Volume 13 | Number 1 | e174

New reptile records from southeastern Gabon

2016 by SM in Lékédi Park (Fig. 13). When encountered,

it was entering a crack in a brick wall. When handled, it

never tried to bite, and displayed several times the spe-

cies’ typical defensive behaviour, 1.e., forming a ball with

its body and raising its rounded tail. Another individual

was photographed by SM in the park on 13 November

2017; while being photographed, it adopted the same de-

fensive behaviour. New record for the Dept. and for the

park (not listed from the Dept. by Pauwels and Vande

weghe 2008). This common species was first recorded

from care ag Prov. Pauwels et al. (2007).

fae aT Ps

Pe aay age a ade Tad

Mpeg | te 7, ; iv ea , : a

Fig. 13. Adult ‘Galabana reinhardtii in Lékédi Pak Haut-

Ogooué Prov., southeastern Gabon. Photograph by S. Morelle.

COLUBRIDAE

Crotaphopeltis hotamboeia (Laurenti, 1768)

An adult individual was photographed by SM on 5 Janu-

ary 2017 in Bakoumba (Fig. 14). New locality record.

Two other individuals (RBINS 18475—76; see Appendix

1) were found by SM in Lékédi Park on 17 March 2017,

and another was photographed by SM in the park on 11

December 2017. All were aggressive, attempting to bite

when approached. The species was already recorded by

Pauwels et al. (2016b) from Lékédi Park where it can be

regarded as common. It was mentioned for the first time

from Haut-Ogooué Prov. by Pauwels and Sallé (2009).

Fig. 14. Adult Crotaphopeltis hotamboeia in Bakoumba, Haut-

Ogooué Prov., southeastern Gabon. Photograph by S. Morelle.

Dasypeltis confusa Trape & Mané, 2006

An adult individual was encountered by SM in Lékédi

Park at 17h00 on 14 June 2017 (Fig. 15). It was inflat-

Amphib. Reptile Conserv.

ing itself and adopted a threatening posture all the time

it was approached. New Dept. record and new record for

the park. This recently described egg-eating species was

first reported from Gabon by Pauwels and Sallé (2009).

Fig. 15. Adult Dasypeltis confusa in Lékédi Park, Haut-Ogooué

Prov., southeastern Gabon. Photograph by S. Morelle.

Dasypeltis fasciata Smith, 1849

The adult specimen RMCA 29891 (see Appendix 1) was

collected by M. Gauduin in 1969-1970 in ““Munana, pres

de Moanda” [= Mounana near Moanda, in Lébombi-Léy-

ou Dept.]. SM encountered an adult individual in Bak-

oumba at 20h00 on 12 February 2017 (RBINS 18477;

Appendix 1; Fig. 16). Contrary to the Dasypeltis confusa

individual above, it was very calm when approached and

handled. Its poorly defined black dorsal bands are due

to the black color of the interstitial skin between scales,

which allows to easily distinguish this species from

Dasypeltis confusa which shows black scales forming

well-defined saddle-shaped marks on the dorsum, like

in the individual shown on Fig. 15. Another individual

was found by NR while it was being preyed upon by a

Naja melanoleuca on the compounds of the CIRMF in

Franceville, Passa Dept., on 25 December 2014 (Fig. 24).

First records for the Prov. (not listed from the Prov. by

Pauwels and Vande weghe 2008; Carlino and Pauwels

2015; Pauwels et al. 2016a). This species was evaluated

as Least Concern by the IUCN; the former species was

not yet evaluated.

Fig. 16. Adult Dasypeltis fasciata in Bakoumba, Haut-Ogooué

Prov., southeastern Gabon. Photograph by S. Morelle.

April 2019 | Volume 13 | Number 1 | e174

Pauwels et al.

Grayia ornata (Barboza du Bocage, 1866)

A young individual was photographed by SM in Lékédi

Park at midday on 28 December 2017 (Fig. 17). New

record for the park and for the Prov. Haut-Ogooué Prov.

is the last province of Gabon from which this common

freshwater species had not yet been recorded (Pauwels

and Vande weghe 2008).

Fig. 17. Young Grayia ornata in Lékédi Park, Haut-Ogooué

Prov., southeastern Gabon. Photograph by S. Morelle.

Hapsidophrys lineatus Fischer, 1856

A dead-on-road adult individual was found by SM on

5 June 2016 between Lékédi Park and Bakoumba (Fig.

18). It shows a round pupil, one loreal, one preocular, no

subocular, three postoculars, two supralabials in contact

with the orbit, two anterior temporals and two posterior

temporals, strongly keeled dorsals with the vertebral

row not widened, and keeled ventrals. New Dept. record

(not listed from the Dept. by Pauwels and Vande weghe

2008). Hapsidophrys lineatus being rarely encountered

in Gabon and difficult to approach due to its shyness

and fastness, Pauwels and Vande weghe (2008:165) also

had to illustrate it with the photograph of a dead-on-road

individual. This strongly contrasts with its congener H.

smaragdinus, which is one of the most commonly en-

countered snakes in the country and easier to approach.

Fig. 18. Dead-on-road adult Hapsidophrys lineatus found near

Bakoumba, Haut-Ogooué Prov., southeastern Gabon. Photo-

graph by S. Morelle.

Amphib. Reptile Conserv.

Hapsidophrys smaragdinus (Schlegel, 1837)

On 10 April 2016 SM caught and photographed an adult

individual in a garden in Bakoumba. It showed (right

side) 9(5-6) supralabials, one loreal, one preocular, no

subocular, two postoculars, one anterior temporal, a

round pupil, keeled dorsals with a non-widened vertebral

row, keeled ventrals and subcaudals, a black lateral stripe

on its green head, and a green body with bluish dots on its

dorsum. The snake repeatedly bit while being handled.

New locality record. Within this Dept., the species was

already known from the nearby Lékédi Park (Pauwels

et al. 2016b). On 6 December 2017 SM photographed

a dead-on-road individual in Moanda, Lébombi-Léyou

Dept. It showed the typical uniformly green body, a lat-

eral black stripe on the head, and strongly keeled dorsal

scales. New Dept. record (not listed from the Dept. by

Pauwels et al. 2016b, 2017b).

Philothamnus carinatus (Andersson, 1901)

On 27 November 2017 at 07hO0 SM found a dead-on-

road individual (RBINS 18503; see Appendix 1) in Léké-

di Park, at 200 m from the guard guerite marking the en-

trance of the park. Given its poor state, 1t was probably

killed by a car the day before. Its pupil is round; its ver-

tebral row is not widened; its temporal formula 1s 2+2+2

on each side. First record for the park and for the Dept.

This species, which is very common in Gabon, was first

mentioned from Haut-Ogooué by Pauwels et al. (2007).

Within the Prov., this snake was until now known only

from Franceville in Passa Dept

Se a

Fig. 19. Juvenile Philothamnus hughesi in Bakoumba, Haut-

Ogooué Prov., southeastern Gabon. Photograph by S. Morelle.

Philothamnus hughesi Trape & Roux-Esteve,

1990

On 13 May 2015 at 13h15 SM encountered a juvenile

individual in Bakoumba (Fig. 19). It showed one pre-

ocular, two supralabials contacting the orbit, two post-

oculars, 1+1 temporals, 15 dorsal scale rows at midbody,

unkeeled ventrals, a dark green head, irregular blue and

black oblique bars on the neck, and a bronze dorsum

irregularly dotted with black. On 25 November 2016

at 15h30 SM met an adult individual in Lékédi Park

April 2019 | Volume 13 | Number 1 | e174

New reptile records from southeastern Gabon

(RBINS 18478; Appendix 1). It showed 1+1 temporals

on each side, a greenish grey head, irregular blue and

black oblique bars on the neck, and a bronze dorsum

irregularly dotted with black. On 10 April 2017 a third

individual was photographed by SM in Lékédi Park. It

showed one loreal, one preocular, two postoculars, 8 su-

pralabials whose 4" and 5" in contact with the orbit, ten

infralabials, 1+1 temporals, unkeeled ventrals, a greenish

grey head, irregular blue and black oblique bars on the

neck, and a bronze dorsum irregularly dotted with black.

The meristic and chromatical characters of these three

individuals perfectly fit with the original description of

the species. Their combination of main diagnostic char-

acters (including a single anterior temporal, two rows of

temporals, two supralabials in contact with the orbit, and

the absence of a vertebral dark band) excludes an identi-

fication as Philothamnus dorsalis, P. heterodermus or P.

nitidus (compare with photographs and scalation char-

acters presented in original description and by Pauwels

and Vande weghe 2008). New record for the park, the

Prov., and for Gabon. The species was said in its original

description to occur in Gabon, based on the examination

by Trape and Roux-Esteéve (1990) of a paratype (MNHP-

A-709) from ‘Gabon.’ However, this paratype is an old

specimen, and the geographical entity then covered by

the term ‘Gabon’ does not necessarily correspond to what

is today called Gabon, and Pauwels and Vande weghe

(2008:10) thus preferred to not include this species in the

country’s herpetofaunal list until the species was prop-

erly documented with a precise locality.

Fig. 20. Dead-on-road Rhamnophis aethiopissa aethiopissa in

Bakoumba, Haut-Ogooué Prov., southeastern Gabon. Photo-

graph by S. Morelle.

Rhamnophis aethiopissa aethiopissa Gunther,

1862

Fig. 20 illustrates an adult dead-on-road individual pho-

tographed on 26 August 2016 by SM in Bakoumba. The

snake showed distinctly enlarged postparietals, a single

loreal, one preocular and two postoculars, one anterior

temporal, 17 smooth dorsal scale rows at midbody with

a widened vertebral row, keeled ventral scales, a divided

anal plate, and divided subcaudals. The female individ-

Amphib. Reptile Conserv.

ual RBINS 18504 (see Appendix 1) was encountered by

SM on 12 September 2017 in Lékédi Park. New Prov.

record and new record for the park (the species was not

listed from the Prov. by Pauwels and Vande weghe 2008;

Pauwels et al. 2016a,b, 2017e). This snake is common

in Gabon where it is found in pristine and moderately

degraded forest.

Thelotornis kirtlandii (Hallowell, 1844)

Fig. 21 shows an individual displaying the typical de-

fensive posture, photographed on 27 June 2016 at 10h00

by SM in Lékédi Park. New record for the Dept. and

for the park (not listed from the Dept. by Pauwels and

Vande weghe 2008). The species was first recorded from

Haut-Ogooué Prov. by Pauwels et al. (2007). This snake,

which avoids primary forests and lives mostly in clear-

ings and cultivated areas, is one of the most commonly

encountered species in Gabon.

Fig. 21. Adult Thelotornis kirtlandii in defensive posture in

Lékédi Park, Haut-Ogooué Prov., southeastern Gabon. Photo-

graph by S. Morelle.

Thrasops flavigularis (Hallowell, 1852)

On 20 May 2016 SM found a dead-on-road adult individ-

ual (RBINS 18479) in Moanda, Lébombi-Léyou Dept.

(Appendix 1). It showed uniformly glossy black dorsal

and ventral surfaces, except a yellowish underside of

head and throat, smooth oblique dorsal scale rows with

the vertebral row not widened. Its temporal formula is

1+1 on each side. New Dept. record (this species was not

listed from the Dept. by Pauwels and Sallé 2009; Pau-

wels et al. 2016b). The distribution and ecology of this

species in Gabon are still poorly known.

Toxicodryas pulverulenta (Fischer, 1856)

The first record of the species for Haut-Ogooué Prov.

(Christy et al. 2008) was based on the examination by

OSGP of photos of an adult individual taken in 2007 by

Paul Aczel near the camp of the Projet Protection des

Gorilles (PPG) of the Aspinall Foundation in the north-

ern part of Plateaux Batéké NP. An adult individual was

found by SM on the evening of 10 August 2017 in the

April 2019 | Volume 13 | Number 1 | e174

Pauwels et al.

kitchen of the laboratory in Lékédi Park (Fig. 22) and

behaved very aggressively. It showed 21 dorsal scale

rows at midbody. New record for the park and for the

Dept. (not listed from the Dept. by Pauwels and Vande

weghe 2008; Pauwels et al. 2017b,e). OSGP examined

two specimens (RMCA 29892-29893) collected by M.

Gauduin in 1969-1970 in ““Munana, pres de Moanda”’

[= Mounana near Moanda, in Lébombi-Léyou Dept.].

The above male and female specimens had their skulls

removed, probably for osteological studies, but their

head skin is present and generally in good condition,

their tails are complete; part of the ventral skin missing

in the second specimen. They show a widened vertebral

row. Their temporal formula is 2+2 on each side. Their

main diagnostic characters are presented in Appendix 1.

X-rays revealed a small bird in the stomach of RMCA

29893. On 22 February 2018 JLA found a dead-on-road

adult individual on the road to Amissa Hospital (H6p1-

tal Amissa), Quartier Epila, at six km from the center of

Franceville, Passa Dept. It showed (left side) one loreal,

one preocular, two postoculars, 8(3-5) supralabials, 2+2

temporals; 2 pairs of sublinguals; an orange tongue; its

total length was about 100 cm. New Dept. records. The

variation in color, pattern and dorsal scale rows numbers

within this species in Gabon requires further studies to

assess if one or more species are involved.

Fig. 22. Adult Toxicodryas pulverulenta in Lékédi Park, Haut-

Ogooué Prov., southeastern Gabon. Photograph by S. Morelle.

ELAPIDAE

Dendroaspis jamesoni jameson (Traill, 1843)

On 5 October 2016 SM examined a dead-on-road adult

individual on the R.24 road between Ndjima and Moanda

in Lébombi-Léyou Dept. The individual, very damaged,

showed the typical wide black vertebral band on a green

dorsum, and a yellow tail with each scale circled with

black. On 16 June 2017 SM encountered another dead-

on-road adult individual in Moanda, Lébombi-Léyou

Dept. Its left side showed no loreal, three preoculars, four

postoculars, eight supralabials of which the 4" contacts

the orbit, a single elongate temporal scale, and eight in-

fralabials. New Dept. record (the species was not listed

from the Dept. by Pauwels and Vande weghe 2008; Car-

Amphib. Reptile Conserv.

lino and Pauwels 2015; Pauwels et al. 2017e, 2018c).

Another dead-on-road adult individual was encountered

by SM in Franceville, Passa Dept., on 22 June 2017;

its head and tail tip were preserved (RBINS 18480); its

available meristic characters are provided in Appendix 1.

This mamba lives in primary and secondary forests and

even ventures in plantations, but is rarely observed due to

its shy and arboreal habits.

Naja annulata annulata Buchholz & Peters in Pe-

ters, 1876

An individual of this freshwater cobra was photographed

by SM on the shore of Lékédi Lake in Lékédi Park (Fig.

23). It quickly escaped to the water after having been

photographed. New record for the park and for the Dept.

(this cobra species was not listed from the Dept. by Pau-

wels and Vande weghe 2008; Pauwels et al. 2017a).

Fig. 23. Naja annulata annulata in Lékédi Lake, Haut-Ogooué

Prov., southeastern Gabon. Photograph by S. Morelle.

Naja melanoleuca Hallowell, 1857

On 1 September 2016 an Anatolian Shepherd Dog killed

an adult black and white cobra in Bakoumba, biting and

chewing its face and throat (RBINS 18481, see Appendix

1). The dog was itself bitten on the face by the snake and

quickly showed neurological and shock symptoms (hy-

persalivation, ataxia, tachycardia, and hyperpnea). The

dog was treated with Medetomidine for anaesthesia, and

three injections each of a third of a dose of FAV-Africa

serum, the first one just after the bite, then 12 hours and

24 hours later. The dog recovered and was eating again

48 hours after the bite. Another cobra was photographed

in Bakoumba by SM on 8 February 2018. New Dept. re-

cord (this cobra species was not listed from the Dept. by

Pauwels and Vande weghe 2008; Pauwels et al. 2017b).

The species was first recorded from Haut-Ogooué Prov.

by Pauwels et al. (2007) based on an individual from

Franceville. Another case of bite of a dog by this ubiq-

uitous cobra in Gabon was mentioned by Pauwels et al.

(2017b). See also above, under the account for Dasy-

peltis fasciata, and Fig. 24.

April 2019 | Volume 13 | Number 1 | e174

New reptile records from southeastern Gabon

Fig. 24. Young Naja melanoleuca swallowing an adult Dasy-

peltis fasciata in Franceville, Haut-Ogooué Prov., southeastern

Gabon. Photograph by N. Rahola.

Naja nigricollis Reinhardt, 1843

On 15 December 2016 an adult individual was photo-

graphed during the day by E. Pendrié in a savanna in Lé-

coni Park, Plateaux Dept. This is a new record for Léconi

Park, and the third record of this savanna-dwelling cobra

from Gabon (a record from Haut-Ogooué and another

from Nyanga provinces had been presented by Pauwels

and Vande weghe 2008; Pauwels et al. 2017d).

LAMPROPHIIDAE

Atractaspis reticulata heterochilus Boulenger,

1901

The female specimen RMCA 29902 was collected in

‘““Munana, pres de Moanda” [= Mounana near Moanda,

in Lébombi-Léyou Dept.] in 1969-1970 by M. Gauduin.

It has a round pupil and only one pair of sublinguals. On

each side the 2™ infralabial is fused with the sublingual.

On both sides the temporal formula is 1+2. Additional

morphological characters are provided in Appendix 1.

New Prov. record. This is only the third individual of this

rare species recorded with certainty from Gabon (Pau-

wels and Vande weghe 2008; Rodel et al. 2019).

Boaedon olivaceus (Dumeril, 1856)

An adult individual was photographed on 28 June 2016

by SM in Bakoumba (Fig. 25). It showed the typical uni-

form dark grey dorsal coloration and orange eyes with

a vertical pupil, an elongate loreal, one preocular, two

postoculars, eight supralabials with the 3" to 5" in con-

tact with the orbit, 1+2+3 temporals, and smooth dorsal

scale rows with the vertebral row not enlarged. It was

very aggressive when handled. New Prov. record (the

species was not recorded from Haut-Ogooué Prov. by

Pauwels and Vande weghe 2008; Pauwels et al. 2016a,

20178):

Boaedon perisilvestris Trape & Mediannikov,

2016

A young individual was encountered by SM at night near

buildings in Bakoumba on 23 March 2016 (Fig. 26). It

Amphib. Reptile Conserv.

Fig. 25. Adult Boaedon olivaceus in Bakoumba, Haut-Ogooué

Prov., southeastern Gabon. Photograph by S. Morelle.

Loe cs

“a4,

Fig. 26. Young Boaedon perisilvestris in Bakoumba, Haut-

Ogooué Prov., southeastern Gabon. Photograph by S. Morelle.

was aggressive and bit when handled. It showed a verti-

cal pupil, one preocular, in contact with the frontal, two

postoculars, one loreal, eight supralabials with three (3"—

5") in contact with the orbit, one anterior temporal, 29

smooth dorsal scale rows at midbody, the vertebral row

not widened, a single anal plate, divided subcaudals, two

poorly contrasted stripes on each side of the head, and a

uniform blackish dorsum without light stripes. An adult

female individual showing the same head scalation char-

acteristics, but no stripes on the sides of the head (as is

typical for adults, according to Trape and Mediannikov

2016) was found on 26 December 2016 inside a dried

well in Lékédi Park. It was resting near the six oblong,

white eggs it had just laid. The adult female RBINS 18505

(see Appendix 1) was found by SM in Lékédi Park on 18

November 2016. It contains six large eggs, the largest

with a length of 32 mm and a maximum width of 12 mm;

its stomach contains the remains of a micromammal. The

young individual RBINS 18482 (see Appendix 1) was

found in Moanda, Lébombi-Léyou Dept. on 20 Novem-

ber 2016 at 7h00. Its temporal formula is 1+2+3 on each

side. Another individual (RBINS 18506; see Appendix

1) was found by SM in Moanda on 5 November 2016;

it also shows a temporal formula of 1+2+3 on each side.

The adult female RMCA 29885 (see Appendix 1) was

collected in ““Munana, pres de Moanda” [= Mounana

near Moanda, in Lébombi-Léyou Dept.] in 1969-1970

April 2019 | Volume 13 | Number 1 | e174

Pauwels et al.

by M. Gauduin. It shows vertical pupils. New record for

both Depts. and for the park. Within Gabon, this recently

described species was known only from Franceville in

Passa Dept. (Pauwels et al. 2017e).

Mehelya poensis (Smith, 1847)

SM encountered an adult individual (RBINS 18483) on

17 April 2016 in Bakoumba. It showed two pairs of sub-

linguals; a strongly widened vertebral row with a double

keel, and all dorsal scales keeled, without secondary

keels; no white spots on dorsals (see other diagnostic

characters in Appendix 1). On each side its temporal for-

mula is 1+2, but on the left side, the anterior temporal is

separated from the postocular by a slight contact between

the parietal and the 5" supralabial. When handled, the

snake never attempted to bite. New Prov. record (the spe-

cles was not recorded from Haut-Ogooué Prov. by Pau-

wels and Vande weghe 2008; Pauwels et al. 2016a).

Psammophis cf. phillipsii (Hallowell, 1844)

On 5 July 2017 JLA found a dead-on-road adult indi-

vidual on the road to Amissa Hospital, at six km from

the center of Franceville, Passa Dept. Its total length was

about 105 cm (including the incomplete tail). A second

individual was found dead-on-road in the same local-

ity on 13 July 2017 (Fig. 27); its total length was about

110 cm. Both showed a round pupil, two internasals, two

prefrontals, eight supralabials of which the 4" and 5" in

contact with orbit, one large preocular, two postoculars,

an elongate loreal, two anterior temporals, 17 smooth

and oblique dorsal scale rows at midbody (vertebral row

not widened), divided anal and subcaudals, and an olive

brown dorsum with a black spot in the posterior part of

each ventral; the dorsal surface of the head 1s olive brown

with irregular black spots. On 11 August 2017 SM found

a dead-on-road individual in Moanda, Lébombi-Léyou

Dept. It showed the same head scalation features as the

two specimens above, and the same coloration. New

Prov. record (not recorded from the Prov. by Pauwels and

Vande weghe 2008; Pauwels et al. 201 6b, 2017a).

y *

Fig. 27. Dead-on-road adult Psammophis cf. phillipsii in

Franceville, Haut-Ogooué Prov., southeastern Gabon. Photo-

graph by J.-L. Albert.

Amphib. Reptile Conserv.

NATRICIDAE

Natriciteres olivacea (Peters, 1854)

An adult female (RBINS 18484, see Appendix 1 and Fig.

28) was encountered by SM at midday on 10 October

2016 in Lékédi Park. It contains four large eggs between

the ventrals 93 and 127, each of a length of about 17 mm.

It shows a round pupil and a temporal formula of 1+2 on

each side. In life the dorsum of this individual was nearly

uniformly black, and the mediodorsal band that is char-

acteristic of this species was nearly invisible. New record

for the park and new Dept. record (not recorded from the

Dept. by Pauwels and Vande weghe 2008; Pauwels et al.

2017a). In Gabon this snake is much less common than

its congener Natriciteres fuliginoides (Gunther, 1858)

which was however not found during our survey; both

were evaluated as Least Concern by the IUCN.

Fig. 28. Adult female Natriciteres olivacea (RBINS 18484) in

Lékédi Park, Haut-Ogooué Prov., southeastern Gabon. Photo-

graph by S. Morelle.

PYTHONIDAE

Python sebae (Gmelin, 1789)

The record of the species from Plateaux Batéké NP by

Christy et al. (2008) was based on a pers. comm. to

OSGP by Gaél Vande weghe who observed an individual

along Mpassa River in the NP in April 2007, and on a

picture of a large adult individual shown in Pearson et

al. (2007). On 15 January 2018 at 13h00 SM discovered

a juvenile in Lékédi Park (Fig. 29). We examined pho-

Fig. 29. Juvenile Python sebae in Lékédi Park, Haut-Ogooué

Prov., southeastern Gabon. Photograph by S. Morelle.

April 2019 | Volume 13 | Number 1 | e174

New reptile records from southeastern Gabon

tographs taken on 16 July 2017 by A.N. Eyeang Mba of

an adult individual of about four meters long that had

been shot by a hunter the previous night in Moanda,

Lébombi-Léyou Dept. New record for the park and new

Dept. records (not recorded from these Depts. by Moc-

quard 1887; Pauwels and Vande weghe 2008; Pauwels

et al. 2016b, 2017e, 2018c). In spite of a heavy hunting

pressure, this python is still common in Gabon, includ-

ing in urban areas, and is known from twelve Gabonese

protected areas (Pauwels, 2016).

TYPHLOPIDAE

Letheobia caeca (Dumeéril, 1856)

On 15 March 2017 at 14h00 SM found an individual in

Bakoumba (RBINS 18507). It shows a uniformly pink-

ish color; a snout-vent length of 260 mm, a tail length

of 4.7 mm, a midbody diameter of 3.8 mm, a ratio total

length/midbody diameter of 69.7; 22-22-22 scale rows,

all smooth; a rostral with parallel sides; a longitudinally

enlarged frontal; long supranasals; and a T-0 supralabial

imbrication pattern. Its eyes are invisible. New Prov. re-

cord (this fossorial species was not recorded from Haut-

Ogooué Prov. by Wallach 2005; Pauwels and Vande

weghe 2008). This snake is possibly widespread in the

country, but rarely observed due to its subterranean hab-

Its.

VIPERIDAE

Atheris squamigera (Hallowell, 1856)

Pearson et al. (2007) presented a picture of a light green

Atheris with well-spaced yellowish transversal rings from

Plateaux Batéké NP, that they identified as a “bush viper

(possibly Atheris squamigera or A. chlorechis).” On 17

September 2016 SM photographed an individual in Bak-

oumba (Fig. 30). New Dept. record (this arboreal viper

species was not recorded from the Dept. by Pauwels and

Vande weghe 2008; Pauwels et al. 2016b, 2017d,e).

Fig. 30. Atheris squamigera in Bakoumba, Haut-Ogooué Prov.,

southeastern Gabon. Photograph by S. Morelle.

Bitis arietans (Merrem, 1820)

An individual was photographed by A. Willaume on

22 March 2015 at 12h33 in a savanna in Lékédi Park.

Amphib. Reptile Conserv.

It showed the typical dorsal chevron pattern, the beige

interorbital band and two beige spots on the back of the

dorsal surface of the head. The weather was sunny and

the viper was aggressive when approached. New record

for the park and for the Dept. This savanna-dwelling vi-

per is known in Gabon only from a handful of localities

in Haut-Ogooué and Nyanga provinces (Pauwels et al.

2012, 2017d, 2018a,b).

Bitis gabonica (Dumeril, Bibron & Dumeéril, 1854)

Pearson et al. (2007) mentioned Bitis gabonica from Pla-

teaux Batéké NP, but they did not provide a photo nor the

reference to a voucher specimen. The species was con-

firmed from Haut-Ogooué Prov. by Pauwels and Sallé

(2009) based on an individual from Franceville in Passa

Dept. On 22 April 2016 SM photographed a young indi-

vidual in Lékédi Park (Fig. 31). The snake was found at

11h15 in a forest within a large enclosure where a group

of habituated mandrills (Cercopithecidae: Mandrillus

sphinx) lives in semi-freedom (Brockmeyer et al. 2015),

and was signalled by their alert calls. New record for the

park and for the Dept.

Fig. 31. Young Bitis gabonica in Lékédi Park, Haut-Ogooué

Prov., southeastern Gabon. Photograph by S. Morelle.

Bitis nasicornis (Shaw, 1802)

An adult individual collected a few years ago in Lékédi

Park is kept in the park’s vivarium. Another adult individ-

ual of this common viper was encountered by SM in the

park at midday on 30 April 2016; its head was preserved

(RBINS 18485; see Appendix 1) and shows 18/19 circu-

mocular scales (the uppermost the largest); 14 scales sep-

arate the orbits. The juvenile RBINS 18508 was found by

SM crossing a road in Bakoumba on 11 October 2016 at

9h30. It shows well-developed nasal horns; 18/18 circu-

mocular scales; its umbilical scar extends from its ven-

trals 111 to 114 included; 15 scales separate its orbits (see

also Appendix 1). New Dept. record and new record for

the park (this viper species was not listed from the Dept.

by Christy et al. 2008; Pauwels and Vande weghe 2008;

Pauwels et al. 2016b, 2017d).

April 2019 | Volume 13 | Number 1 | e174

Pauwels et al.

Causus lichtensteinii (Jan, 1859)

On 19 June 2017 at 11h00 SM encountered a dead-on-

road individual in a forest road in Lékédi Park (RBINS

18486; see Appendix 1). Its pupil is round. The dorsal

surface of the head is grey. The dorsum is dark olive

green with regularly-spaced black chevrons. New re-

cord for the park and for the Prov. (the species was not

recorded from the Prov. by Pauwels and Vande weghe

2008; Pauwels et al. 2016b, 2017b). The distribution and

ecology of this forest-dwelling viper in Gabon are still

poorly documented.

Causus maculatus (Hallowell, 1842)

In 2017 an adult individual was photographed by R. Pacta

in Moanda, Lébombi-Léyou Dept. (Fig. 32). New Dept.

record (the species was not recorded from the Dept. by

Pauwels and Vande weghe 2008; Pauwels et al. 2017b,

2018c). In his unpublished Masters’ thesis on Cau-

sus, de Massary (1993) listed two specimens (MNHN

1886.233—234) from “Franceville, Gabon’, and three

more (MNHN 1980.1100—1102) from “‘Mounana, 80 km

de Francev. [= Franceville], Gabon,” thus in Lébombi-

Léyou Dept. This savanna-dwelling viper seems rela-

tively localized in Gabon.

se hy

Prov., southeastern Gabon. Photograph by R. Pacta.

Discussion

The first reptile records for Plateaux Batéké NP, the only

national park in Haut-Ogooué Prov., were made by Pear-

son et al. in 2007. Its herpetofauna has never been sys-

tematically surveyed, but a compilation of opportunistic

observations allowed Pauwels (2016) to provide a pre-

liminary list of 11 species, obviously extremely incom-

plete. Not a single herpetological record currently exists

for the only Ramsar Site in the Prov., the Rapides de

Mboungou Badouma et de Doumé site (Pauwels 2016).

In this context, records from sites such as the Lékédi sa-

fari park, where the savanna-forest mosaic is well rep-

resented (Peignot et al. 2008; Brockmeyer et al. 2015)

and benefits from a certain level of protection, are im-

portant both for the conservation of these species and for

Amphib. Reptile Conserv.

the development of local ecotourism. Zoogeographically,

Lékédi Park and its surroundings belong to the Chaillu

Massif (Massif du Chaillu), and our new records from

the park, Bakoumba and Lémanassa add to the already

rich herpetofaunal list gathered for the massif (Pauwels

et al. 2002, 2008, 2016a,b, 2017b,d; Carlino and Pauwels

2014; Dewynter et al. 2018).

The total number of reptile species currently known

from Lékédi Park is 33. This 1s obviously an incomplete

list, as it is the case for all protected areas of Gabon, but

for comparison this is much more than the current totals

respectively known for the Arboretum Raponda Walk-

er (18 species), Akanda NP (16), Plateaux Batéké NP

(16), Mayumba NP (16), Minkébé NP (17), Pongara NP

(25), Mwagna NP (1), Waka NP (2), or Wonga-Wongué

Presidential Reserve (26) (Pauwels 2016; Pauwels et al.

2017a,b,c,d, 2018b). Seven of the 28 new species records

for Lékédi Park also represent new provincial records.

The discovery in Lékédi Park of a new population of

Mecistops cataphractus is of particular importance for

the conservation of this crocodile, evaluated as Critically

Endangered by the IUCN (Shirley 2014). It is to be noted

that another lake, Missombo Lake (Lac Missombo), ex-

ists in Lékédi Park, which has never been herpetologi-

cally surveyed, and seems to be ecologically appropriate

to house a population of M. cataphractus. Searches for

crocodiles in the lake will likely also reveal the presence

of more aquatic species such as trionychids (Cycloderma

aubryi and Trionyx triunguis), more pelomedusids (Pelu-

sios spp.) and water snakes of the genus Hydraethiops.

Our finding of Osteolaemus tetraspis in Lékédi Park is

also good news for the conservation of this small croco-

dile species evaluated as Vulnerable by the IUCN. The

first records for Haut-Ogooué Prov. of three venomous

snakes (Atractaspis reticulata heterochilus, Causus lich-

tensteinii, and Psammophis cf. phillipsii) are of medical

importance. Including our new data, all six viperid spe-

cies currently known from Gabon are presently recorded

from Haut-Ogooué Prov. in addition to, for instance, all

three Naja species. Although not yet documented, Gold-

ie’s Tree Cobra Pseudohaje goldii and more burrowing

asps of the genus Atractaspis are most certainly present

in Haut-Ogooué Prov., based on their respective global

distributions.

With the recent additions -or confirmations of occur-

rence- of the gekkonids Hemidactylus echinus by Car-

lino and Pauwels (2015) and Lygodactylus conraui by

Pauwels et al. (2016a), the lacertid [chnotropis bivittata

by Ineich and Le Garff (2015), the colubrid 7hrasops

Jacksonii by Carlino and Pauwels (2013), the lamprophi-

id Polemon gracilis by Pauwels et al. (2018c), and the

present confirmation of Philothamnus hughesi, the rep-

tile fauna of Gabon is now known to include 130 docu-

mented species (the record of Natriciteres variegata by

Hughes in 2017 proved to be based on a misidentified

specimen of N. fuliginoides, see Pauwels et al. 2017c;

the record of Trachylepis makolowodei by GvoZzdik et al.

156 April 2019 | Volume 13 | Number 1 | e174

New reptile records from southeastern Gabon

in 2018 was based on a misidentified 7. albilabris, see

Pauwels et al. 2018a). This number should still increase

with further field research, especially along the borders

with Cameroon and the Republic of the Congo.

Acknowledgements.—We are grateful to Daniel

Franck Idiata and Aurélie Flore Koumba Pambo (CENA-

REST, Libreville) who facilitated the research permit for

the MSNS (permit n° AROO50/17/MESRS/CENAREST/

CG/CST/CSAR) and to Antonio Durante (MSNS) for

providing working facilities. We thank Eric Willaume

(Sodepal) for logistic support, Agathe Nicaise Eyeang

Mba (COMILOG, Moanda), Raphaél Pacta (Moanda),

Estelle Pendrié (Moanda), and Aurélie Willaume for

letting us use their photographs, and Paul Aczel (John

Aspinall Foundation), Bernard Le Garff (Université de

Rennes I), Nicolas Longin (COMILOG, Moanda), and

Jean Pierre Vande weghe (Kigali) for useful information.

Danny Meirte gave OSGP access to the RMCA herpe-

tological collections. Garin Cael (RMCA) X-rayed a

Toxicodryas specimen. SM thanks the Ecole des Mines

et de la Métallurgie de Moanda (Moanda) for working

facilities, Noémie De Botton Martin, Kevin Morelle,

Max-Anaclet Makaba, Clauther Lepoko, Joel Tsamouna,

Cédric Moudengue, Stanislas Lekosso, and Maick Mous-

sodji for their assistance in the field, Daniel Potier and

Geoffrey Aseglio who taught him how to handle reptiles,

and his family and most particularly his grandfather

Jean-Pierre Nicot who encouraged his passion for rep-

tiles. We thank Werner Conradie and the late Bill Branch

for editorial support.

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375-383. Cameroon & Gabon, with comments on Agama me-

Vande weghe J-P. 2008. Les Parcs Nationaux du Gabon. helyi Tornier, 1902. Bonner zoologische Beitrdge

Plateaux Batéké. Wildlife Conservation Society, Li- 56(4): 285-297.

breville, Gabon. 272 p. Wallach V. 2005. Letheobia pauwelsi, a new species of

Amphib. Reptile Conserv.

blindsnake from Gabon (Serpentes: Typhlopidae). Af

rican Journal of Herpetology 54(1): 85-91.

Olivier S.G. Pauwels, born in 1971 in Belgium and graduated in Zoology from the Free University of

Brussels (ULB). After having managed environmental programs for a dozen years in Gabon, then for

several years in Kazakhstan, he became the permanent curator of the recent vertebrate collections of

the Royal Belgian Institute of Natural Sciences. A conservationist and taxonomist, Olivier co-authored

about 300 papers and books on the biodiversity of the Old World, and co-described about 100 new taxa

from tropical Africa and Asia, mostly reptiles, but also amphibians and some insects.

Stephan Morelle, born in 1989 in Germany, graduated as a doctor in veterinary medicine from the

University of Liege in Belgium. Interested in wildlife medicine, he followed courses on chemical and

physical restraint of wild animals in Zimbabwe. He worked for two years as a veterinarian in a great

apes sanctuary in Gabon, and took the opportunity to be based in Gabon to photograph all reptiles and

amphibians he encountered. Stephan is currently working in France in a clinic for small animals.

Jean-Louis Albert was born in France in 1951. Since a young age he has had a strong interest in

Nature, especially entomology. He has worked as a biomedical and technical manager in the tropics

since 1973. He spent seven years in Gabon in the 70’s, then three years in Guatemala, 13 years in

Congo Brazzaville, and eventually came back in 1995 to Gabon where he is still based, in Haut-

Ogooue Province. He spends all his free time in Gabon exploring all provinces of the country and

photographing and documenting its fauna and flora, with an emphasis on butterflies, reptiles, and birds

(see http://www.jeanlou.fr/).

Piero Carlino was born in 1978 in Italy. He is currently curator of the herpetology department of the

Natural History Museum of Salento, and is involved in various projects on biodiversity conservation.

His studies focus mainly on the systematics, taxonomy, biogeography, and ecology of the Palaearctic

and Afrotropical herpetofaunas. He organizes and participates in numerous expeditions in Europe, the

Middle East, and tropical Africa to evaluate the conservation status of various animal species, and is

the manager of the Salento sea turtles rescue center.

Nil Rahola, born in 1983 in France, was passionate about entomology from his earliest age. In 2008 he

began work at the IRD (Research Institute for Development in Montpellier, France). He made his first

field trips to Gabon in 2010 and lived there from 2012 to 2018. He is a medical entomologist interested

in taxonomy and systematics of arthropods of medical and veterinary importance, with emphasis on

mosquitoes and sand flies. He has discovered and described several new species and is a herpetology

and nature photography enthusiast.

Jean-Francois Trape is a French medical doctor, biologist, and herpetologist with lengthy experience

in Africa, where he was born in 1949. Since 1980 he has worked continuously in Central and West

Africa for the Institut de Recherche pour le Développement (IRD, formerly ORSTOM), a French public

institution for research in Southern countries. Jean-Francois has authored or co-authored over 300

peer-reviewed papers and books on tropical medicine and herpetology, including the books “Guide des

serpents d’ Afrique occidentale. Savane et desert” (2006) and “Lézards, crocodiles et tortues d’ Afrique

occidentale et du Sahara” (2012), and has described 35 reptile and five tick species. Jean-Francois

is also a malaria expert for the World Health Organization, where he has served in several steering

committees. In 2010 he received the first IRD prize for research, and in 2013 the Lucien Tartois prize

from the French Foundation for Medical Research.

159 April 2019 | Volume 13 | Number 1 | e174

Pauwels et al.

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April 2019 | Volume 13 | Number 1 | e174

161

Amphib. Reptile Conserv.

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

13(1) [General Section]: 162-172 (e175).

Annotated checklist of the herpetofauna

(Amphibia, Reptilia) of Mount Ararat and surroundings

John Mulder

Natural History Museum Rotterdam, Department of Vertebrates, Westzeedijk 345, 3015AA, Rotterdam, the NETHERLANDS

Abstract.—Presented is a comprehensive overview of data on the herpetological species occurring in the

Ararat region of Turkey. A total of 41 species, including their first records, published historical localities, and

respective sources, are recorded in this assessment of surprisingly high regional species richness.

Keywords. Zoogeography, Turkey, species richness, reptiles, amphibians, survey

Citation: Mulder J. 2019. Annotated checklist of the herpetofauna (Amphibia, Reptilia) of Mount Ararat and surroundings. Amphibian & Reptile

Conservation 13(1) [General Section]: 162-172 (e175).

ternational (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in any medium,

provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced, are as follows:

Official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org.

Received: 19 October 2017; Accepted: 18 December 2018; Published: 1 May 2019

Introduction

Mount Ararat, Turkey’s highest mountain peak, reaches

an elevation of 5,137 ma.s.l. The Ararat massif is about

40 km in diameter and is of volcanic origin. As the pre-

sumed resting place of Noah’s Ark, Mount Ararat, or

Buytkk Agri Dag1, as the mountain is called in Turkish,

has for some people reached mythical dimensions. The

region has a very rich diversity of flora and fauna. It is

positioned on Turkey’s northeasternmost national fron-

tier and is bordered by Armenia, Nakhchivan, and Iran.

The mountain and its surroundings have been visited

by several zoological expeditions over the past century.

Early herpetological records were collected by Boettger

(1892), Méhely (1894), Basoglu (1945), and Clark and

Clark (1973). Since the 1980s, publications from explora-

tions followed more rapidly, for instance Flardh (1983),

Teynié (1987, 1991), Mulder (1995), Garzoni and Ge-

niez (2004), Ilgaz et al. (2005), Baran et al. (2005), and

Tosuno$lu et al. (2010).

The principal objective of this survey 1s to provide a

checklist of the amphibians and reptiles known to inhabit

Mount Ararat and surroundings.

Materials and Methods

The basis of this survey is formed by several personal her-

petological observations recorded during multiple explo-

rations in the region. Most of these observations, including

some first records for the area, have been published previ-

ously (Mulder 1995). However, in an attempt to assemble

Correspondence. contact@ecologischadviesbureau.nl

Amphib. Reptile Conserv.

a comprehensive species list for the area, these are supple-

mented by a full literature and online source search.

The area under study is the entire province (il) of I&dir

(i.e., the communities Tuzluca, [&dir, Karakoyunlu, and

Aralik) and the community (ilce) of DoSubayazit in the

province of Agri, as shown in Fig. 1. Mount Ararat is situ-

ated in both provinces. The area under study also includes

the north face of Mount Tendurek, another high mountain

(3,533 m). The locality ‘Tuzluca north’ in Teynié (1991) is

provisionally considered to be located within the borders

of the province of Igdir, though it was not possible to ob-

tain further information from the author.

Scientific species names have been adapted to the most

recent taxonomical and systematic reviews, superseding

the names used in the original publications. In cases of

indistinct derivation of a species in a source an explana-

tion is given. The names used follow The Reptile Database

(www.reptile-database.org).

It was decided to refrain from providing available alti-

tudinal data here, since those data can be easily found in

the original publications.

Results

Currently 41 species are recognized for the area under

study, including five amphibian and 36 reptile species.

All recorded amphibians are anurans. Within the class

of reptiles two species are chelonians, 15 are lizards and

19 are snakes. Table 1 presents an overview of all recog-

nized species, year of first record, and respective source

citations. More comprehensive data are presented in the

checklist that follows.

May 2019 | Volume 13 | Number 1 | e175

Mulder

Table 1. Recognized species within the area under study with references of first records.

' Not mentioned, given here is the year of publication;

? Only a few days prior to Teynié (1987),

> If ‘north of Tuzluca’ is situated within the area under study.

Species Year of recording Author

Bufotes variabilis 1980 Flardh 1983

Pelophylax ridibundus 1987 Mulder in this paper

Pelobates syriacus 1988 Franzen and Sigg 1989

Rana macrocnemis 2011 AdaMerOs 2014

Hyla savignyi 199] Mulder 1995

Mauremys caspica 1893 Méhely 1894

Testudo graeca 1967 Clark and Clark 1973

Phrynocephalus helioscopus 1893 Méhely 1894

Paralaudakia caucasia 1967 Clark & Clark 1973

Mediodactylus kotschyi 1980 Flardh 1983

Darevskia bendimahiensis 1982 Schmidtler et al. 1994

Darevskia raddei 1980 Flardh 1983

Darevskia valentini 1994! Schmidtler et al. 1994

Eremias pleskei 1943 Basoglu 1945

Eremias strauchi i952! Mertens 1952

Eremias suphani 1967 Clark and Clark 1973

Lacerta media 1967 Clark and Clark 1973

Lacerta strigata 1969 Baran 1980

Ophisops elegans 1987 Baran 1980

Heremites septemtaeniata 1987? Mulder 1995

Eumeces schneideri foot? Teynié 1991

Pseudopus apodus 1969 Baran 1977a

Xerotyphlops vermicularis 1987 Mulder 1995

Eryx jaculus 1988 Franzen and Sigg 1989

Natrix natrix 1896 Boettger 1899

Natrix tessellata 1896 Boettger 1899

Dolichophis schmidti 1942 Mertens 1952

Eirenis collaris 1988 Franzen and Sigg 1989

Eirenis modestus 1980! Basoélu and Baran 1980

Eirenis punctatolineatus 1991! Teynié 19913

Elaphe dione 1988 Garzoni and Geniez 2004

Elaphe sauromates 1896 Boettger 1899

Hemorrhois ravergieri 1940 Mertens 1952

Platyceps najadum 1987 Mulder 1995

Telescopus fallax 1987-1991 Teynié 1991?

Zamenis longissimus 1993 Schweiger 1994

Zamenis hohenackeri 1896 Boettger 1899

Malpolon insignitus 1967 Clark and Clark 1973

Macrovipera lebetina 1942 Mertens 1952

Montivipera raddei 1888 Boettger 1890

Vipera eriwanensis 1896 Boettger 1899

Amphib. Reptile Conserv. 163 May 2019 | Volume 13 | Number 1 | e175

Herpetofauna of Mount Ararat

43.2 43.3 43.4 43.5 43.6 43.7 43.8 43.9 44.0 44.1 442 443 44.4

445 446 44.7 44.8 44.9 45.0

Fig. 1. The region under study, comprising the province of I&dir and the community of DoSubayazit in the province of Agri.

Checklist

Class Amphibia; Order Anura; Family Bufonidae

Bufotes variabilis (Pallas, 1769) Variable Toad

This taxon has a complex nomenclatural history. It had

long been regarded as a synonym of the taxon Bufo viri-

dis Laurenti 1768 and allocated to the genus Bufo, but

was then briefly assigned to Pseudepidalea before Bu-

fotes was chosen for this group (Frost et al. 2006; Dubois

and Bour 2010; Frost 2013). The records from the area

under study were undoubtedly recognized as belonging

to this taxon, irrespective of the names used.

Flardh (1983): west slope of Ararat, 1980; Franzen

and Sigg (1989): surroundings of Dogubayazit, 1988;

Norstrom (1990): west slope of Ararat and [dir plain;

Mulder (1995): northwest slope of Ararat, 1987; Franzen

and Heckes (1999): northeast of Karabulak (Agri) and

south of [&dir, 1990; Fonters and Fonters (web): [dir

plain at the height of Suveren; Tosuno$lu et al. (2010):

Tasburun, 2009; AdaMerOs (web): Aralik, 2012; AdaM-

erOs (web): Yukari Cinikli k6yt/Tuzluca, 2012.

Family Ranidae

Pelophylax ridibundus (Pallas, 1771) Marsh Frog

Amphib. Reptile Conserv.

Mulder (not previously published): Karabulak, prov-

ince of Aén, 1987; Franzen and Sigg (1989): surround-

ings of Dogubayazit, 1988; Schneider and Schneider (in

litt): Tasburun, 2008; Tosunoglu et al. (2010): Tasburun,

2009; AdaMerOs (web): Tuzluca, 2013.

Pelobates syriacus Boettger, 1889 Eastern Spadefoot

Franzen and Sigg (1989): surroundings of Dogubayazit,

1988; Franzen and Heckes (1999): northeast of Karabu-

lak (Agri) and south of [&dir, 1990; Trapp (web): Ara-

rat; AdaMerOs (web): Yukari Cigrikli koyt, province of

Isdir (2011); AdaMerOs (web): Aralik, 2012.

Rana macrocnemis Boulenger, 1885 Caucasus Frog

AdaMerOs (web): Yukari Cryrikli mahallesi, Tuzluca,

province of [&dir (2011).

Family Hylidae

Hyla savignyi Audouin, 1827 Middle East Tree Frog

Mulder (1995): Aralik, 1991; Schneider and Schneider

(in litt): I&dir, 2009; Tosunoglu et al. (2010): Tasburun,

province of Igdir, 2009; AdaMerOs (web): Tuzluca,

2013.

May 2019 | Volume 13 | Number 1 | e175

Mulder

Class Reptilia; Order Testudines; Family Geoemydi-

dae

Mauremys caspica (Gmelin, 1774) Caspian Turtle

Méhely (1894): Aralik, 1893; Eiselt and Spitzenberger

(1967): Aralik, 1893 (Méhely’s data); Basoglu and Ba-

ran (1977): Aralik, 1893 (Méhely’s data); Schneider

and Schneider (in litt): AsaSi¢amurlu north of Ararat

(39°55’33”N 44°24’39”E), 2008, 2009, 2014; Tosunoglu

et al. (2010): Tasburun, 2009.

Family Testudinidae

Testudo graeca Linnaeus, 1758 Spur-thighed Tortoise

Clark and Clark (1973): ISdir, 1967; Baran (1980):

Tasburun Koyti ISdir, 1977; Basoglu and Baran (1977):

Dogubayazit (Aér1); Norstrom (1990): I&dir plain; Mul-

der (1995): northwest slope of Ararat, province of [&dir,

1987; Baran et al. (2004): around Aralik, 2000; Schneider

and Schneider (in litt): six km south of I&dir (39°51710”°N

44°4°13”E) and Aralik (39°50°10"N 44°31’21”E)

and Tasburun (39°57°11”N 44°13°40”E), 2008-2014;

Tosuno§lu et al. (2010): Tasburun, 2008; Tosuno$lu et al.

(2011): between Karakoyunlu and Tasburun, I&dir, 2008.

Order Squamata; Suborder Lacertilia;

Agamidae

Family

Phrynocephalus helioscopus (Pallas, 1771) Sunwatcher

Toadhead Agama

The distribution of this species in Turkey is confined al-

most entirely within the area treated here and the plains

around the foot of Mount Ararat. These plains are cur-

rently under threat due to development activities. Only

Mulder (1995) mentioned its occurrence along the river

Aras near Tuzluca.

Méhely (1894): Aralik, 1893; Mertens (1952):

“Baskoyt” (=Aralik), 1942; Clark and Clark (1973):

three km west of Dogubayazit, two km north of

Dogubayazit, 10 km south of ISdir, 1967; Baran (1980):

Devlet Uretme Ciftlidi (State farm), 1969 and Tasburun

Koyt [gdir, 1977; Daszak and Cawthraw (1991): two

km east of DoSubayazit, 1989; Mulder (1995): east of

Tuzluca, 1991 and 1992; Franzen and Heckes (1999):

northeast of Karabulak (A&r1) and south of [gdir, 1990;

Baran et al. (2004): six km southwest of Aralik, 2000;

Schneider and Schneider (in litt): Aralik (39°50’10°N

44°31°21”E), 2008, 2009, 2011, 2014; Gogcmen (web):

Aralik, 2011; Tosunoglu et al. (2011): between Kara-

koyunlu and Tasburun, [&dir, 2008; Cicek et al. (2012):

Aralik.

Paralaudakia caucasia (Eichwald, 1831) Caucasian

Agama

Amphib. Reptile Conserv.

Clark and Clark (1973): 10 and 25 km south of [&dir

and 20 km north of Dogubayazit, 1967; Baran (1980):

Tasburun Koyti [gdir, 1977; Baran et al. (1989): north

slope of Tendurek between Dogubayazit and the Ziyaret

pass, 1987 and Ishak Pasa Sarayi1, 1969; Flardh (1983):

west slope of Ararat, 1980; Norstrom (1990): west slope

of Ararat; Mulder (1995): Ararat northwest (Igdir), 1987;

Teynié (1987): Ararat southwest (province of Agri) and

north of Tuzluca, 1987; Franzen and Sigg (1989): sur-

roundings of Dogubayazit, 1988; Daszak and Cawthraw

(1991): three km east of DoSubayazit, 1989; Franzen

and Heckes (1999): northeast of Karabulak (Agri), 1990;

Schneider and Schneider (in litt): six km south of [&dir

(39°50°10"N 44°31’21”E), 2008, 2009, 2011, 2012,

2014.

Family Gekkonidae

Mediodactylus kotschyi (Steindachner, 1870) Kotschy’s

Gecko

This gecko has been found in the region by just one au-

thor and urgently needs confirmation by new observa-

tions.

Flardh (1983): west slope of Ararat, 1980.

Family Lacertidae

Darevskia bendimahiensis (Schmidtler, Eiselt, and

Darevsky, 1994)

Schmidtler et al. (1994): DoSubayazit, southwest Ararat,

province of Aér1.

Darevskia raddei (Boettger, 1892) Radde’s Lizard

Flardh (1983): west slope of Ararat, 1980; Mulder (not

previously published): west slope of Ararat, province

of ISdir, 1987; Norstrom (1990): west slope of Ararat;

Schmidtler et al. (1994): Dogubayazit; Trapp (web): Ara-

rat; Panner (web): Ararat, 2005, 2010.

Darevskia valentini (Boettger, 1892) Valentin’s Lizard

Schmidtler et al. (1994): On the way up to Mount Ararat

(north of DoSubayazit), province of Agri.

Eremias pleskei Nikolsky, 1905 Pleske’s Racerunner

The distribution of this species in Turkey is almost en-

tirely confined to the focal area treated here. First record

was by Basoglu (1945), namely four females caught in

1943 at Baskoy (=Aralik), province of [&dir.

Clark and Clark (1973): 10 km south of I&dir, 1967;

Baran (1980): Tasburun Koyiti Igdir, 1977, Devlet Uret-

me CiftliSi (State farm), province of Igdir, 1969; Nor-

strom (1990): I&dir plain; Franzen and Heckes (1999):

May 2019 | Volume 13 | Number 1 | e175

Herpetofauna of Mount Ararat

northeast of Karabulak (Aér1) and south of [gdir, 1990;

Baran et al. (2004): Gédekli (nine km SE of Aralik) and

five km SW of Aralik, 2000; Tosunoglu et al. (2011):

between Karakoyunlu and Tasburun, [&dir, 2008; AdaM-

erOs (web): Aralik, 2012.

Eremias strauchi Kessler, 1878 Strauch’s Racerunner

The distribution of this species in Turkey is entirely re-

stricted to the focal area under consideration here and

encompasses the plains around the foot of Mount Ararat

and along the river Aras. The locality “east of Karakurt”

in the Aras valley mentioned by Mulder (1995) is the

westernmost finding of the species in Turkey.

Mertens (1952): Tuzluca; Baran (1980): Devlet Uret-

me CiftliSi (State farm), province of I&dir, 1969 and Cilli

Gecidi, DoSubayazit, 1975 and Tasburun Koyu [édir,

1977; Clark and Clark (1973): 10 km south of I&dir, 1967;

Norstrom (1990): plain of I&dir; Mulder (1995): plain

south of I&dir, 1990 and east of Karakurt, 1989; AdaM-

erOs (web): Dogubayazit, 2011; Franzen and Heckes

(1999): northeast of Karabulak (Agri), 1990 and 25 km

northwest of I&dir and south of [&dir; Baran et al. (2004):

Godekli (nine km southeast of Aralik), Torulpasa Kisasi,

Aralik and five km southwest of Aralik, 2000; AdaMerOs

(web): Tuzluca, 2011; Tosunoglu et al. (2011): between

Karakoyunlu and Tasburun, [&dir, 2008.

Eremias suphani Basoglu and Hellmich, 1968 Suphan

Racerunner

The Suphan Racerunner has long been seen as a Turk-

ish endemic species. Mulder (1995) found the species in

1988 a few kilometres from the Iranian border, so an oc-

currence in Iran was predictable. Rastegar-Pouyani et al.

(2013) proved the occurrence on the Iranian side of the

border.

Franzen and Heckes were the first to mention the oc-

currence of this species within the region under study,

viz. near Dogubayazit in 1999, though previously Clark

and Clark had already found them in 1967 but published

under the taxon strauchi.

Clark and Clark (1973): two km north of DoSubayazit,

1967; Franzen and Heckes (1999): near Ishak Pasa

Saray1, Dogubayazit, 1988, 1990.

Lacerta media Lantz and Cyrén, 1920 Eastern Emerald

Lizard

Baran et al. (2004) used the name Lacerta trilineata tri-

lineata for a specimen from the Aras valley, probably in-

dicating this species.

Clark and Clark (1973): 25 km south of ISdir, 1967;

Flardh (1983): west slope of Ararat, 1980; Norstrom

(1990): west slope of Ararat; Mulder (1995): Cetenli

(north slope of Tendiirek), 1993; Schneider and Schnei-

der (in litt): northwest slope of Ararat (39°47’21”°N

Amphib. Reptile Conserv.

44°11°35”E), 2009; Trapp (web): Ararat.

Lacerta strigata Eichwald, 1831 Caspian Emerald Lizard

Baran (1980): Igdir, 1969 and Tasburun Koyt [&dir,

1977; Mulder (1995): Aralik, 1991.

Ophisops elegans Ménétries, 1832 Snake-eyed Lizard

Baran (1980): Tasburun Koyu I&dir, 1977; Mulder (1995):

Ararat northwest, 1987, 1992; Franzen and Sigg (1989):

surroundings of DoSubayazit, 1988; Trapp (web): Ararat.

Family Scincidae

Heremites septemtaeniatus (Reuss,

Grass Skink

1834) Southern

Species affiliation and genus name of a few Middle-

Eastern skinks were obscure for a long time. Karin et al.

(2016) replaced them into the old genus name Heremites

to solve polyphyly. Bahmani et al. (2017) solved the au-

ratus-septemtaeniatus problem for this region.

Mulder (1995): northwest slope of Ararat, province of

Isdir, 1987, 1990, 1992; Teynié (1987): Ararat northwest,

province of Idir, 1987; Franzen and Sigg (1989): sur-

roundings of DoSubayazit, 1988; Norstrom (1990): west

slope of Ararat; Trapp (web): Ararat; Fonters and Fonters

(2006): Igdir plain at the height of Suveren; Schneider

and Schneider (in litt.): northwest slope of Ararat, prov-

ince of Igdir (39°49°21”N 44°7°47°E and 39°47’58°N

44°10°44”E), 2008; Tosunoglu et al. (2011): between

Karakoyunlu and Tasburun, [&dir, 2008.

Eumeces schneideri (Daudin, 1802) Schneider’s Skink

Teynié (1991): northwest of Tuzluca, province of [&dir

and Agabey (19 km west of Tuzluca); Schneider and

Schneider (in litt): six km south of ISdir (39°51?10°N

44°4°13”E), 2009; AdaMerOs (web): Melekh, 2012.

Familiy Anguidae

Pseudopus apodus (Pallas, 1775) European Glass Lizard

Baran (1977b): I&dir side of Ararat, 1969; Mulder (1995):

northwest slope of Ararat, province of I&dir, 1987, 1990,

1992; Teynié (1987): northwest slope of Ararat, province

of ISdir, 1987; Norstrom (1990): west slope of Ararat;

Schneider and Schneider (in litt): Tuzluca, 2008.

Suborder Serpentes; Family Typhlopidae

Xerotyphlops vermicularis (Merrem, 1820) Eurasian

Blindsnake

First record in 1987 by Mulder (1995) on the plain of

Isdir at the foot of the northwest slope of Ararat.

May 2019 | Volume 13 | Number 1 | e175

Mulder

Teynié (1991): ISdir south/Ararat north; Tosuno$lu et al.

(2010): Tasburun, 2009; Go¢gmen (web): Melekli, 2012;

AdaMerOs (web): Tuzluca, 2014.

Family Boidae

Eryx jaculus (Linnaeus, 1758) Sandboa

Erroneously presented as first record for the region by

Tosunoglu et al. (2010).

Franzen and Sigg (1989): surroundings of

Dogubayazit, 1988; Teynié (1991): ASabey (19 km west

of Tuzluca); Norstrom (1990): west slope of Ararat,

1989; Schneider and Schneider (in litt.): Tasburun, 2009;

Tosunoglu et al. (2010): Melekli, 2008; Tosunoglu et al.

(2011): between Karakoyunlu and Tasburun, I&dir, 2008.

Family Natricidae

Natrix natrix (Linnaeus, 1758) Grass Snake

Boettger (1899): Kasikoparan (=Kazkoparan), province

of [Sdir, 1896; Baran (1976): Ararat [g&dir, 1969; Flardh

(1983): west slope of Ararat, 1980; Teynié (1987): north-

west of Ararat, province of ISdir, 1987; Norstrom (1990):

west slope of Ararat; AdaMerOs (web): Tuzluca, 2012.

Natrix tesselata (Laurenti, 1768) Dice Snake

Boettger (1899): Kasikoparan (=Kazkoparan), province

of Igsdir, 1896; Mertens (1952): ISdir, 1942; BasoSlu and

Baran (1980): Igdir; Flardh (1983): west slope of Ararat,

1980; Teynié (1987): southwest slope of Ararat, prov-

ince of Agri, 1987; Norstro6m (1990): Igdir plain; Mul-

der (1995): northwest slope of Ararat, province of [&dir,

1987; Franzen and Heckes (1999): northeast of Karabu-

lak (A&r1), 1990; Franzen and Sigg (1989): surroundings

of Dogubayazit, 1988; Schneider and Schneider (in litt):

six km south of [&dir (39°51’10”N 44°4713”E) and two

km east of Asaitavla/ 20 km southeast of Dogubayazit

(39°30’10”N 44°16751”E), 2008, 2009.

Family Colubridae

Dolichophis schmidti (Nikolsky, 1909) Red-bellied Rac-

Around 1990, a woman with a dead Dolichophis schmid-

ti stabbed onto and twisted around a stick was witnessed

standing along the road from I&dir to DoSubayazit at the

sideway to Kavaktepe (J. Mulder, not previously pub-

lished). This locality is one kilometer across the border

in the province of Agri. This probably is the first find of

this species in that province.

Mertens (1952): Igdir, at the foot of Ararat, 1942;

Basoélu and Baran (1980): slope of Ararat, [&dir, Flardh

(1983): west slope of Ararat, 1980; Mulder (1995): north-

Amphib. Reptile Conserv.

west slope of Ararat, province of [Sdir, 1987; Mulder

(not published before): Kavaktepe, west slope of Ararat,

province of Agri, around 1990; Norstrom (1990): west

slope of Ararat; Franzen and Heckes (1999): northeast

of Karabulak (Agri), 1990; AdaMerOs (web): Tuzluca,

2014.

Eirenis collaris (Ménétries, 1832) Collared Dwarf Snake

The subspecies macrospilota is only known from Mount

Takjaltu, near Kazkoparan southwest of Tuzluca. Hith-

erto only two specimens have been found, in 1903 and

1913. See Darevsky and Bakradze (1982).

Franzenand Sigg (1989): surroundings of DoSubayazit,

1988; Teynié (1991): north of Tuzluca, 1987, 1988 and

Dogubayazit, 1988; Mulder (1995): northwest slope of

Ararat, province of Igdir, 1990; AdaMerOs (web): Tu-

zluca, 2009, 2014.

Eirenis modestus (Martin, 1838) Dwarf Snake

Basoélu and Baran (1980): slope of Ararat (ISdir); Fran-

zen and Sigg (1989): surroundings of Dogubayazit, 1988.

Eirenis punctatolineatus (Boettger, 1892) Dotted Dwarf

Snake

Teynié (1991): north of Tuzluca, between 1987— and

1989; Schneider and Schneider (in litt.): plain south of

Isdir (39°50°44”N 44°5’20”E), 2008.

Elaphe dione (Pallas, 1773) Steppe Ratsnake

In 1988 three specimens were found by Garzoni and Ge-

niez (2004) a couple of kilometers west of the city of

Isdir. This was the first observation in Turkey. No subse-

quent observations are known.

Elaphe sauromates (Pallas, 1811) Blotched Snake

Boettger (1899): Kasikoparan (=Kazkoparan), prov-

ince of ISdir, 1896; Flardh (1983): west slope of Ararat,

1980; Teynié (1987): northwest side of Ararat, province

of ISdir, 1987; Norstrom (1990): west slope of Ararat;

Teynié (1991): I&dir south/Ararat north.

Hemorrhois ravergieri (Ménétries, 1832) Ravergier’s

Whip Snake

Mertens (1952): Ararat, 1940; Basoglu and Baran (1980):

slope of Ararat (I&dir) and Aralik; Flardh (1983): west

slope of Ararat, 1980; Norstrém (1990): west slope of

Ararat; Mulder (1995): northwest side of Ararat, prov-

ince of ISdir, 1987; Teynié (1987): northwest side of Ara-

rat, province of ISdir, 1987 and southwest side of Ararat,

province of Adri, 1987; Teynié (1991): Agabey (19 km

west of Tuzluca).

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Herpetofauna of Mount Ararat

Platyceps najadum (Eichwald, 1831) Slender Whip

Snake

Mulder (1995): northwest slope of Ararat, province

of ISdir 1987, 1991, and 1992; Trapp (web): “Ararat,”

2007; Schneider and Schneider (in litt.): seven km south

of [dir (39°51713”N 44°4°19”E), 2008.

Telescopus fallax Fleischmann, 1831 European Cat

Snake

Teynié (1991): Tuzluca north, between 1987 and 1991.

Zamenis longissimus (Laurenti, 1768) Esculapean Snake

The only observation in the area treated was at Aralik and

concerned two male specimens (Schweiger 1994).

Zamenis hohenackeri (Strauch, 1873) Transcaucasian

Ratsnake

Boettger (1899): Kasikoparan (=Kazkoparan), province

of ISdir, 1896; Flardh (1983): west slope of Ararat, 1980;

Teynié (1987): northwest slope of Ararat, province of

Isdir, 1987; Norstrom (1990): west slope of Ararat; Mul-

der (1995): northwest slope of Ararat, province of I&dir,

1992.

Family Lamprophiidae

Malpolon insignitus (Geoffroy de St-Hilaire, 1809) East-

ern Montpellier Snake

Clark and Clark (1973): 10 km South of ISdir, 1967;

Basoflu and Baran (1980): north of Ararat: Devlet

Uretme Ciftlizi (State farm) and Hasanhan near Aralik;

Teynié (1991): Tuzluca west; Mulder (1995): Northwest-

ern foot of Ararat/ [dir plain, 1990; Franzen and Heckes

(1999): south of Igsdir, 1990; Tosunoglu et al. (2010):

Melekli k6yt, 2008; Tosunoglu et al. (2011): between

Karakoyunlu and Tasburun, [&dir, 2008.

Family Viperidae

Macrovipera lebetina (Linnaeus, 1758) Blunt-nosed Vi-

per

Mertens (1952): “Tuzluca near ISdir,” 1942; Eiselt and

Baran (1970): Devlet Uretme Ciftligi (State farm), prov-

ince of Igdir, 1969; Mulder (1995): northwest slope of

Ararat, 1987, 1992; Franzen and Heckes (1999): south of

Igdir, 1990; Schneider and Schneider (in litt): Tasburun,

2008; AdaMerOs (web): Tuzluca, 2013.

Montivipera raddei (Boettger, 1890) Armenian Viper

The first record in the area under study was in the spe-

Amphib. Reptile Conserv.

cies description by Boettger (1890) on the basis of two

specimens collected in 1888. The locality was mentioned

as “Kasikoparan in Armenia,” nowadays Kazkoparan or

Kozkoparan is in the province of [gdir, Turkey.

Mertens (1952): Tuzluca near Igdir, 1942; Eiselt and

Baran (1970): near Igdir, on the slope of Ararat, 1969;

Flardh (1983): west slope of Ararat; Mulder (1995): west-

ern slopes of Ararat, province of I&dir, 1987, 1988, 1989,

1991, 1992; Teynié (1987): Ararat northwest, province of

Isdir, 1987; Norstrom (1990): west slope of Ararat.

Vipera eriwanensis (Reuss, 1933) Armenian Steppe Vi-

per

The locality “Kasikoparan” (=Kazkoparan, province of

Igdir) was mentioned by Boettger (in Radde 1899) as

a site for Vipera berus, by Nikolsky (1916) referred to

as Coluber berus dinnicki and subsequently assigned to

Coluber renardi. Taking into account the known distribu-

tion of eriwanensis (just outside the province of I&dir),

assigning that record to this taxon is the only credible

solution. Nilson et al. (1988) already indicated this.

Discussion

Mount Ararat and its surroundings belong among Tur-

key’s richest regions for reptile species diversity, with 36

species. Sindaco et al. (2000) presented a map of Ana-

tolian reptile species richness based on a grid with one-

degree latitude and longitude. The region under study

has a surface considerably smaller than one of those grid

squares. In their publication, the richest square (province

of Adana and surroundings) comprised 43 species, being

the only square containing more than 40 species. Only

seven out of the total of around 90 squares showed 31—40

species, all of them along the south coast and especially

around the aforementioned richest square.

Although the number of observed species in and

around Mount Ararat is high, the fact that many of them

are represented by only one or just a few observations

cannot be neglected. This scarcity of observations can be

the result of insufficient research, but could also indicate

relative rareness. Within the borders of Turkey some of

the species are only encountered in the area under study,

especially the species living on the plains around Ararat

and along the river Aras.

Mountain-dwelling species can be regarded as rela-

tively safe, but those living on the plains are under direct

threat due to ongoing and expected urban expansion and

intensified agricultural land use.

A summary of the herpetofauna of the province of

Igdir, which only comprizes the northern part of Mount

Ararat, was published by Tosuno§lu et al. in 2010. The

number of consulted sources in that paper was small (next

to their own observations, just eight other sources), caus-

May 2019 | Volume 13 | Number 1 | e175

Mulder

ing them to miss many species. Moreover, even some

species from the consulted sources were not mentioned.

First records of Sandboa and Grass Snake in the province

of I&dir were wrongly claimed. For the Grass Snake this

was later corrected by one of the authors (Gul 2011).

A doubtful article (just a German translation avail-

able) about Urodela in Turkey by Boglu and Hayman

(1978) mention Batrachuperus for the Ararat region:

“Wir haben ... einige Molchlarven erhalten welche aus

dem Araratgebiet stammen und mit Sicherheit den Batra-

chuperus zugeordnet werden mussen.” The occurrence

of Iranodon (the name recently in use for Batrachuperus)

has to be disclaimed as not reasonable. As no newts or

salamanders are known for the area, any larva would be

interesting though. In the same article, the ‘species’ Neuw-

rergus caucasicus 1s treated as occurring at Ararat. As lo-

cality they gave “Bis 2,500 m hohe Gebirgsregionen des

Ararat. Aralik, Igdir, Tasburun” (high mountain region of

Ararat up to 2,500 m, Aralik, Igdir, Tasburun). This spe-

cies account is given next to N. strauchii and N. croca-

tus. The non-existent binominal name gives no clue as to

which species the authors would have meant and makes

the source highly unreliable. See also Schmidtler (1984),

Schatti and Sigg (1989) and Olgun et al. (1997).

The locality ‘Tasburun Koyt, [&dir’ was visited by

Baran in 1977. For instance, Phrynocephalus heliosco-

pus and Paralaudakia caucasia were found there (Baran

1980) and subsequently by other authors. Later, these

finds were surprisingly quoted as located at “Tasburun

Koyti near KaSizman’ (Baran et al. 1989), which is a

mere 110 km to the west. Though the collector listed is

one of the authors, this must be erroneous. The habitat

seems to be inappropriate for Phrynocephalus.

Several additional species can be expected for the re-

gion, as they are known to occur relatively close to the

area under study. They include the following taxa:

Tranolacerta brandti (De Filippi, 1863) Persian Lizard

About 20 km south of the area under study the Persian

lizard was recently found in Karadulda, province of Van,

2014 (Yildiz and I§ci 2015).

Darevskia unisexualis (Darevsky, 1966) White-bellied

Lizard

Schmidtler et al. (1994): Diyadin, province of Agri.

Parvilacerta parva (Boulenger, 1887) Dwarf Lizard

Near the area under study observations were made, e.g.,

between Selim and Karakurt in 1999 (Baran et al. 2004)

and in the province of Agri in 2014 (Anonymous 2014).

Coronella austriaca Laurenti, 1768 Smooth Snake

The Smooth Snake has been found in 1942 in Boctiklii at

Amphib. Reptile Conserv.

the north border of the Aras valley (Mertens 1952; Baran

1977a), about 30 km from the area under study.

Eirenis eiselti Schmidtler and Schmidtler, 1978 Eiselt’s

Dwarf Snake

Mentioned as occurring in the province of Aéri, 2014

(Anonymous 2014).

Hemorrhois nummifer (Reuss, 1834) Coin-marked Snake

The species is known from the Aras valley in Armenia

(Schatti and Agasian 1985).

Rhynchocalamus melanocephalus (Jan, 1862) Black-

headed Ground Snake

The Black-headed Ground Snake occurs in nearby Ar-

menia on mountain slopes bordering the Aras Valley in

Yerevan and Ararat (Arakelyan et al. 2011).

The mentioning by both Kamali (2017) and Rajabi-

zadeh (2017) of Dolichophis jugularis, next to Dolicho-

Dhis schmidti, in Yran directly bordering the Ararat region

must be erroneous.

During the publication of this checklist, it became obvi-

ous that a paper was going to be published on more or

less the same subject. At the Fourth International Sym-

posium Mount Ararat and Noah’s Ark in October 2017

an oral presentation was given, from which a published

text became available in October 2018 (Yildiz et al.

2018).

A comparison of the results is briefly treated here. As

a result of a different scope of area (more strictly around

the mountain) their list is of course shorter.

The literature search is quite limited and several classic

publications of well-known herpetologists treating her-

petological distribution in Turkey are not included, like

Méhely (1894), Mertens (1952), Eiselt and Baran (1970),

Clark and Clark (1973) and Baran (1976, 1977a,b, 1980).

Also, other informative sources like Flardh (1983), Fran-

zen and Sigg (1989), Norstrom (1990) and Teynié (1991)

are not incorporated. This resulted in an incomplete view

of the known distribution and dates of first publication

and, even considering the different scope of area, the lack

of several species, 1.e., Mediodactylus kotschyi, Lacerta

media, Eirenis modestus, Eirenis punctatolineatus, and

Elaphe sauromates, all of which occur within their scope

of area.

Species affiliation of the local Toad Head Agama is

still not resolved sufficiently. It is given as Phrynoceph-

alus horvathi by them and as P. helioscopus here (fol-

lowing The Reptile Database), but undoubtedly the same

taxon is intended. The Heremites species in the region

is treated by them as the species auratus, while here it

is presented as septemtaeniatus, the latter according to

Bahmani et al. (2017).

May 2019 | Volume 13 | Number 1 | e175

Herpetofauna of Mount Ararat

Acknowledgements.—Jaap van Wingerde is thanked

for his help in obtaining some hard to get references and

Wolfgang Wuster for critically reviewing the text.

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John Mulder was born in Rotterdam, the Netherlands, and he received his M.Sc. degree in biology from Utrecht University.

His interests include ecology, natural history, and biogeography of the Palaearctic herpetofauna and orthopteran fauna. John has

been associated with the Natural History Museum Rotterdam since the 1980s. John has travelled extensively in the Middle East

and visited Turkey, Syria, Iran, Yemen, and Georgia. John has extensive experience studying the herpetofauna in the respective

countries, especially that of Eastern Anatolia. John has authored or co-authored several papers on herpetology, including papers on

biogeography (Turkey), the Zagrosian lizard (Iran), unexpected ophiophagy by a snake (Iran), ecology and morphology of the rare

Anatolian Meadow Viper (Turkey), morphology of Montivipera albizona (Turkey), northern distribution limits of the Bridled Skink

(Turkey) and distribution of the Transcaucasian Sandviper.

Amphib. Reptile Conserv. 172 May 2019 | Volume 13 | Number 1 | e175

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

13(1) [General Section]: 173-197 (e176).

urn:lsid:zoobank.org:pub:0AB829CA-DAFF -4EE9-B7D7-FBC037B4B2B0

Taxonomic assessment of Craugastor podiciferus

(Anura: Craugastoridae) in lower Central America

with the description of two new species

123F rick Arias, *°Andreas Hertz, and '*Gabriela Parra-Olea

'Departamento de Zoologia, Instituto de Biologia, UNAM, AP 70-153 Ciudad Universitaria, CP 04510, Ciudad de México, MEXICO *Posgrado

en Ciencias Biologicas, Universidad Nacional Autonoma de Mexico, Av. Ciudad Universitaria 3000, C.P. 04360, Coyoacan, Ciudad de México,

MEXICO Escuela de Biologia, Universidad de Costa Rica, San Pedro, 11501-2060 San José, COSTA RICA ‘Department of Biology, University of

Massachusetts Boston, 100 Morrissey Blvd., Boston, MA 02125, USA °Senckenberg Forschungsinstitut und Naturmuseum, Senckenberganlage 25,

60325, Frankfurt am Main, GERMANY

Abstract.—The systematics and taxonomy of polytypic species Craugastor podiciferus are poorly understood

due to the high level of phenotypic polymorphism between and within species and the lack of molecular data

from topotypic specimens. Herein are reported results of a well-sampled study including all known species

of the C. podiciferus species group, several localities from highlands in Costa Rica and western Panama, and

for the first time, samples from the type locality of C. podiciferus. A phylogenetic analysis based on the DNA

sequences of the mitochondrial 16S rRNA (16S) and cytochrome oxidase 1 (COI) genes and a morphometric

analysis are also included. Based on the results, we restrict C. podiciferus to the populations from the Cordillera

Volcanica Central and Cordillera de Talamanca in Costa Rica and western Panama. Craugastor podiciferus

sensu stricto and six additional clades from the highlands of Costa Rica constitute the well-supported C.

podiciferus sensu lato clade. These analyses support the existence of three additional species from the Pacific

slope of southwestern Costa Rica and western Panama. Herein, two lineages are described as new species and

revised descriptions for C. podiciferus and C. blairi are provided. The name C. blairi is resurrected and used

for populations from the Cordillera de Talamanca and Cordillera Central in western Panama. Two additional

species are named. One is easily differentiated by the presence of nuptial pads in adult males, a smooth venter,

and flat subarticular tubercles. The other, named for populations from southwestern Costa Rica, is recognized

by its coarsely areolate venter, projecting subarticular tubercles, and heel without a projecting tubercle. The

recognition of these three species from the lower montane rainforest highlights the role of the highlands on the

Pacific slope of Costa Rica and Panama in the diversification of the C. podiciferus species group.

Keywords. Brachycephaloidea, cryptic species, DNA barcoding, Talamanca, Terrarana, Costa Rica, Panama

Citation: Arias E, Hertz A, Parra-Olea G. 2019. Taxonomic assessment of Craugastor podiciferus (Anura: Craugastoridae) in lower Central America

with the description of two new species. Amphibian & Reptile Conservation 13(1): [General Section]: 173-197 (e176).

4.0 International (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in any

medium, provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced, are

as follows: official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org.

Received: 18 January 2019; Accepted: 4 April 2019; Published: 13 May 2019

Introduction

The direct-developing frogs of the Craugastor podic-

iferus species group (Hedges et al. 2008) are found from

eastern Honduras to Central Panama (AmphibiaWeb

2019; Savage 2002) with most of the species (nine out of

ten) restricted to Isthmian Central America. The distribu-

tion of this group ranges from sea level to 2,700 m a.s.l.

in a wide variety of habitats, from tropical rain forest

and cloud forest, to montane forest (Savage 2002). The

morphological delimitation between members of the C.

podiciferus species group 1s difficult due to the extremely

conserved morphological characters and the high level

of phenotypic polymorphism within species and popu-

lations. The systematics and taxonomy of the C. podic-

iferus species group has been poorly studied; however,

previous molecular studies have suggested the existence

of several unnamed species that are masked under the

current names (Savage 2002; Crawford 2003; Crawford

and Smith 2005; Streicher et al. 2009). For example,

Crawford and Smith (2005) examined 10 samples (seven

species) of the C. podiciferus species group and found

Correspondence. ' gparra@ib.unam.mx (corresponding author); '?* eapiedra@gmail.com; ** andreas. hertz@umb.edu

Amphib. Reptile Conserv.

May 2019 | Volume 13 | Number 1 | e176

Arias et al.

the presence of two unnamed species (Craugastor sp. B

and Craugastor sp. C); they also found C. stejnegerianus

(Cope, 1893) to be paraphyletic. Recently, Arias et al.

(2016) showed that populations formerly considered to

be part of the C. stejnegerianus from southwestern Costa

Rica and western Panama belonged to a different species

and described them as C. gabbi Arias, Chaves, Crawford,

and Parra-Olea, 2016.

Craugastor podiciferus (Cope, 1875) is the most com-

plicated taxon within the C. podiciferus species group. Its

morphological polymorphism has been recognized since

its description by Cope (1875), who described it based on

four varieties plus two additional species (C. muricinus

and C. habenatus) in the same paper, with specimens from

the same locality. In addition, Cope (1875) stated that “the

colors of this species vary remarkably, more than I have

observed to be the case in any other frog.” Subsequently,

Barbour (1928) described C. blairi using specimens from

western Panama, which Taylor (1952) later synonymized

together with C. muricinus and C. habenatus under C.

podiciferus. An additional species, C. jota (Lynch 1980),

was named using specimens from western Panama, and

was suggested to be related to C. podiciferus. More re-

cently, the name C. podiciferus has been used for speci-

mens from several highland populations (1,089—2,650 m

a.s.l.) on both slopes of the mountain ranges in Costa Rica

and western Panama (Savage 2002). Streicher et al. (2009)

used specimens from several populations referred to as C.

podiciferus to perform a well-sampled phylogenetic study

of this complex. They found C. podiciferus is represented

by six clades, each likely representing distinct species.

In addition, the populations from western Panama were

not grouped with the main C. podiciferus clade, but were

called Craugastor sp. B.

Although the molecular and geographical evidence

shown by Streicher et al. (2009) supported the presence of

several species under the name C. podiciferus, taxonomic

changes have not yet been implemented, mainly due to the

lack of topotypic specimens of C. podiciferus. The correct

type locality has been discussed by Savage (1970), and

Arias and Chaves (2014) concluded it is on the Caribbean

slopes of Cerro Kamuk.

Here, specimens of Craugastor podiciferus from the

type locality are included for the first time together with

those from several additional localities from Costa Rica

and western Panama. Mitochondrial DNA sequences were

used to address three goals: 1) to identify C. podiciferus

sensu Stricto in a phylogenetic context, 2) to evaluate the

phylogenetic relationships of the C. podiciferus species

complex, and 3) to evaluate the taxonomic status of popu-

lations from the Pacific slopes of southwestern Costa Rica

and western Panama. The result is a comprehensive revi-

sion of the C. podiciferus species complex, in which two

new species are described from southwestern Costa Rica

and western Panama. Additionally, an old name is resur-

rected for a third species in mountainous western Panama.

Amphib. Reptile Conserv.

Materials and Methods

Species criterion: The general metapopulation lineage

species concept is followed here (Simpson 1951; Wiley

1978; de Queiroz 2007). Consistent with this concept, a

species is recognized when there is evidence of metapopu-

lation lineage separation, preferably based on multiple

lines of evidence following a consensus protocol for inte-

grative taxonomy (Dayrat 2005; Padial et al. 2010).

Taxon sampling: The frogs were collected in the field, eu-

thanized, fixed in 10% formalin, and processed in 70% eth-

anol for long-term storage. A tissue sample was preserved

in 96% ethanol or RNAlater and used for genetic analysis.

Museum collection acronyms follow Frost (2019) with the

addition of AH for Andreas Hertz field numbers, EAP for

Erick Arias field numbers, and CRARC for the Costa Rica

Amphibian Research Center private collection.

Amplification and sequencing: Partial sequences of the

large subunit ribosomal RNA (16S) mitochondrial gene

were determined for six specimens of Craugastor sp. |

(Arias et al. 2018), Craugastor sp. 2 (Arias et al. 2018),

and Craugastor sp. B (Crawford and Smith 2005) from

the Pacific slopes of southwestern Costa Rica and west-

ern Panama (Fig. 1). The sequences obtained herein were

compared with those available in GenBank for the C.

podiciferus species group. The protocols for DNA extrac-

tion, amplification, sequencing, and editing follow those

of Arias et al. (2018). The list of vouchers and GenBank

accession numbers used in this study are provided in Ap-

pendix I.

Phylogenetic analyses: Sequence alignments used the

MAFFT software (Katoh et al. 2017) under the “auto”

strategy and default parameters, and were trimmed to

the point where a majority of the taxa had sequence

data. The sequence data were partitioned by gene, and

the COI data were further partitioned by codon posi-

tion. PartitionFinder v1.1.1 (Lanfear et al. 2012) and

the Bayesian information criterion (BIC) were used to

select both the best partition scheme and the best mod-

el of sequence evolution for each partition. A single

set of branch-lengths were used across all partitions

(branchlengths=linked), and the search for the best par-

tition scheme was implemented using a heuristic search

(scheme=greedy). Four partitions were defined, a priori,

one for 16S and three for COI (one for each codon).

Phylogenetic analyses were performed using both the

maximum likelihood (ML) and Bayesian inference (BI)

methods. The maximum likelihood analysis was _per-

formed using Garli 2.01 (Zwickl 2006), with 10 search

replicates with the following default setting values:

streefname=random, attachmentspertaxon=24, genthresh-

fortopoterm=100000, __ significanttopochange=0.00001.

For bootstrapping, 1000 replicates were run with the

previous settings with the following changes: genthresh-

May 2019 | Volume 13 | Number 1 | e176

Craugastor podiciferus species group in Central America

83° W 82°W

84°W

Pacific

Ocean

Species

() Craugastor aenigmaticus

@ Craugastor blairi

@ C. podiciferus sensu stricto} Elevation (m)

O C. podiciferus sensu lato 50

A Craugastor sagul Sp. NOV. | 1000-2500 ET |

@ Craugastor zunigai sp. nov.\ I >2500 Kilometers

85°W

83°W 82°W

Fig. 1. Map showing the known populations of Craugastor blairi, C. sagui sp. nov., and C. zunigai sp. nov. from the lower montane

rainforest in southwestern Costa Rica and western Panama, and populations of other species of the C. podiciferus species group

inhabiting the highlands of Costa Rica and western Panama. The arrow indicates the type locality of C. podiciferus.

fortopoterm=10000, significanttopochange=0.01, treere-

jectionthreshold=20, as suggested in the Garli manual, to

accelerate the bootstrapping. The bootstrap consensus tree

was performed using Sumtrees (Sukumaran and Holder

2010b) from the DendroPy package version 4.4.0 (Suku-

maran and Holder 2010a). Bayesian phylogenetic analy-

sis was performed using MrBayes 3.2.6 (Ronquist et al.

2012) with the partition scheme and model of sequence

evolution for each partition as selected previously. Two

separate analyses were run, each consisting of 20 million

generations, sampled every 1,000 generations, and four

chains with default heating parameters. A time-series plot

of the likelihood scores of the cold chain was examined

to check the stationarity using Tracer 1.6 software (Ram-

baut et al. 2014). The first 25% of trees were discarded as

burn-in and the remaining trees were used to estimate the

consensus tree along with the posterior probabilities for

each node and each parameter. Maximum likelihood and

Bayesian analyses were run on the CIPRES portal (Miller

et al. 2010). Genetic distances (uncorrected p-distances)

were computed using MEGA6 (Tamura et al. 2013).

Morphometric analyses: A morphometric analysis was

performed to compare the three populations from the high-

lands of southwestern Costa Rica and western Panama.

Specimens examined included 19 specimens of Craugas-

tor sp. 1, seven specimens of Craugastor sp. 2, and 25

specimens of Craugastor sp. B (Appendix II). The speci-

Amphib. Reptile Conserv.

mens were deposited in Museo de Zoologia (UCR), San

José, Costa Rica, and the Senckenberg Research Institute

and Nature Museum, Frankfurt, Germany (SMF). The fol-

lowing morphological measurements were recorded as de-

scribed by Savage (2002), Duellman and Lehr (2009), and

Arias et al. (2016): snout-vent length (SVL), head length

(HL), head width (HW), inter orbital distance (IOD),

width of the upper eyelid (EW), eye-nostril distance (EN),

eye diameter (ED), and tympanum diameter (TY). Mea-

surements were performed using dial calipers and rounded

to the nearest 0.1 mm. To avoid allometric effects rela-

tive to differences in size and shape between species and

between individuals, data were transformed using the

method of Lleonart et al. (2000). Additional proportions

reported herein include: EW/IOD, IOD/HW, TY/ED, EN/

ED, ED/HL, HL/HW, and EN/HL. The sex of individuals

was determined by gonadal morphology; specimens with

opaque seminal vesicles were assumed to be adult males,

and those with developed oviducts were assumed to be

adult females. The general terminology for morphological

characteristics follows Duellman and Lehr (2009). Savage

(2002) was followed for the term “supernumerary tuber-

cles,” to refer to tubercles on the phalanges (between sub-

articular tubercles), which is different from the tubercles

referred to herein as accessory palmar or plantar tubercles.

The mean, standard deviation, and range for each mor-

phometric variable were calculated without correction. All

variables were used to perform a linear discriminant analy-

May 2019 | Volume 13 | Number 1 | e176

Arias et al.

Craugastor gollmerii AJC1926

76/96

kK is

19/--~ | 8

61/78 [

39/67|

50/9

C. podiciferus s.1.

59/79

/92/*

26/72

27/5]

58/96

30/94

74/*

15/48

81/*

| UCR22703 Las Alturas

All :

SMF104015 La Nevera .

SMF 104017 LaNevera |Craugastor sagui Sp. nov.

SMF 104014 Cerro Sagui

UCR22709 Las Alturas

UCR20429 Potrero Grande|-/"Ugastor zunigai sp. Nov.

UCR20389 Potrero Grande

SMF 104034 Guayabito

SMF 104027 Volcan Bart

SMF 104037 La Estrella

USNM563039 Palo Alto rh 8

FMNH257689 La Fortuna|Craugastor blairi

| SMF104023 Fortuna

SMF 104033 Fortuna

SMF 102024 Fortuna

*/a- SMF104020 Jurutungo

UCR22737 Cerro Haku

C. aenigmaticus

UCR21951 Cerro Utyum

__*/*- UCR22643 Guayacan

~“t UCR22269 San Saale: bransfordii

*/%*, UCR22668 Bribri

UCR20050 aera Pp oly P tych Us

“= UCR22625 Cascajal

UCR22619 Tapanti Ic underwoodi

*/*— [JCR22211 Tausito + ys

UCR22671 Ras persimilis

UCR21864 San Vito :

UCR21863 San aC gabbi

*/*» UCR20352 Potrero Grande

EAP0514 Palmar Norte

*/%) SMF79759 Selva Negra

USNMS559393 Tapalwas

FMNH257673 Monte Verde

UCR16361 Tapesco

| UCR22675 Monte Verde

FMNH257669 Monte Verde

UCR22146 Cascajal

«af CRARCO12 Volcan Turrialba

UCR20992 Nectandra

UCR22201 El Empalme

SMF104005 Changena

UCR23175 Alto Uren

UCR19862 Lori

UCR19856 Lori

UCR19860 Lori

UCR19853 Lori

UCR23169 Alto UrenIC, podiciferus ““Siola”

*/4 UCR22228 Pico Blanco ae “Di ”

DOR aaae pe BIC. podiciferus “Pico Blanco

61) CRARC0247 Monte Verde

dor FMNH257671 Monte Verde

UCR16353 Montafia Azul

Ueno 0 A eC. podiciferus “Chumacera”

UCR16585 Copey

EAP0509 Fila Costefia

FMNH257651 Fila Costefia

UCR22091 Quebradas

* [7

64/87

ok /% *

Ic. stejnegerianus

Ic . lauraster

C. podiciferus “Monte Verde”

C. podiciferus sensu stricto

C. podiciferus “San Gerardo”

C. podiciferus “Fila Costefia”

Fig. 2. Maximum likelihood phylogram of the Craugastor podiciferus species group based on the 16S and COI mitochondrial DNA

gene markers. Bootstrap proportions and posterior probability (multiplied by 100) values obtained with MrBayes before and after

the slash, respectively. The scale bar refers to the estimated substitutions per site. Asterisks represent support > 95. The blue clade

corresponds to C. podiciferus sensu stricto.

sis to determine whether the morphometric variables were

effective in predicting the species. The proportion of cor-

rectly classified individuals was validated using jackknife

accuracy (Manly 1994). All analyses were performed us-

ing R v3.3.3 (R Core Team 2017).

Results

Molecular: The resulting mitochondrial data matrix in-

cluded 59 sequences with a total sequence length of 1,222

bp including gaps; 565 bp for 16S and 657 bp for COI. The

best strategy partition contains four partitions, one for 16S

and one for each codon in COI. The following substitution

models were selected: GTR+G for 16S, K80+I+G for COI

Amphib. Reptile Conserv.

176

codon position 1, HK Y+I+G for COI codon position 2,

and GTR+I+G for COI codon position 3. Mitochondrial

genetic distances are shown in Table |. Genetic distances

between Craugastor sp. | and all other members of the

Craugastor podiciferus species group are 4.9-15.2% for

16S and 13.5—21.6% for COI. Craugastor sp. 2 is sepa-

rated by an uncorrected genetic distance to other members

of the Craugastor podiciferus species group of 2.9-15.7%

for 16S and 14.3-21.3% for COI. Genetic distances be-

tween Craugastor sp. B and other members of the Crau-

gastor podiciferus species group are 2.9-16.2% for 16S

and 14.3-19.6% for COI.

The ML and Bayesian trees were similar in topology

and show six well-supported clades (Fig. 2). The first is

May 2019 | Volume 13 | Number 1 | e176

Craugastor podiciferus species group in Central America

x ®

e &

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oy °

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Species @ C. biairi A C. sagui sp. nov.™ C. zunigai sp. nov.

Fig. 3. Linear discriminant analysis showing the morphological separation among the three species from the lower montane

rainforest in southwestern Costa Rica and western Panama.

formed by specimens from La Nevera and Cerro Sagui

from western Panama. The second and third clades are

formed by specimens from southwestern Costa Rica and

western Panama, respectively. The fourth clade comprises

C. aenigmaticus from the montane rainforest of the Cor-

dillera de Talamanca. A fifth major clade includes seven

species of the C. podiciferus species group that mainly

occur from the lowlands to mid-elevations from eastern

Honduras to central Panama. Finally, a sixth major clade

contains specimens from the type locality of C. podicifer-

us and several other localities from the highlands of Costa

Rica and western Panama that are tentatively referred to

this species. The main difference between the ML and the

Bayesian topology is the position of the clade containing

the samples from La Nevera and Cerro Sagui. In the ML

tree, this group was the sister to all other members of the

C. podiciferus species group (although with very low sup-

port), while in the Bayesian tree (not shown), it was the

sister to a clade containing the samples from Las Alturas

and Potrero Grande and C. blairi.

Morphometry: Morphometric variation and comparisons

among the species are shown in Table 2. The proportion

of specimens correctly assigned to the species was 82%,

showing a clear morphological separation between the

specimens of the three populations of southwestern Costa

Rica and western Panama (Fig. 3).

Systematics

Redefinition of Craugastor podiciferus (Cope, 1875)

and C. blairi (Barbour, 1928)

Craugastor podiciferus: The precise type locality of Crau-

Amphib. Reptile Conserv.

VEE

gastor podiciferus has been a matter of some uncertainty.

The taxon was described by Cope (1875) based on material

collected by W.M. Gabb from “slope of Cerro Pico Blan-

co” in 1874. Savage (1974) corrected it to Cerro Utyum,

Canton de Talamanca, Provincia de Limon, 1,524-2,134

m a.s.l. but collected no additional specimens because he

did not reach that site. Recently, Arias and Chaves (2014)

corrected the type locality to a place between Cerro Pat

and the headwaters of the Lari River, elev. 1,520—2,135,

Provincia de Limon, Caribbean slope of Cerro Kamuk.

The type locality is a remote site within the Parque Inter-

nacional La Amistad, in the Cordillera de Talamanca. It is

only accessible on foot and requires three days of hiking

from the last village (Amubri, Talamanca). For the pres-

ent study, specimens were collected in the surroundings

of the type locality according to Arias and Chaves (2014)

[Figs. 1 and 4A]. Based on phylogenetic relationships, we

restrict the taxon Craugastor podiciferus to the popula-

tions from Cordillera Volcanica Central from Costa Rica

and Cordillera de Talamanca (Caribbean slopes) in Costa

Rica and western Panama (Fig. 1).

Craugastor jota: This species was described by Lynch

(1980) based on specimens from the Changena River in

western Panama, at 760 m a.s.I., collected by Linda Trueb

in 1966. It was placed in the C. podiciferus species group

based on morphology (Hedges et al. 2008). In this study,

specimens from Changena River, western Panama, very

near the type locality of C. jota (Linda Trueb, pers. comm.)

are included. In the phylogenetic analyses these specimens

fall within the C. podiciferus sensu stricto clade. As a re-

sult, we suggest that C. jota should be referred to as a ju-

nior synonym of C. podiciferus.

May 2019 | Volume 13 | Number 1 | e176

Arias et al.

Table 1. Mean uncorrected genetic distances among lineages of the

Craugastor podiciferus species group using the COI (above) and 16S

(below) mitochondrial genes.

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Amphib. Reptile Conserv. 178

Craugastor blairi: The precise type locality of Craugas-

tor blairi is unknown. Barbour (1928) indicated the type

locality as “from Gutierrez, Bocas del Toro Province,

Panama (near Costa Rican frontier).” The type series

was collected by Emmett R. Dunn and Chester Duryea

in the summer of 1923. Their expedition followed a trail

from Chiriquicito at the Laguna de Chiriqui (today in the

Corregimiento of Miramar, Bocas del Toro) to Boquete,

Province of Chiriqui. Most of their trail climbs up the

Atlantic slopes of the Cordillera Central (Savage 1970)

before descending into the high valley of Boquete. On the

same route, Dunn and Duryea collected several other new

species and identified the type localities as either “Gutier-

rez” or “La Loma.” Species with type locality “La Loma”

are Bolitoglossa colonnea (Dunn, 1924), Dermophis par-

viceps (Dunn, 1924), Hyloscirtus colymba (Dunn, 1931),

Pristimantis pardalis (Barbour, 1928), and Pristimantis

caryophyllaceus (Barbour, 1928). Each of these species

has a vertical distribution range that enters the lowlands

to almost sea level (K6hler 2011). Dunn gives the eleva-

tion of La Loma as 2,000 feet (610 m) in the descrip-

tion of D. parviceps and as 1,500 feet (460 m) in the de-

scription of H. colymba. In comparison to “La Loma,” a

species with the type locality “Gutierrez” is Craugastor

obesus (Barbour 1928). Dunn (1940) also mentioned col-

lecting C. monnichorum at “Gutierrez,” and both species

had a more premontane to montane distribution. There-

fore, we believe that “Gutierrez” is further uphill than

“La Loma” and thus closer to the continental divide and

much closer to Boquete than to the Caribbean lowlands.

Several specimens collected at different sites in the vicin-

ity of Boquete on the foothills of Bari Volcano form a

separate clade with specimens collected at Cerro La Es-

trella, Cerro Guayaba, Guayabito, and La Fortuna (Fig. 1

and 4B). For the above reasons, we resurrect the name C.

blairi for this clade.

Restricting Craugastor podiciferus to populations

inhabiting the Caribbean slopes of Cordillera de Tala-

manca of Costa Rica and extreme western Panama and

C. blairi to populations inhabiting the Cordillera Central

from Bart Volcano to the east, results in two allopatric

lineages on the Pacific slopes of Cordillera de Talamanca,

Costa Rica and those from Cordillera Central, Panama

without assignment to an existing taxon. With no avail-

able name present in the synonymy of C. podiciferus for

either of these populations, these two lineages are herein

described as new species, and redescriptions are provided

for C. podiciferus and C. blairi.

Craugastor podiciferus (Cope, 1875)

Common name: Polymorphic Dirt Frog

(Figs. 4A and 6)

Syntypes: USNM 30662, USNM 30664—75, and MCZ

11841. All specimens from “5,000 to 7,000 feet (eleva-

tion), on the Caribbean slopes of Cerro Pico Blanco,” col-

lected by William M. Gabb in 1874.

May 2019 | Volume 13 | Number 1 | e176

Craugastor podiciferus species group in Central America

Table 2. Mean, standard deviation (S.D.), and range for morphometric variables by species.

Variable

Craugastor podiciferus

Craugastor blairi

Craugastor sagui sp. nov.

Craugastor Zunigai sp. nov.

Mean + S.D. Range Mean+S.D. Range Mean + S.D. Range Mean + S.D. Range

SVL 25.6+4.3 16.9-33.9 23.2+4.8 13.4-30.6 21.546.5 13.0-30.1 20.843.9 13.8-26.5

HE 10.4+1.5 7.0-12.9 8.3+1.5 5.6—11.0 8.1+2.5 4.7-10.9 8.4+1.5 5.7-10.3

HW 10.2+1.6 6.5-13.3 8.9+1.8 5.5-12.0 8.0+2.4 4.9-11.5 8.2+1.6 5.6—10.7

BD 3.0+0.5 2.0-3.9 2.9+0.6 1.9-3.8 2.9+0.7 2.04.0 2.5+0.3 1.9-2.9

GY! 1.8+0.4 1.0-2.7 1.8+0.4 0.8-2.6 1.7+0.6 0.8-2.6 2.0+0.3 1.6—2.9

EW 19033 1.2-2.4 1.6+0.3 0.9-2.1 1.5+0.4 0.9-2.0 1.6+0.3 1.2—2.1

IOD 3.3+0.5 2.5-4.2 3.3+0.6 1.84.3 2.9+0.8 1.7-3.9 2.7+0,5 1.8-3.7

EN 2.60.4 1.9-3.2 2.2+0.4 1.3-3.0 2.1+0.8 1.1-3.2 2.0+0.4 1.3-2.6

EW/IOD 28+033 2.3—3.6 0.50+0.09 0.34-0.73 0.52+0.06 0.46—0.61 0.59+0.08 0.44—0.76

IOD/HW 0.6+0.1 0.40.7 0.37+0.04 0.30-0.45 0.37+0.03 0.34-0.40 0.34+0.02 0.30—0.38

TY/ED 0.6+0.1 0.4—1.0 0.66+0.23 0.40-1.09 0.58+0.18 0.40—0.81 0.81+0.17 0.62—1.34

EN/ED 0.9+0.1 0.7-1.0 0.77+0.08 0.63-0.91 0.71+0.11 0.55—0.88 —0.81+40.10 0.63—0.98

ED/HL 0.3+0.1 0.20.3 0.35+0.03 0.30-0.43 0.37+0.04 0.31-0.43 0.29+0.02 0.26—0.33

HL/HW 1.0+0.1 0.9-1.1 0.94+0.05 0.84-1.05 1.01+0.06 0.95—1.08 1.03+0.05 0.95—1.14

EN/HL 0.2+0.1 0.2—0.3 0.27+0.02 0.23-0.30 0.26+0.02 0.23-0.29 =0.24+0.02 0.20—0.27

Genetic reference specimen: UCR 23175 (EAP 0810),

an adult female from Costa Rica: Provincia de Limon:

Canton de Talamanca: Distrito de Telire: Parque Inter-

nacional La Amistad, (9.366°, -83.042°; 1,860 m a.s.l.),

collected by Erick Arias and Omar Zufiga on 27 October

2016.

Referred specimens: UCR 23155 (EAP 0803), adult fe-

male, same data as the genetic reference specimen. UCR

23145 (EAP 0792), an adult male from Costa Rica: Pro-

vincia de Limon: Canton de Talamanca: Distrito de Te-

lire: Cerro Pat, Parque Internacional La Amistad, (9.393°,

-83.025°; 1,450 m a.s.l.), collected by Erick Arias and

Omar Zufiga on 26 October 2016.

Assignment to group: Assigned to Craugastor based on

molecular analysis and the following characters: cranial

crests absent and Toe III larger than Toe V.

Diagnosis: The following combination of characteristics

distinguish Craugastor podiciferus (Fig. 5A—6) from oth-

er described species in the genus: 1) skin on the dorsum

is smooth to scattered tubercles; 2) skin on the venter is

smooth, at least along the midline; 2) vocal slits in adult

males; 3) nuptial pads absent; 4) unwebbed toes; 5) heel

with a projecting tubercle; 6) accessory palmar and plan-

tar tubercles absent, usually no supernumerary tubercles

under the digits; and 7) subarticular tubercles flat in form.

Craugastor (Craugastor) podiciferus is a small species

with the following characteristics: (1) skin on the dor-

sum smooth to scattered tubercles; head smooth; venter

smooth; flanks smooth with scattered tubercles to warty;

posterior surface of hind limbs surrounding cloaca strong-

ly areolate; some specimens with a pair of scapular, dor-

solateral or lateral folds; discoidal fold complete laterally

Amphib. Reptile Conserv.

and posteriorly; (2) tympanic membrane round, heavily

pigmented; prominent in males, evident in females; annu-

lus evident through the skin; (TY/ED = 44.9-100%); usu-

ally with a pair of supratympanic folds; (3) snout subovoid

in dorsal view, rounded in profile; loreal region concave;

canthus rostralis usually rounded; (4) eyelid granular, with

several low tubercles forming a more or less distinct ridge

on outer edge of the eyelid continuous with supratympanic

fold (EW/IOD = 40.5—71.2%); cranial crests absent; (5)

vomerine teeth in two transverse fasciculi, behind choa-

nae; choanae smaller than dentigerous; (6) vocal slits

and large single vocal sac in adult males; nuptial pads

absent; (7) fingers I and II subequal; discs absent, some

specimens with terminal transverse grooves on fingers,

especially in males; tips symmetric, usually rounded but

pointed in fingers III-IV in some specimens; pads ovoid

to triangular; (8) fingers lack lateral fringes; webbing ab-

sent; thenar and palmar tubercles low, ovoid, similar in

size; supernumerary tubercles absent; accessory palmar

tubercles usually absent but 1—2 low barely distinct tu-

bercles visible in some specimens; subarticular tubercles

round in basal outline, flat in form and globular in profile:

(9) ulnar fold absent but tubercles sometimes visible; (10)

heel with a projecting tubercle; inner edge tarsal with an

indistinct short ridge, outer smooth or with tubercles; (11)

toes lacking lateral fringes; inner metatarsal tubercle elon-

gate, outer rounded, much smaller than inner, inner and

outer metatarsal tubercles projecting; supernumerary and

plantar tubercles absent; subarticular tubercles rounded to

ovoid in basal outline, flat in form and obtuse in profile;

(12) Toe III larger than Toe V; discs and terminal trans-

verse grooves present on all fingers; tips symmetrical,

disc covers spadate; pads triangular; webbing absent; (13)

coloration highly variable; dorsum tan to light or dark

brown, nearly uniform or suffused with black or reddish

May 2019 | Volume 13 | Number 1 | e176

Arias et al.

a .

ogi as

Fig. 4. In life photographs of (A) Craugastor podiciferus (UCR 23169) from the Caribbean slope of Cerro Kamuk, Costa Rica, (B)

C. blairi (SMF 104032) from Fortuna, Panama, (C) C. sagui sp. nov. (SMF 104018) from La Nevera, Panama, and (D) C. zunigai

sp. nov. (UCR 20389) from Potrero Grande, Costa Rica. Photos by E. Arias (A), A. Hertz (B—C), and E. Boza-Oviedo (D).

pigment; frequently with paired suprascapular dark spots,

area between dorsal folds usually contrasting with flank

color; several specimens with a dark mask from the snout

continuing above the tympanum and often bordered above

by a narrow light line; venter yellow, grayish, or reddish,

uniform or with light or dark spots; usually forelimbs and

hind limbs with dark bars, some specimens with paired

dark spots on anterior surface of hind limbs; upper lip has

dark bars with white pigment in the form of faded bars;

and (14) SVL in males 21—28 mm; SVL in females 23-40

mm.

Comparison: Craugastor podiciferus differs from all other

craugastorids of Isthmian Central America, excluding those

in the C. podiciferus species group, by having unwebbed

toes and a narrow head (HW 37.0-43.3% of SVL). Crau-

gastor podiciferus differs from other members of the C.

podiciferus species group by having the following charac-

teristics (condition for C. podiciferus in parentheses).

Craugastor bransfordii (Cope, 1886), C. gabbi, C. laura-

ster (Savage, McCranie, and Espinal, 1996), C. persimilis

(Barbour, 1926), C. polyptychus (Cope, 1886), C. stejne-

gerianus (Cope, 1893), and C. underwoodi (Boulenger,

1896) differ from C. podiciferus by the following features:

a) dorsum usually granular or warty (dorsum smooth to

scattered tubercles); b) subarticular tubercles projecting

(subarticular tubercles flat); and c) venter coarsely areo-

late, including the midline (venter smooth, at least in mid-

Amphib. Reptile Conserv.

line). Craugastor aenigmaticus Arias, Chaves, and Parra-

Olea, 2018 differs from C. podiciferus by the following

features: a) absence of prominent calcar tubercle on heel,

although some specimens can have one to three small tu-

bercles (prominent calcar tubercle on the heel); b) venter

violet-brown with white blotches (venter yellow, orange,

grayish or olive in adults); and c) white prominent folds

between subarticular tubercles on hands of adults (absence

of white folds between subarticular tubercles on hands of

adults). Craugastor blairi (Barbour, 1928) differs from C.

podiciferus by the following features: a) skin on venter

coarsely areolate (venter smooth, at least in midline); b)

absence of prominent calcar tubercle on heel (prominent

calcar tubercle on heel); c) having accessory palmar tu-

bercles (accessory palmar tubercles absent); and d) having

subarticular tubercles projecting (subarticular tubercles

flat). Craugastor sagui differs from C. podiciferus by the

following features: a) nuptial pads in adult males (nuptial

pads absent); b) absence of prominent calcar tubercle on

heel (prominent calcar tubercle on heel); and c) absence

of vocal slits in adult males (vocal slits in adult males).

Craugastor zunigai differs from C. podiciferus by the fol-

lowing features: a) skin on venter coarsely areolate (venter

smooth, at least in midline); b) absence of prominent cal-

car tubercle on heel (prominent calcar tubercle on heel);

c) accessory palmar tubercles (accessory palmar tubercles

absent), and d) subarticular tubercles projecting (subar-

ticular tubercles flat).

May 2019 | Volume 13 | Number 1 | e176

Craugastor podiciferus species group in Central America

Fig. 5. Variation of the ventral views of the right hands. (A) Craugastor podiciferus (UCR 23175), (B) C. blairi (SMF 104032), (C)

C. sagui sp. nov. holotype (SMF 104018), and (D) C. zunigai sp. nov. holotype (UCR 22703). Photos by E. Arias (A and D) and G.

Kohler (B-C).

Natural history: Craugastor podiciferus inhabits the

lower montane rainforest (Holdridge 1967; Bolafios et al.

2005), which is characterized by a very short dry season

(one to three months), annual precipitation ranging from

3,600—7,500 mm, and annual temperature from 12—17 °C.

Very little is known about the natural history of C. pod-

iciferus; however, it 1s noteworthy that the species was

abundant during the months of fieldwork. Individuals

were always found on the forest floor and observed jump-

ing during the active search. Schlaepfer and Figeroa-Sandi

(1998) described the call of C. podiciferus from Las Cru-

ces, herein referred to as C. podiciferus “Fila Costefia.”

The advertising call of C. podiciferus sensu stricto is un-

known, although it is known to vocalize.

Distribution: Craugastor podiciferus sensu stricto is re-

stricted to the highlands of the Cordillera Volcanica Cen-

tral in Costa Rica and Caribbean slopes of the Cordillera

de Talamanca in Costa Rica and western Panama (Fig. 1).

The altitudinal range of C. podiciferus is 1,700—2,700 m

a.s.l. All known populations of C. podiciferus are found in

primary forests, and several localities are within protected

areas, (1.e., Parque Internacional La Amistad, Parque Nacio-

nal Tapanti-Macizo de la Muerte, Parque Nacional Braulio

Carrillo, and Parque Nacional Juan Castro Blanco). More

fieldwork is necessary to clarify the distribution of C. pod-

iciferus, especially on the northern end of the Cordillera de

Talamanca and in the adjacent zone between the Cordillera

Volcanica Central and the Cordillera de Tilaran.

Amphib. Reptile Conserv.

Remarks: Several populations from the highlands of Cos-

ta Rica are tentatively assigned to Craugastor podiciferus.

However, these populations are phylogenetically struc-

tured and show uncorrected p-distances in the 16S rRNA

gene between 2.45 and 6.37% (Table 1), and some dif-

fer morphologically from typical C. podiciferus and thus

may represent as many as six additional unnamed species.

These six populations are as follows: 1) the C. podiciferus

“Monte Verde” clade, which corresponds to clade A of

Streicher et al. (2009), restricted to Cordillera de Tilaran

and Volcanica Central, at 1,500—1,931 m a.s.l. The speci-

mens in this clade are morphologically very similar to C.

podiciferus sensu stricto. 2) The C. podiciferus “San Ge-

rardo” clade, which corresponds to clade B of Streicher et

al. (2009), restricted to Cordillera de Tilaran and Cordil-

lera Volcanica Central, at 1,470—1,500 m a.s.l. The speci-

mens in this clade differ from C. podiciferus s.s. in having

projecting subarticular tubercles. 3) The C. podiciferus

“Pico Blanco” clade, which contains a single population

from the northern end of the Cordillera de Talamanca in

the Central valley, at 2,242 m a.s.l. The specimens in this

clade differ from C. podiciferus s.s. by having an areo-

late venter and projecting subarticular tubercles. 4) The C.

podiciferus “Fila Costefia” clade, corresponding to clades

E and F of Streicher et al. (2009), restricted to South Pa-

cific Costa Rica, at 1,350—1,550 m a.s.l. The specimens

in this clade differ from C. podiciferus s.s. by having an

areolate venter and accessory palmar tubercles. 5) The C.

podiciferus “Chumacera” clade, which is known for only

May 2019 | Volume 13 | Number 1 | e176

Arias et al.

{

Fig. 6. Dorsal and ventral views in ethanol of Craugastor podiciferus (A, B) syntype (USNM 30672) and (C, D) molecular specimen

reference (UCR 23175). Photos by E.M. Langan (A—B) and E. Arias (C—D).

one population on the Pacific slopes of Cordillera de Ta-

lamanca, at 1,750—1,850 m a.s.l. The specimens in this

clade differ from C. podiciferus s.s. by having accessory

palmar tubercles. 6) The C. podiciferus “Siola,” which is

known for only one population on the Caribbean slope of

the Cordillera de Talamanca, at 1,300—1,350 m a.s.l. The

specimens in this clade differ from C. podiciferus s.s. by

having an areolate venter and by the prominent pungent

calcar tubercle on the heel.

Despite the molecular results and morphological differ-

ences between some of these clades, there is little to dis-

tinguish some populations for particular characters, espe-

cially C. podiciferus sensu stricto, C. podiciferus “Monte

Verde,” and C. podiciferus “Chumacera.” In addition, this

group forms a monophyletic group, thus agreeing with the

definition of the taxon by Savage (2002), Leenders (2016),

and Cossel and Kubicki (2017). For these reasons and until

new morphological evidence support the distinctiveness,

we refrain from raising these clades to the species level.

Craugastor blairi comb. new. (Barbour, 1928)

Common name: Blair’s Dirt Frog

(Figs. 4B and 7)

Holotype: MCZ 13036 from Gutierrez, Bocas del Toro

Province, Panama (near Costa Rican border), collected by

E.R. Dunn and Chester Duryea in summer 1925.

Genetic reference specimen: SMF 104032 (AH 379), an

adult male from Panama: Provincia de Chiriqui: Distrito

Amphib. Reptile Conserv.

de Gualaca: La Fortuna, western slope of Cerro Pata de

Macho (8.6799, -82.193°; 1,793 ma.s.l.), collected by An-

dreas Hertz and Sebastian Lotzkat on 20 May 2010.

Referred specimens: SMF 104030 (AH 377) and SMF

104031 (AH 378), adult males; same date as genetic refer-

ence specimen. SMF 104025 (AH 196), adult female from

Panama: Provincia de Chiriqui: Distrito de Gualaca: La

Fortuna, (8.672°, -82.200°; 1,400 m a.s.l.), collected by

Andreas Hertz and Sebastian Lotzkat on 20 March 2009.

SMF 104026 (AH 238) and SMF 104027 (AH 239), adult

females from Panama: Provincia de Chiriqui: Distrito de

Bugaba: Volcan Bart, Sendero Los Quetzales (8.849°,

-82.515°; 2,134 m a.s.l.), collected by Andreas Hertz and

Sebastian Lotzkat on 8 April 2009.

Assignment to group: Assigned to the genus Craugastor

based on molecular data and on the following characters:

cranial crest absent and Toe III larger than Toe V. Assigned

to the C. podiciferus species group based on the following

features: narrow head (HW/SVL = 34.6-42.8%), dorsum

smooth to scattered tubercles, unwebbed toes, vocal slits

in adult males, and nuptial pads absent.

Diagnosis: The combination of the following character-

istics can be used to distinguish Craugastor blairi (Figs.

5B and 7) from other described species of the genus: 1)

skin on dorsum is smooth or has scattered tubercles; 2)

skin on venter is coarsely areolate; 3) vocal slits and vocal

May 2019 | Volume 13 | Number 1 | e176

Craugastor podiciferus species group in Central America

Fig. 7. Dorsal and ventral views in ethanol of Craugastor blairi (A, B) holotype (MCZ A-13036) and (C, D) molecular specimen

reference (SMF 104032). Photos by the Museum of Comparative Zoology, Harvard University (A—B) and G. Kohler (CD).

sac present in adult males; 4) nuptial pads absent; 5) un-

webbed toes; 6) heel without an enlarged calcar tubercle,

although one to three small tubercles or granules can be

present on heel; 7) accessory palmar and plantar tubercles

present, and absence of supernumerary tubercles; and 8)

subarticular tubercles projecting.

Craugastor (Craugastor) blairi is a small species with

the following characteristics: (1) skin on dorsum smooth

or has scattered tubercles; head smooth; venter coarsely

areolate: flanks smooth with scattered tubercles to warty;

posterior surface of hind limbs surrounding cloaca strongly

areolate; some specimens with a pair of scapular, dorsolat-

eral or lateral folds; discoidal fold complete laterally and

posteriorly; (2) tympanic membrane round, heavily pig-

mented in females, translucent in adult males; prominent

in males, evident in females; annulus evident through the

skin; (TY/ED = 40—-109%); usually with a pair of supra-

tympanic folds; (3) snout subovoid in dorsal view, rounded

in profile; loreal region concave; canthus rostralis usually

rounded; (4) eyelid granular, with an evident supraocular

tubercle and a more or less distinct ridge on outer edge

of eyelid continuous with supratympanic fold (EW/IOD =

37.2—72.7%); cranial crests absent; (5) vomerine teeth in

two transverse fasciculi, behind choanae; choanae smaller

than dentigerous; (6) vocal slits and a single vocal sac that

is large in adult males; nuptial pads absent; (7) Finger I

and II subequal; discs absent, some specimens with ter-

minal transverse grooves on fingers, especially in males;

Amphib. Reptile Conserv.

tips symmetric, usually rounded; pads ovoid to triangular;

(8) fingers lacking lateral fringes; webbing absent; thenar

and palmar tubercles low, ovoid, similar in size; supernu-

merary tubercles absent; 1-2 accessory palmar tubercles;

subarticular tubercles round in basal outline, projecting in

form and globular in profile; (9) ulnar fold absent but tu-

bercles visible; (10) heel without an enlarged calcar tuber-

cle, although one to three small tubercles or granules can

be present on heel; inner edge tarsal smooth, outer with an

incomplete fold and/or tubercles; (11) toes lacking lateral

fringes; inner metatarsal tubercle elongate, outer rounded,

much smaller than inner, inner and outer metatarsal tu-

bercles projecting; supernumerary tubercles absent; sev-

eral low plantar tubercles; subarticular tubercles rounded

to ovoid in basal outline, projecting in form and obtuse

in profile; (12) Toe III larger than Toe V; discs and ter-

minal transverse grooves present on all fingers; tips sym-

metrical, disc covers palmate to spadate; pads triangular;

webbing absent; (13) coloration very variable; dorsum tan

to light or dark brown, nearly uniform or suffused with

black or reddish pigment; frequently paired suprascapular

dark spots; some specimens with dark mask from snout

continuing above tympanum and downward behind axilla,

often bordered above by a narrow light line; venter yellow,

grayish, or reddish, uniform or with light or dark spots;

usually forelimbs and hind limbs with dark bars, some

specimens with paired dark spots on anterior surface of

hind limbs; upper lip with dark bars with white pigment

May 2019 | Volume 13 | Number 1 | e176

Arias et al.

Table 3. Main diagnostic characteristics and character states for secondary sexual characteristics of the species forming the Craugastor

podiciferus species group.

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May 2019 | Volume 13 | Number 1 | e176

184

Amphib. Reptile Conserv.

Craugastor podiciferus species group in Central America

Amplitude

(dB) 0

Frequency (kHz)

Amplitude

Time (s)

Frequency (kHz)

Amplitude

( 0

-5

ile" |

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Fig. 8. Spectrograms and oscillogram of (A) the advertisement call (squeak) and (B) pulsed call (chirp) of Craugastor blairi.

in the form of faded bars; and (14) SVL in males 15.9-21

mm; SVL in females 13.4—30.6 mm.

Comparisons: Craugastor blairi differs from all the other

Isthmian Central America craugastorids (except for those

in the C. podiciferus species group) by having unwebbed

toes and a narrow head (HW 34.642.8% of SVL).

Craugastor blairi differs from other members of the C.

podiciferus species group by having the following char-

acteristics (condition for C. blairi in parentheses). Crau-

gastor bransfordii, C. gabbi, C. lauraster, C. persimilis, C.

polyptychus, C. stejnegerianus, and C. underwoodi differ

from C. blairi by the following features: a) dorsum usu-

ally granular or warty (dorsum smooth to having scattered

tubercles); b) subarticular tubercles obtuse to pointed, at

least the distal subarticular tubercles under Toe III and IV

(subarticular tubercles projecting, globular); and c) altitu-

dinal range, 0-1,600 m a.s.1. (altitudinal range is 1,280-

2,134 maz.s.l. for C. blairi). Craugastor podiciferus differs

from C. blairi by the following features: a) prominent cal-

car tubercle on heel (absence of prominent calcar tubercle

on heel); b) subarticular tubercles flat (subarticular tuber-

cles projecting); and c) venter smooth, at least in midline

(venter coarsely areolate, including midline). Craugastor

aenigmaticus differs from C. blairi by the following fea-

tures: a) venter smooth (venter coarsely areolate); b) sub-

articular tubercles flat (subarticular tubercles projecting,

globular); and c) prominent white folds between subartic-

ular tubercles (absence of white folds between subarticular

tubercles). Craugastor sagui differs from C. blairi by the

following features: a) venter smooth (venter coarsely areo-

late); b) nuptial pads in adult males (nuptial pads absent);

c) absence of vocal slits (vocal slits in adult males); and

d) subarticular tubercles flat (subarticular tubercles pro-

jecting). Craugastor zunigai differs from C. blairi by the

absence of an evident supraocular tubercle (eyelid with an

evident supraocular tubercle).

Amphib. Reptile Conserv.

185

Natural history: Craugastor blairi inhabits the lower

montane rainforest (Holdridge 1967; Bolafios et al. 2005),

which is characterized by a very short dry season (one to

three months). Annual precipitation ranges from 3,600-—

7,500 mm and annual temperature from 12-17 °C. Very

little is known about the natural history of C. blairi; how-

ever, it is noteworthy that the species was abundant dur-

ing the months of fieldwork. The specimens were always

found on the floor in leaf litter, possibly active during the

day. Males call during periods of lower light due to dark

clouds or in evening hours. Calling activity is greater dur-

ing rain.

Vocalization: Vocalizations of four male specimens (AH

375, 1,430 m, dusk, 19.5 °C, 100% RH, AH 377-379,

1,793 m, 18.5 °C, 100% RH) have been recorded on the

slopes of Cerro Pata de Macho. All specimens were call-

ing at dusk, between 18:00 h and 19:00 h after a moderate

rain. Calling sites were elevated positions only a few cen-

timeters above the ground such as low vegetation, twigs,

or roots. Calling stopped completely after dark. One male

(AH375) was recorded at 19.5 °C and 100% relative hu-

midity. The other three males were recorded at 18.5 °C

and 100% relative humidity. Two very different call types

could be distinguished (Fig. 8). The first that was emitted

by all four males was a “squeak” of 0.035—0.065 (0.050 +

0.007) seconds in length, given in frequent intervals after

7.938—-29.124 (11.974 + 4.825) seconds, resulting in a call

rate of 4.66—9.63 (6.67 + 2.41) calls per minute. The call

has two harmonics, the fundamentals of which also con-

tain the dominant frequency at 6,844—8,063 (7,306 + 313)

Hz. The dominant frequency is reached in the middle or

towards the end of the call, 0.009—0.053 (0.030 + 0.012)

seconds after ignition of the call. The frequency range of

the fundamental harmonic is between 4,705—6,674 (5,680

+ 634) Hz low frequencies and 7,181—9,343 (7,965 + 467)

Hz high frequencies. The higher harmonic has a frequency

range of 9,566—14,057 (12,057 + 1,159) Hz low frequen-

May 2019 | Volume 13 | Number 1 | e176

Arias et al.

Fig. 9. Dorsal and ventral views in ethanol of the holotype (SMF 104018) of Craugastor sagui sp. nov. Photos by G. Kohler.

cies and 14,182—16,617 (15,175 + 650) Hz high frequen-

cies, with the highest acoustic intensity at 13,313-15,656

(14,147 + 626) Hz.

The second call type could only be recorded in two of

the four males. It was identified as a trilling, 1e., strongly

pulsed “chirp” composed of mostly two, or less often one

or three, notes. The notes are 0.266—0.692 (0.450 + 0.116)

seconds in length, and the total call duration is 0.745—

1.551 (0.914 + 0.206), depending on how many notes are

contained in the call. Calls are repeated after 5.911—18.265

(10.196 + 3.522) seconds. A single note contains 12—25

(19 + 4) pulses that are emitted at a pulse rate of 36-46

(43 + 2) pulses per second. The dominant frequency of

8,813—10,313 (9,654 + 236) Hz is reached in the middle

or towards the end of the note at pulse 6-21 (13 + 5). Calls

are frequency modulated with the lowest frequencies of

6,044—7,640 (6,617 + 508) Hz in the first pulses gradually

rising towards 10,298—12,002 (10,848 + 466) Hz high fre-

quencies in the last pulses of each note.

Distribution: Craugastor blairi is restricted to western

Panama, on the Cordillera Central from Bart Volcano to

the east over the La Fortuna depression into the Serrania

de Tabasara. The specimens were collected around Baru

Volcano, Cerro La Estrella, Cerro Guayaba, Guayabito,

and La Fortuna (Fig. 1). The altitudinal range of the spe-

cies 1s 1,280—2,134 m a.s.l. To our current knowledge, the

continuous distribution of C. blairi is interrupted between

La Fortuna and Guayabito. Both sites are separated by ~80

airline kilometers. Within this gap, another distinct clade

was found that seems restricted to the surroundings of

Cerro Sagui and Cerro Santiago, and is described below as

C. sagui. More fieldwork will be necessary to clarify the

distribution of C. blairi with respect to C. sagui, especially

in the area between La Fortuna and Guayabito. It would

also be interesting to explore the eastern distribution limits

of C. blairi, as far east as Santa Fé, Veraguas.

Craugastor sagui sp. nov.

urn: Isid:zoobank.org:act:27F02325-D86E-4A 4E-93 11-B6FDD977C1E0

Amphib. Reptile Conserv.

Common name: Sagui Dirt Frog

(Figs. 4C and 9)

Holotype: SMF 104018 (AH481), an adult male from

Panama: Comarca Ng6be-Bugleé: Distrito de Nole Duima:

southeastern slope of Cerro Sagui (8.563°, -81.821°; 1,991

m a.s.l.), collected by Andreas Hertz and Sebastian Lotz-

kat on 9 September 2010.

Paratypes: SMF 104014 (AH483), adult female; same

date as the holotype. SMF 104017 (AH342), adult male

and SMF 104015 (AH045), adult female from Panama:

Comarca Ngdébe-Bugleé: Distrito de Nole Duima: La Ne-

vera, southern slopes of Cerro Santiago (8.500°, -81.772°;

1,700 m a.s.l.), collected by Andreas Hertz and Sebastian

Lotzkat on 13 November 2009.

Assignment to group: Assigned to genus Craugastor

based on our molecular data and on the following morpho-

logical characters: cranial crests absent, and Toe III lon-

ger than Toe V. Assigned to C. podiciferus species group

based on the following features: narrow head (HW/SVL =

35.8-40.9%), length of Finger I equal to Finger II, dorsum

smooth, toes unwebbed, and nuptial pads in adult males.

Diagnosis: The combination of the following characteris-

tics can distinguish Craugastor (Craugastor) sagui (Figs.

AC, 5C, and 9) from the other described species in the ge-

nus: 1) skin on dorsum smooth; 2) skin on venter smooth;

3) vocal slits absent; 4) nuptial pads in adult males; 5) un-

webbed toes; 6) heel without an enlarged calcar tubercle,

although one to three small tubercles or granules may be

present on heel; 7) accessory palmar, plantar, and supernu-

merary tubercles absent; and 8) subarticular tubercles flat.

Comparison: Craugastor sagui differs from all other Isth-

mian Central America craugastorids (except for those in C.

podiciferus species group) by having unwebbed toes and a

narrow head (HW 35.8-40.9% of SVL). Craugastor sagui

differs from other members of the C. podiciferus species

May 2019 | Volume 13 | Number 1 | e176

Craugastor podiciferus species group in Central America

group by having the following characteristics (condition

for C. sagui in parentheses). Craugastor bransfordii, C.

gabbi, C. lauraster, C. persimilis, C. polyptychus, C. stej-

negerianus, and C. underwoodi differ from C. sagui by the

following features: a) dorsum usually granular or warty

(dorsum smooth to shagreen with scattered tubercles); b)

subarticular tubercles projecting (subarticular tubercles

flat); and c) venter coarsely areolate, including the midline

(venter smooth, at least in midline). Craugastor podicifer-

us differs from C. sagui by the following features: a) prom-

inent calcar tubercle on heel (Fig. 6) (calcar tubercle ab-

sent, although some specimens can have one to three small

tubercles); b) vocal slits present in adult males (vocal slits

absent); and c) absence of nuptial pads (nuptial pads pres-

ent in adult males). Craugastor aenigmaticus differs from

C. sagui by the following features: a) venter coloration of

violet-brown with white blotches (venter yellow, orange,

grayish, or olive in adults); b) white prominent folds be-

tween subarticular tubercles on hands of adults (absence

of white folds between subarticular tubercles on hands of

adults); and c) absence of nuptial pads (nuptial pads present

in adult males). Craugastor blairi differs from C. sagui by

the following features: a) venter coarsely areolate (venter

smooth); b) absence of nuptial pads (nuptial pads present

in adult males); c) vocal slits present in adult males (vocal

slits absent in adult males); and d) projecting subarticular

tubercles (flat subarticular tubercles). Craugastor zunigai

differs from C. sagui by the following features: a) venter

coarsely areolate (venter smooth); b) subarticular tubercles

projecting (subarticular tubercles flat); c) absence of nup-

tial pads (nuptial pads present in adult males); and d) vocal

slits in adult males (vocal slits absent).

Description of the holotype: Adult males have an SVL

of 18.7 mm (Figs. 4-5). Head relatively narrow, HW =

35.8% of SVL; snout subovoid in the dorsal view, rounded

in profile; snout relatively long (HL = 7.0 mm, 37.4% of

SL), with nostrils directed laterally; in the ventral view, tip

of the snout protruding slightly beyond the edge of lower

lip. Internarial area convex; canthus rostralis rounded;

intercanthal area flat; loreal region slightly concave; vo-

cal slits absent. Eye moderate (EN/ED = 74.07%), not

protruding beyond dorsal and ventral outline of head, di-

rected laterally. Tympanic membrane distinct, covered in

skin; tympanic annulus prominent, round, relatively large

(77.77% of ED). Skin on all dorsal and lateral surfaces

of head smooth to shagreen. Upper eyelid granular, with-

out superciliar or supraocular tubercles but with a more or

less discernible ridge on outer edge of eyelid continuous

with supratympanic fold and downward behind the axilla.

Postrictal tubercles fused to form a short ridge postero-

ventral to tympanum. Skin on dorsum and limbs smooth to

shagreen. Skin of chest and throat smooth, venter smooth;

ventral surfaces of thighs smooth; skin of groin nearly

smooth. Flanks shagreen, especially along antero-ventral

flank region. Discoidal fold complete.

Amphib. Reptile Conserv.

Forelimb relatively short and slim; fingers moderately

long and slim without lateral fringes. Discs absent; fin-

gers with grooves; tips of fingers unexpanded, rounded in

dorsal view; pads ovoid. Supernumerary tubercles absent;

accessory palmar tubercles absent; subarticular tubercles

rounded in basal outline, flat in form and globular in pro-

file; thenar and palmar tubercles elongate and flat; thenar

and palmar tubercles similar in size. Ulnar fold absent but

several tubercles visible. Fingers not webbed.

Legs relatively long and slim; heel granular, lacking

enlarged tubercles. Discs and grooves on all toes, palmate

on Toe IV, spadate on others; pads triangular on Toe IV,

ovoid in others. Supernumerary and plantar tubercles ab-

sent; subarticular tubercles rounded in basal outline, flat in

form and obtuse in profile; inner metatarsal tubercle elon-

gate, globular; outer metatarsal tubercle rounded, globu-

lar; outer much smaller than inner; outer edge tarsal with

several tubercles, inner smooth. Cloacal opening directed

posteriorly at mid-level of thighs.

Coloration of the holotype in life: Dorsal background

color dark orangeish-brown with dark blotches, and a dark

interorbital bar. Dorsal surfaces of legs and arms with dark

bars, anterior surface of legs with white spots. Upper lip

brown-orange with dark bars and scattered white spots.

Flanks with a similar dorsal coloration but paler orange,

with fine white-bluish mottling. Ventral surface of body

and legs dark reddish-brown with bluish-white pigment

forming blotches; ventral surface of throat similar to ven-

ter but with fewer pale spots. Soles of feet and hands dark

brown with cream-colored tubercles.

Coloration of the holotype in ethanol: After eight years

in ethanol (70%), the overall dark orangeish-brown on

dorsum faded to pale cream-pinkish with dark brown

blotches. The dark orangeish-brown on ventral surface of

the body and legs faded to dark brown.

Measurements of the holotype (mm): SL 18.7; HL 7.0;

HW 6.7; IOD 2.7; EW 1.4; EN 2.0; ED 2.7; TY 2.1. Mea-

surements in related percentages: EW/IOD 51.85%; IOD/

HW 40.3%; TY/ED 77.78%; EN/ED 74.07%; ED/HL

38.57%; HL/HW 104.48%; EN/HL 28.57%.

Etymology: The species was discovered on Cerro Sagui,

in western Panama. The specific name is a noun in the ap-

position.

Natural history: Craugastor sagui inhabits the lower

montane rainforest (Holdridge 1967; Bolafios et al. 2005),

characterized by a very short dry season (one to three

months), annual precipitation ranging from 3,600—7,500

mm and annual temperature from 12—17 °C. Very little is

known about the natural history of C. sagui; however, it

is noteworthy that it was relatively abundant during the

months of fieldwork. The specimens were exclusively

May 2019 | Volume 13 | Number 1 | e176

Arias et al.

Fig. 10. Dorsal and ventral views in ethanol of the holotype (UCR 22703) of Craugastor zunigai sp. nov. Photos by E. Arias.

found in the leaf litter on the forest floor, and also typically

observed jumping during the active search. We never re-

corded a vocalization that could be attributed to C. sagui;

however, it is possible that it does vocalize.

Distribution: Craugastor sagui is restricted to western

Panama, on the pacific slopes of Cordillera Central. The

specimens were collected on the slopes of Cerro Sagui and

Cerro Santiago (Fig. 1). The known altitudinal range is

1,700—-1,991 ma.s.l.

Craugastor Zunigai sp. nov.

urn:Isid:zoobank.org:act:70A61278-3E9F-454D-8F08-436F453F7116

Common name: Zuniga’s Dirt Frog

(Figs. 4D and 10)

Holotype: UCR 22703 (EAP 0618), adult male from Cos-

ta Rica: Provincia de Puntarenas: Canton de Coto Brus:

Distrito de Sabalito: Finca Las Alturas, Zona Protectora

Las Tablas, (8.976°, -82.834; 1,732 m a.s.l.), collected by

Erick Arias, Gerardo Chaves, and Omar Zufiiga on 15

September 2015.

Paratypes: UCR 23014 (EAP 0725), UCR 23016 (EAP

0727), UCR 23017 (EAP 0728), and UCR 23018 (EAP

0729), adult females from Costa Rica: Provincia de Pun-

tarenas: Canton de Buenos Aires: Distrito de Potrero

Grande: Tres Colinas, Parque Internacional La Amis-

tad, (9.123°, -83.066°; 1,846 m a.s.l.), collected by Erick

Arias, Fanny Hernandez, and Omar Zufiga on 10 Sep-

tember 2016. UCR 23176 (EAP 0831), adult male and

UCR 23177 (EAP 0832), adult female from Costa Rica:

Provincia de Puntarenas: Canton de Coto Brus: Distrito

de Pittier: Santa Maria de Pittier, Parque Internacional La

Amistad, (9.031°, -82.962; 1,920 maz.s.l.), collected by Er-

ick Arias and Omar Zufiga on 27 March 2018.

Assignment to group: Assigned to the genus Craugas-

tor based on molecular data and on the following char-

Amphib. Reptile Conserv.

acters: cranial crests absent and Toe III larger than Toe V.

Assigned to the C. podiciferus species group based on our

phylogeny and on the following characters: narrow head

(HW/SVL = 35.8-41.9%), dorsum smooth to shagreen

with scattered tubercles, unwebbed toes, vocal slits in

adult males, and absence of nuptial pads.

Diagnosis: The combination of the following character-

istics can be used to distinguish Craugastor (Craugastor)

zunigai (Figs. 4D, 5D, and 10) from other described spe-

cies in the genus: 1) skin on dorsum smooth to shagreen

with scattered tubercles; 2) skin on venter coarsely areo-

late; 3) vocal slits present in adult males; 4) nuptial pads

absent; 5) unwebbed toes; 6) heel without a projecting

tubercle, although one to three low tubercles or granules

can be present; 7) accessory palmar and plantar tubercles

present, usually with no supernumerary tubercles under

the digits; and 8) subarticular tubercles projecting.

Comparison: Craugastor zunigai differs from all other

Isthmian Central America craugastorids (except for those

in the C. podiciferus species group) by having unwebbed

toes and a narrow head (HW 35.8—-41.9% of SVL). Crau-

gastor zunigai differs from other members of the C.

podiciferus species group by having the following char-

acteristics (condition for C. zunigai in parentheses). Crau-

gastor bransfordii, C. gabbi, C. lauraster, C. persimilis, C.

polyptychus, C. stejnegerianus, and C. underwoodi differ

from C. zunigai by the following features: a) dorsum usu-

ally granular or warty (dorsum smooth to shagreen with

scattered tubercles); b) subarticular tubercles obtuse to

pointed, at least distal subarticular tubercles under Toe III

and IV (subarticular tubercles projecting, globular); and

c) altitudinal range, 0—1,600 m a.s.l. (altitudinal range is

1,500-2,100 m a.s.l. for C. zunigai). Craugastor podic-

iferus differs from C. zunigai by the following features: a)

prominent calcar tubercle on heel (Fig. 6) (calcar tubercle

absent, although some specimens can have one to three

small tubercles in C. zunigai); b) venter smooth (venter

coarsely areolate); c) subarticular tubercles flat (subar-

ticular tubercles projecting); and d) absence of accessory

May 2019 | Volume 13 | Number 1 | e176

Craugastor podiciferus species group in Central America

palmar tubercles (accessory palmar tubercles present).

Craugastor aenigmaticus differs from C. zunigai by the

following features: a) venter smooth (venter coarsely areo-

late); b) subarticular tubercles flat (projecting subarticular

tubercles, globular); and c) prominent white folds between

subarticular tubercles (absence of white folds between

subarticular tubercles). Craugastor blairi differs from C.

zunigai by the presence of an evident supraocular tubercle

(eyelid smooth to granular but without an evident supraoc-

ular tubercle). Craugastor sagui differs from C. zunigai by

the following features: a) nuptial pads in adult males (nup-

tial pads absent); b) absence of vocal slits (vocal slits in

adult males); c) venter smooth (venter coarsely areolate);

and d) flat subarticular tubercles (projecting subarticular

tubercles, globular).

Description of the holotype: Adult male having an SVL

of 19.8 mm (Fig. 10). Head relatively narrow, HW =

35.86% of SVL; snout subovoid in the dorsal view, round-

ed in profile; snout relatively long (HL = 7.6 mm, 38.38%

of SL), with nostrils directed laterally; in ventral view, tip

of snout protruding markedly beyond edge of lower lip.

Internarial area convex (IN 2.2 mm); canthus rostralis

rounded; intercanthal area flat IC = 3.6 mm); loreal re-

gion slightly concave; vomerine teeth transverse, in two

fascicles behind choanae. Tongue round in shape, lacking

a distinct posterior notch; teeth absent; choanae moder-

ately large, rounded on the posterior half but flat on ante-

rior half, hemispherical; paired elongate vocal slits under

posterolateral margins of tongue and a single internal sub-

gular vocal sac. Eye moderate (EN/ED = 84.1%), protrud-

ing beyond dorsal outline of head in ventral view, directed

laterally. tympanic membrane prominent, translucent, and

slightly pigmented; tympanic annulus prominent, round,

large (134% of ED). Skin on dorsal and lateral surfaces of

head smooth. Upper eyelid smooth, without superciliar or

supraocular tubercles but with a more or less discernible

ridge on outer edge of eyelid continuous with supratym-

panic fold and downward behind axilla. Elongate and pro-

jecting postrictal tubercle, postero-ventral to tympanum.

Skin on dorsum and limbs smooth to shagreen with a pair

of incomplete dorso-lateral folds, extending from axillary

to inguinal level. Skin of chest and throat smooth, ven-

ter coarsely areolate with low granules; ventral surfaces

of thighs areolate; skin of groin smooth. Flanks shagreen

with scattered tubercles to areolate, especially along an-

tero-ventral flank region. Discoidal fold complete.

Forelimb relatively short and robust; fingers moder-

ately long and slim without lateral fringes. Discs absent;

fingers I-IV with grooves; tips of fingers unexpanded,

rounded in dorsal view; pads ovoid. Supernumerary tu-

bercles absent; four small and rounded accessory palmar

tubercles; subarticular tubercles rounded in basal outline,

slightly projecting in form, and globular in profile; thenar

tubercle elongate and palmar tubercle rounded, flat, simi-

lar in size. Ulnar fold absent but some tubercles visible.

Fingers not webbed.

Amphib. Reptile Conserv.

Legs relatively long and slim (TL = 55.3% SVL); heel

smooth with two barely visible low granules, lacking a

projecting tubercle. Discs and grooves on all toes, palmate

on Toe IV, spadate in others; pads triangular on Toe IV,

ovoid in others. Supernumerary tubercles absent; plantar

tubercles small and rounded; subarticular tubercles ovoid

in basal outline, projecting in form, and obtuse in profile;

inner metatarsal tubercle elongate, projecting; outer meta-

tarsal tubercle rounded, globular; outer metatarsal tubercle

much smaller than inner; outer edge tarsal with an indis-

tinct short ridge and low granules, inner smooth. Cloacal

opening directed posteriorly at mid-level of thighs.

Coloration of the holotype in life: Dorsal background

color dark brown suffused laterally with pale brown, head

uniform dark brown. Dorsal surfaces of legs and arms with

dark bars, posterior surface of legs cream suffused with

red. Cloacal opening darker than posterior surface of legs.

Flanks pale brown, groin suffused with red. Ventral sur-

face of body and legs yellowish-cream with dark pigment;

throat cream with dark pigment. Absence of bars on lips,

mask, and blotches on dorsal surface.

Coloration of the holotype in ethanol: After three years

in ethanol (70%), the overall dark brown on dorsum re-

mained very similar to that in life. The yellowish-cream

color on ventral surface of body and legs faded to pale

brown.

Measurements of the holotype (mm): SL 25.9; HL 10.3;

HW 10.3; IOD 3.7; EW 1.7; EN 2.5; ED 2.9; TY 1.9. Mea-

surements in related percentages: EW/IOD 68.82%; IOD/

HW 28.18%; TY/ED 54.12%; EN/ED 81.12%; ED/HL

27.24%; HL/HW 107%; EN/HL 24.34%.

Variation: The morphometric variation is summarized

in Table 2. Craugastor zunigai shows a relatively high

level of intraspecific polymorphism. In some specimens,

a pair of lateral folds is present, and some specimens show

a supraocular tubercle. In some specimens, two unfused

postrictal tubercles are visible. Palmar and thenar tuber-

cles are equal in size in some specimens, thenar slightly

smaller than palmar in others. The palmar tubercle is

heart-shaped in some specimens and ovoid in others. UCR

20389 with a pair of lateral folds from the axillar level to

cloaca bordered by black pigment, the area between folds

and from snout to cloaca a lighter, cream-yellowish color

in ethanol. UCR 20428 with a pattern of dark brown on the

dorsum with a lighter interorbital mark and area anterior to

it more pale than dorsum. Mask absent in some specimens.

Throat has a nearly uniform cream color in UCR 20401,

heavily mottled in UCR 20421, and nearly uniform dark

brown in UCR 20389. Venter ranges from nearly uniform

cream color without dark pigment in UCR 20401 to heav-

ily mottled in UCR 20389.

Etymology: The name zunigai is a patronym, honoring

the field-guide Omar Zufiiga in recognition of his im-

May 2019 | Volume 13 | Number 1 | e176

Arias et al.

portant aid during the fieldwork within the Cordillera de

Talamanca. Omar Zufiiga took part in the fieldwork that

yielded the specimens from Tres Colinas, Las Alturas, Las

Tablas, and Santa Maria de Pittier.

Natural history: Craugastor zunigai inhabits the lower

montane rainforest (Holdridge 1967; Bolafios et al. 2005),

which is characterized by a very short dry season (one to

three months), annual precipitation ranging from 3,600—

7,500 mm and annual temperature from 12-17 °C. Very

little is known about the natural history of C. zunigai;

however, is noteworthy that the species was relatively

abundant during the months of fieldwork. The specimens

were always found on the forest floor jumping among the

leaf litter during the active search. We never recorded a vo-

calization that could be attributed to C. zunigai; however,

it is possible that it vocalizes. At Las Alturas, C. zunigai

occurs very near the known C. gabbi localities; however,

we did not find them in sympatry since they are not known

to overlap altitudinally. In Tres Colinas, the above-men-

tioned pattern is similar to C. stejnegerianus.

Distribution: Craugastor zunigai 1s restricted to south-

western Costa Rica, on the Pacific slopes of Cordillera

de Talamanca. The specimens were collected from Santa

Maria de Pittier, Tres Colinas, Las Alturas de Coton, and

road to Las Tablas (Fig. 1). The altitudinal range of the

new species is 1,500—2,100 m a.s.l. All populations were

found in primary forests, the populations from Santa Maria

de Pittier and Tres Colinas are in La Amistad International

Park and the population from Las Alturas de Coton and

road to Las Tablas is in Zona Protectora Las Tablas. The

known populations of C. zunigai are fragmented over ~43

km. This species was not found on the Pacific slope of

Cerro Utyum and Cerro Durika (to the west). Addition-

al fieldwork is necessary to clarify the distribution of C.

zunigai. The locality “road to Las Tablas” is very close to

the Costa Rica-Panama border, so C. zunigai is likely also

present in Panama.

Discussion

With the recognition of Craugastor blairi, C. zunigai, and

C. sagui, the C. podiciferus species group is now formed

by 12 species, all of which are collectively distributed

in Costa Rica and western Panama (Savage 2002;

AmphibiaWeb 2019). The species in this group have been

difficult to delineate historically because of morphological

variability between and within populations. However,

using molecular sequence data, we found well-supported

clades and large genetic distances between several of these

populations (Table 1). Although the sole use of genetic

distances for species delimitation is not recommended,

we believe that the combination of large genetic distances

and close geographic proximity between phylogenetically

related species provide an important measure to identify

cryptic species with a conservative morphology. The

Amphib. Reptile Conserv.

genetic distances presented herein between members of

the C. podiciferus species group are above the thresholds

of 3% in the 16S rRNA and 10% in the COI mitochondrial

genes suggested by Fouquet et al. (2007) and Vences et al.

(2005). For amphibians, the 16S rRNA gene fragment has

been suggested as a DNA barcode marker for amphibian

diversity inventories (Vences et al. 2005) to complement

the standard COI-5’ marker used for animals in general

(Smith et al. 2008). Although the species recognized

herein are not cryptic sensu stricto (Pérez-Ponce de Leon

and Nadler 2010), they are in taxonomic practice. As

suggested by Pérez-Ponce de Leon and Nadler (2010), the

use of molecular data within morphologically conserved

groups allow delineation of taxa that otherwise would be

considered a single taxon. The use of phylogenies based

on molecular data has been essential to solve taxonomic

problems in this group.

Taylor (1952) synonymized C. blairi under C. podic-

iferus without further discussion, nor was it discussed in

the comprehensive seminal work of Savage (2002). The

results presented here support the validity of Craugastor

blairi (previously shown as Craugastor sp. B), and it is

not a sister to C. podiciferus (Crawford and Smith 2005;

Streicher et al. 2009). However, C. blairi and C. podic-

iferus are morphologically similar and show a high level

of polymorphism, and thus the taxonomic decision of Tay-

lor (1952) is not surprising given the technical limitations

at the time of that publication. The distinctiveness of C.

blairi is provided by its phylogenetic position. The same

rules apply to C. sagui, the phylogenetic position of which

provides significant evidence for its distinct evolutionary

trajectory as a species beyond the observation that it re-

sembles C. podiciferus and C. blairi morphologically. The

morphological similarity among nonsister species can be

explained either as plesiomorphy or convergence (Castro-

viejo-Fisher et al. 2017). All species analyzed herein have

inhabiting the ground leaf litter in highland forest habitats

(lower montane zonation) in common, which has likely

played a role in the maintenance of this morphology.

A more complicated situation lies in the recognition of

Craugastor zunigai, given its morphological resemblance

to its sister species C. blairi. Although the morphological

divergence between C. blairi and C. zunigai is weak, we

believe that both are valid species supported by phylo-

genetic distinctiveness based on mitochondrial analysis.

These species are geographically very close (~10 airline

km); thus, it seems unlikely that the large genetic distances

(2.9% in 16S and 14.3% in COI) are explained by isolation

due to distance. Our taxonomic decision to recognize these

two as separate species will be strengthened with addition-

al evidence, especially the comparison of mating calls of

C. zunigai with the call of C. blairi. Future sampling ef-

forts must also be conducted on the highlands of extreme

western Panama and adjacent Costa Rica between Bart

Volcano and Cerro Pando to explore the species distribu-

tion limits and possible contact zones of C. aenigmaticus,

C. blairi, C. zunigai, and C. podiciferus.

May 2019 | Volume 13 | Number 1 | e176

Craugastor podiciferus species group in Central America

The presence of several additional species within Crau-

gastor podiciferus is uncertain. Streicher et al. (2009) sug-

gested that the current concept of C. podiciferus could

be masking six distinct species that are geographically

structured, but with two instances of sympatry. However,

the findings of Streicher et al. (2009) and our own results

agree with C. podiciferus being a monophyletic entity;

thus, there is currently no need to split C. podiciferus into

several species until such division is supported by detailed

morphological or other evidence. The performance of an

extensive, integrative analysis of the C. podiciferus spe-

cies complex is strongly suggested to reevaluate the taxo-

nomic status of its different genetic clades. For now, the

recognition of C. podiciferus sensu lato as a single species

results in a highly variable morphological group, given

that even those few useful characters (i.e., skin on ven-

ter, subarticular tubercles, vocal slits, and nuptial pads) are

highly variable between populations. The clades shown

herein should be reevaluated using other types of data that

can provide evidence of lineage divergence, such as geo-

graphical distribution, ecological niche, mating calls, or

detailed morphometric data.

The members of the Craugastor podiciferus species

group have qualities such as high abundance, broad dis-

tribution (collectively), and high genetic diversity that

makes them suitable for use in various studies in ecology

and evolution. However, these species have been poorly

studied, likely because of the difficulty in clearly identify-

ing the species. Therefore, it is necessary to clarify the tax-

onomy of this species group. Upon revisiting study objec-

tives, the following conclusions are offered: 1) Craugastor

podiciferus sensu stricto 1s restricted to those populations

at Cordillera Volcanica Central, Costa Rica and Cordil-

lera de Talamanca (Caribbean slopes) in Costa Rica and

western Panama; 2) seven well-supported clades are found

within the current concept of C. podiciferus, and these

seven clades require extensive taxonomic revision; and 3)

the populations from southwestern Costa Rica and western

Panama are grouped in three clades, one potentially refer-

ring to an existing name, C. blairi, which is resurrected

herein, and two representing new species that are herein

named C. sagui from western Panama and C. zunigai from

southwestern Costa Rica.

Acknowledgements.—We thank Laura Marquez-

Valdelamar and Andrea Jiménez-Marin for their labo-

ratory assistance and Federico Bolafos for the use of

specimens from the Museo de Zoologia of the Universi-

dad de Costa Rica. We thank Linda Acker and Gunther

Kohler of the Senckenberg herpetology collection (SMF)

for providing photographs of some of the specimens ex-

amined herein. Sebastian Lotzkat, Frank Hauenschild,

Nadim Hamad, Omar Zuniga, Olmer Cordero, Justo

Layam Gabb, and Xavier Baltodano provided valuable

assistance in the field during the expeditions. We thank

Rogelio Moreno, former general chief of the Ngdbe-

Buglé indigenous community, for granting us access to

Amphib. Reptile Conserv.

the Comarca Ngdébe-Buglé, as well as all the Ngdbe and

Buglé who helped us logistically during the field work.

AH is thankful to the owners and staff of the Lost and

Found Ecohostel in Valle de las Minas, Chiriqui for their

enduring support and hospitality. EA thanks the Posgra-

do en Ciencias Bioldgicas for its support of this study,

the CONACyT for the student grant (CVU/Becario)

626946/330343, and the Programa de Innovacion y Capi-

tal Humano para la Competitividad PINN-MICITT for

the student grant (PED-0339-15-2). The laboratory work

was funded by a grant from PAPITT-UNAM (IN203617)

to GP-O. The fieldwork was partially supported by the

National Geography Society (Grant number W-346-14).

We acknowledge the Costa Rican Ministry of Environ-

ment and Energy (MINAE) for providing the corre-

sponding scientific collecting permits for this expedition

(SINAC-SE-GAS-PI-R 007-2013 and 59-2015). Collect-

ing permits for Panama SE/A-30-08, SC/A-8-09, SC/A-

28-09, and SC/A-21-10, as well as the corresponding ex-

portation permits, were issued by the Direccion de Areas

Protegidas y Vida Silvestre of the Autoridad Nacional del

Ambiente (ANAM), recently renamed the Ministerio de

Ambiente (MiAmbiente), Panama City, Panama.

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Amphib. Reptile Conserv.

Erick Arias is a Costa Rican herpetologist; who earned a bachelor’s degree in biology at

Universidad de Costa Rica in 2013. Erick then joined Parra’s Lab at Universidad Nacional

Autonoma de México (UNAM) where he received a doctorate in systematics in 2019 focusing

on molecular systematics, taxonomy, and biogeography of the Craugastor podiciferus species

group. Erick’s current research focuses on the systematics, taxonomy, and biogeography of the

amphibians and reptiles of Costa Rica, especially of direct-developing frogs and salamanders.

Andreas Hertz earned a diploma degree in biology at Goethe-University, in cooperation

with the Senckenberg Research Institute in Frankfurt am Main, Germany, where he studied

reptiles and amphibians from Venezuela. In a subsequent doctorate, Andreas worked on the

systematics, taxonomy, and conservation of Panamanian amphibians. Andreas then joined

University of Massachusetts Boston (USA) as a DFG postdoctoral fellow to work on amphibian

chytridiomycosis. His current research is on recovering amphibian populations in upland areas

of Panama that have survived mass extinction through chytridiomycosis. This research focuses

on the mechanisms of host adaption in recovering frog populations which allow coexistence

with the pathogen.

Gabriela Parra Olea earned a Ph.D. at University of California, Berkeley working on the

molecular systematics of neotropical salamanders. In 2001, Gabriela obtained a full-time

research position at the Instituto de Biologia, UNAM in Mexico City. Her current research

focuses on molecular systematics, taxonomy, and conservation of neotropical amphibians,

especially salamanders. A second line of Gabriela’s research is based on studying the impact of

chytridiomycosis on the conservation of Mexican amphibians.

193 May 2019 | Volume 13 | Number 1 | e176

Arias et al.

Appendix I. Institutional voucher numbers, locality information, and GenBank accession numbers for the specimens used in the

molecular phylogenetic analyses. Museum collection acronyms follow Frost (2019) with the addition of EAP to refer to Erick

Arias field numbers and CRARC to refer to the Costa Rica Amphibian Research Center private collection. CR = Costa Rica, HN =

Honduras, NI = Nicaragua, PA = Panama.

: Institutional Collection Elevation | Geographic coordinates GenBank Number

Species .

vouchers locality (m) Lat Lon 16S COI

Changuinola,

C. aenigmaticus SMF: 104020 Bocas del 2388 8.9139 -82.7088 MK211615 MK211577

Toro, PA

Telire,

C. aenigmaticus UCR: 21951 Talamanca, 2700 9.3488 -83.1750 MK211616 MK211578

Buenos Aires,

C. aenigmaticus UCR: 22737 Puntarenas, 2660 9.3224 -83.2028 MK211617 MK211579

Ls, FMNH: Gualaca,

C. blairi 157689 Chirigili; PA 1000 8.7500 -82.2170 EF562353 a

C. blairi SMF: 102024 iualaca, 1730 8.6775 -82.1980 MK211627. MK211583

Chiriqui, PA

C. blairi SMF: 104023 Cualaca, 1280 8.6781 -82.2101 MK211628 MK211584

Chiriqui, PA

al Bugaba,

C. blairi SMEF: 104027 ie a 2134 8.8494 -82.5154 MK211629 MK211585

Chiriqui, CR

C. blairi SMF: 104033 oualaca. 1456 8.6740 -82.2154 MK279367 —

Chiriqui, PA

Nurum,

C. blairi SMF: 104034 Negdbe Bugle, 1541 8.5512 -81.4833 MK279368 a

C. blairi SMF: 104037 _Doduete, 1952 8.7757 -82.3901 MK279369 ome

Chiriqui, PA

USNM: Boquete,

C. blairi 563039 Chiriqui, PA 1663 8.3136 -82.4000 EF562356 —

C. bransfordii UCR: 22269, Alajuela, 466 10.3121 -84.1778 KT950295. = MK211571

Alajuela, CR

C. bransfordii UcR:saeag - Dtauimes, 537 10.0595 -83.5452. | _MK211610 MK211572

Limon, CR

Coto Brus,

C. gabbi UCR: 21863 Puntarenas, 1200 8.7889 -82.9583 KT950271 MK211567

Coto Brus,

C. gabbi UCR: 21864 Puntarenas, 1200 8.7889 -82.9583 KT950272 MK211568

Matagalpa,

C. lauraster SME: 79759 Matagalpa, 1300 12.9993 -85.9092 MK211608 MK211565

Puerto

C. lauraster Scone mie ie 190 14.9275 -84.5339 _KU323364 = MK211566

559393 Gracias a

Dios, HN

4 eo Paraiso,

C. persimilis UCR: 22211 1050 9.7841 -83.7517 KT950293 MK211570

Cartago, CR

C. persimilis Cer oem: ~Atamance. 121 9.5773 -82.9343. | MK211609 =_MK211569

Limon, CR

Co pocsclans a eg Se 2250 10.0192 -83.7132. | _MK211633. = MK211589

sensu stricto 0012 Cartago, CR

Amphib. Reptile Conserv. 194 May 2019 | Volume 13 | Number 1 | e176

Craugastor podiciferus species group in Central America

Appendix I (continued). Institutional voucher numbers, locality information, and GenBank accession numbers for the specimens

used in the molecular phylogenetic analyses. Museum collection acronyms follow Frost (2019) with the addition of EAP to refer to

Erick Arias field numbers and CRARC to refer to the Costa Rica Amphibian Research Center private collection. CR = Costa Rica,

HN = Honduras, NI = Nicaragua, PA = Panama.

Species

C. podiciferus

sensu stricto

C. podiciferus

sensu stricto

C. podiciferus

sensu stricto

C. podiciferus

sensu stricto

C. podiciferus

sensu stricto

C. podiciferus

sensu stricto

C. podiciferus

sensu stricto

C. podiciferus

sensu Stricto

C. podiciferus

sensu stricto

C. podiciferus

“Chumacera”

C. podiciferus

“Chumacera”

C. podiciferus

“Fila Costefia”

C. podiciferus

“Fila Costefia”

C. podiciferus

“Fila Costefia”

C. podiciferus

“Fila Costefia”

C. podiciferus

“Monte Verde”

C. podiciferus

“Monte Verde”

C. podiciferus

“Monte Verde”

Institutional

vouchers

SMF: 104005

UCR: 19853

UCR: 19856

UCR: 19860

UCR: 19862

UCR: 20992

UCR: 22146

UCR: 22201

UCR: 23175

UCR: 22120

UCR: 22690

EAP: 0509

FMNH:

257651

UCR: 16585

UCR: 22091

FMNH:

257669

FMNH:

257673

UCR: 16361

Amphib. Reptile Conserv.

Collection

locality

Changuinola,

Bocas del

Toro, PA

Telire,

Talamanca,

Telire,

Talamanca,

Telire,

Talamanca,

Telire,

Talamanca,

Alfaro Ruiz,

Alajuela, CR

Vazquez de

Coronado,

San José, CR

Dota, San

José, CR

Talamanca,

Limon, CR

Buenos Aires,

Puntarenas,

Pérez

Zeledon, San

José, CR

Golfito,

Puntarenas,

Coto Brus,

Puntarenas,

Dota, San

José, CR

Pérez

Zeledon, San

José, CR

Monte Verde,

Puntarenas,

Monte Verde,

Puntarenas,

Alfaro Ruiz,

Alajuela, CR

Elevation

(m)

1766

1817

2108

2143

1700

2395

1860

1821

1793

1546

1350

1400

1488

1500

1930

Geographic coordinates

Lat

8.9908

9.3580

9.3645

10:22772

10.0263

9.7126

9.3659

9.3218

9.3267

8.7878

8.7833

Fats hs:

9.4410

10.2773

10.2176

195

Lon

-82.6716

-83.2294

-83.2164

-84.3482

-83.9448

-83.9488

-83.0417

-83.4546

-83.4706

-83.0306

-82.9833

-83.8580

-83.6830

-84.5891

-84.3671

GenBank Number

16S COI

MK211641 MK211597

MK211639 MK211595

MK211637 MK211593

MK211636 MK211592

MK211638 MK211594

MK211632 MK211588

MK211635 MK211591

MK211634 MK211590

MK211640 MK211596

MK211642 —

MK211631 MK211587

—— MK211605

EF562367 a

MK211647 ——

MK211646 MK211604

EF562372 MK211598

EF562343 MK211603

EF562371 —-

May 2019 | Volume 13 | Number 1 | e176

Arias et al.

Appendix I (continued). Institutional voucher numbers, locality information, and GenBank accession numbers for the specimens

used in the molecular phylogenetic analyses. Museum collection acronyms follow Frost (2019) with the addition of EAP to refer to

Erick Arias field numbers and CRARC to refer to the Costa Rica Amphibian Research Center private collection. CR = Costa Rica,

HN = Honduras, NI = Nicaragua, PA = Panama.

, Institutional Collection Elevation | Geographic coordinates GenBank Number

Species :

vouchers locality (m) Lat Lon 16S COI

OC podiciferis Puntarenas,

net P i UCR: 22675 Puntarenas, 1726 10.3202 -84.7987 — MK211606

Monte Verde

OTe. dich oGm ke es Un22d8 9.8646 -84.1429 MK211644 — MK211601

Pico Blanco José, CR

CRORE cripeognig? WES eat 545 9.8646 -84.1429 _MK211643. ~ MK211600

Pico Blanco José, CR

er Tilaran,

C ROn eres: CRARC i Guanacaste. 1470 10.3600 -84.8000 MK211645 —

San Gerardo 0247

or } Monte Verde,

C ROn ers: Snes Puntarenas, 1500 10.2773 -84.5891 EF562374. = MK211599

San Gerardo 257671 CR

ren eter UcRri6sssa .pampae, 1500 10.2022 84.1625 EF562349. = MK211602

“San Gerardo” Heredia, CR er

POT” suCRTaIEe. se eee sep 9.3987 -83.0200 _MK211630 — MK211586

Siola Limon, CR

Talamanca,

C. polyptychus UCR: 20050 Limén, CR 900 9.6178 -83.2681 MK211614 MK211576

Talamanca,

C. polyptychus UCR: 22668 DEMORER. 198 9.6064 -82.9115 MK211613 MK211575

Nole Duima,

C. saguisp. nov. SMF: 104014 Negdbe Bugle, 1762 8.5571 -81.8245 MK211623 —-

Nole Duima,

C. sagui sp.nov. SMF: 104015 Ngdbe Bugle, 1700 8.4997 -81.7724 MK211624 MK211580

Nole Duima,

C. sagui sp.nov. SMF: 104017 Ngdbe Bugle, 1815 8.4955 -81.7672 MK279370 —-—

Osa,

C. stejnegerianus EAP: 0514 Puntarenas, 45 8.9655 -83.4411 MK211607 MK211563

Buenos Aires,

C. stejnegerianus UCR: 20352 Puntarenas, 900 9.0863 -83.1105 KT950284 MK211564

; Paraiso,

C. underwoodi UCR: 22619 1412 9.7518 -83.7792 MK211611 MK211573

Cartago, CR

Vazquez de

C. underwoodi UCR: 22625 Coronado, 1708 10.0254 -83.9456 MK211612 MK211574

San José, CR

be Eben Buenos Aires,

ante BOT oes UCR: 20389 Puntarenas, 1500 9.1112 -83.1006 MK211625 MK211581

be Ebanl Buenos Aires,

a! SP. UCR: 20428 Puntarenas, 1800 9.1381 -83.0700 MK279371 —

nov.

Amphib. Reptile Conserv. 196 May 2019 | Volume 13 | Number 1 | e176

Craugastor podiciferus species group in Central America

Appendix I (continued). Institutional voucher numbers, locality information, and GenBank accession numbers for the specimens

used in the molecular phylogenetic analyses. Museum collection acronyms follow Frost (2019) with the addition of EAP to refer to

Erick Arias field numbers and CRARC to refer to the Costa Rica Amphibian Research Center private collection. CR = Costa Rica,

HN = Honduras, NI = Nicaragua, PA = Panama.

nee Institutional Collection Elevation | Geographic coordinates GenBank Number

Pp vouchers locality (m) Lat Lon 16S COI

C. zunigai s Coto Brus,

; EGE SD UCR: 22703 Puntarenas, 1732 8.9759 -82.8344 MK279372 saa:

nov. cn

C. zunigai s Coto Brus,

eae Bai UCR: 22709 Puntarenas, 1980 8.9751 -82.8243 MK211626 MK211582

Appendix IT. Specimens used in the morphometric analysis. Museum collection acronyms follow Frost (2019) with the addition of

AH to refer to Andreas Hertz field numbers and HAU to refer to Frank Hauenschild field numbers.

Craugastor blairi: PANAMA: Chiriqut: Bajo Mono, Los Naranjos, Boquete (AH: 0289-0290; SMF: 104028): Cerro La Estrella,

Jaramillo, Boquete (SMF: 104037); Cerro Guayaba, Caldera, Boquete (HAU: 023; SMF: 104035); Volcan Bart, Los Naranjos,

Boquete (AH: 0240-1; SMF: 104026—7); Fortuna, Hornito, Gualaca (AH: 0079-0080, 0372, 00376; HAU: 008, 010; SMF: 102024,

104023, 104025, 104029-33). Ngdbe Buglé: Guayabito, Nuriim (SMF: 104034).

Craugastor sagui sp. nov.: PANAMA: Negdbe Bugle: Cerro Sagui, Jadeberi, Nole Duima (SMF: 104014, 104018—9); La Nevera,

Nole Duima (AH: 0168; SMF: 104015—7).

Craugastor zunigai sp. nov.: COSTA RICA: Puntarenas: Las Alturas, Pittier, Coto Brus (UCR: 22703—4, 2270910); Tres Colinas,

Potrero Grande, Buenos Aires (UCR: 20257—8, 20389, 20395, 20401, 20411, 20419, 20421, 20423, 20428, 23014, 23016—8); road

to Las Tablas, Sabalito, Coto Brus (UCR: 23170).

Amphib. Reptile Conserv. 197 May 2019 | Volume 13 | Number 1 | e176

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

13(1) [General Section]: 198-208 (e177).

urn:lsid:zoobank.org:pub:E9E72353-3C2F -47AD-B6F8-06D72D568E04

A new species of endemic day gecko (Reptilia: Gekkonidae:

Cnemaspis) from a wet zone forest in the second peneplain

of Southern Sri Lanka

‘Anslem de Silva, 7Aaron M. Bauer, *Madhava Botejue, and **Suranjan Karunarathna

‘Amphibia and Reptile Research Organization of Sri Lanka, 15/1, Dolosbage Road, Gampola, SRI LANKA *Department of Biology, Villanova

University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA *Biodiversity Conservation Society, 150/6, Stanly Thilakaratne Mawatha,

Nugegoda 10250, SRI LANKA *Nature Explorations and Education Team, No: B-1 / G-6, De Soysapura Flats, Moratuwa 10400, SRI LANKA

Abstract.—A new day gecko species of the genus Cnemaspis Strauch, 1887 is described from a geographically

separated forested area in Ensalwatte, Matara district, Sri Lanka. This species is medium (34-—35mm SVL) in

size and can be differentiated from all other Sri Lankan congeners by a suite of distinct morphometric, meristic,

and color characters. The species is recorded from rock outcrop habitats in wet, cool, and shady forest with

minimal anthropogenic disturbance at low-elevations (below 860 m). Existing data suggest this gecko is a point

endemic. Being a rupicolous microhabitat specialist with a scansorial mode of life, this species is susceptible

to both localized and widespread threats. Therefore, isolated forest patches warrant special conservation

action, including habitat protection, in-depth research, and species-specific hands-on management practices.

Keywords. Conservation, critically endangered, microhabitat, natural history, point endemic, redlist, Sri Lankan war-

rior, threats

Citation: de Silva A, Bauer AM, Botejue M, Karunarathna S. 2019. A new species of endemic day gecko (Reptilia: Gekkonidae: Cnemaspis) from a

wet zone forest in the second peneplain of Southern Sri Lanka. Amphibian & Reptile Conservation 13(1) [General Section]: 198-208 (e177).

4.0 International (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in any

medium, provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced, are

as follows: official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org.

Received: 16 April 2019; Accepted: 10 May 2019; Published: 3 June 2019

Introduction

During the past decade, the number of day gecko species

recognized in genus Cnemaspis has grown rapidly, reach-

ing over 140 species (Uetz et al. 2019), and it is now con-

sidered the second most diverse gecko genus in the Old

World (Grismer et al. 2014). Cnemaspis species are dimin-

utive, slender-bodied geckos. They possess comparatively

large, forward and upwardly-directed eyes with round

pupils, and widely-splayed limbs bearing elongate slen-

der digits that are bent at an angle with entire subdigital

lamellae (Deraniyagala 1953; Manamendra-Arachchi et

al. 2007; Wood et al. 2017). These crepuscular geckos are

mostly rupicolous, though a few are arboreal or ground-

dwelling, and most Cnemaspis have a cryptic morphology

and coloration, both of which help with camouflage (Gris-

mer et al. 2014; Wood et al. 2017). The genus Cnemaspis

has been the subject of numerous taxonomic revisions and

new species descriptions in recent years. Currently, there

are 25 Cnemaspis species inhabiting Sri Lanka, all of

which are endemic to the island (Manamendra-Arachchi

Correspondence. *suranjan.karu@gmail.com

Amphib. Reptile Conserv.

et al. 2007; Wickramasinghe and Munindradasa 2007;

Vidanapathirana et al. 2014; Wickramasinghe et al. 2016;

Batuwita and Udugampala 2017; Batuwita et al. 2019; Ka-

runarathna et al. 2019a). Herein, another species of Cne-

maspis is reported from a lowland rainforest of southern

Sri Lanka that could not be assigned to a known species

and it is described as new.

Materials and Methods

Specimens: Museum acronyms follow Sabaj Pérez

(2015). The type material discussed in this paper is depos-

ited in the National Museum of Sri Lanka (NMSL), Co-

lombo. Specimens were caught by hand and were photo-

graphed in life. They were euthanized using halothane and

fixed in 10% formaldehyde for two days, washed in water

and transferred to 70% ethanol for long-term storage. Be-

fore fixation, tail tips were collected as tissue samples for

future genetic analyses and stored in 95% ethanol under

cool conditions (25 °C). For comparison, 394 Cnemaspis

specimens (catalogued and uncatalogued) representing all

June 2019 | Volume 13 | Number 1 | e177

de Silva et al.

recognized Sri Lankan species were eaxmined, including

all type specimens housed at the National Museum, Sri

Lanka (NMSL.NH), The Natural History Museum, Lon-

don (BMNH), and Anslem de Silva’s field series (ADS).

Specimens that formerly belonged to the Wildlife Heri-

tage Trust (WHT) collection and bear WHT numbers are

currently deposited in the NMSL, catalogued under their

original numbers. Specimens in this study were collected

during a survey of the lizards of Sri Lanka under permit

numbers WL/3/2/1/14/12 and WL/3/2/42/18 issued by the

Department of Wildlife Conservation, and permit num-

bers FRC/S5 and FRC/6 issued by the Forest Department

of Sri Lanka. Additional information on the morphology

and natural history of Sri Lankan Cnemaspis species was

extracted from the relevant literature (Bauer et al. 2007;

Manamendra-Arachchi et al. 2007; Wickramasinghe and

Munindradasa 2007; Vidanapathirana et al. 2014; Amar-

asinghe and Campbell 2016; Wickramasinghe et al. 2016;

Batuwita and Udugampala 2017; Agarwal et al. 2017;

Batuwita et al. 2019; Karunarathna et al. 2019a). Assign-

ment of unidentified specimens to species was based on

the presence of shared morphometric and meristic char-

acters (Wickramasinghe et al. 2016; Batuwita and Udu-

gampala 2017; Agarwal et al. 2017; Batuwita et al. 2019;

Karunarathna et al. 2019a).

Morphometric characters: Forty morphometric mea-

surements were taken using a Mitutoyo digital Vernier cal-

liper (to the nearest 0.1 mm), and detailed observations

of scales and other structures were made through a Leica

Wild M3Z dissecting microscope. The following symmet-

rical morphometric characters were taken on the left side

of the body: eye diameter (ED), horizontal diameter of

eye ball; orbital diameter (OD), greatest diameter of orbit;

eye to nostril length (EN), distance between anteriormost

point of orbit and posterior border of nostril; snout length

(ES), distance between anteriormost point of orbit and tip

of snout; snout to nostril length (SN), distance between tip

of snout and anterior most point of nostril; nostril width

(NW), maximum horizontal width of nostrils; eye to ear

distance (EE), distance between posterior border of eye and

anteriormost point of ear opening; snout to axilla distance

(SA), distance between axilla and tip of snout; ear length

(EL), maximum length of ear opening; interorbital width

(IO), shortest distance between left and right supraciliary

scale rows; inter-ear distance (IE), distance across the head

between the two ear openings; head length (HL), distance

between posterior edge of mandible and tip of snout; head

width (HW), maximum width of head in-between the ears

and the orbits; head depth (HD), maximum height of head

at the level of the eye; jaw length (JL), distance between

tip of snout and corner of mouth; internarial distance (IN),

smallest distance between the inner margins of nostrils;

snout to ear distance (SED), distance between tip of snout

and anteriormost point of ear; upper-arm length (UAL),

distance between the axilla and the angle of the elbow;

lower-arm length (LAL), distance from elbow to wrist

Amphib. Reptile Conserv.

with palm flexed; palm length (PAL), distance between

wrist (carpus) and tip of longest finger excluding claw;

length of digits I-V of manus (DLM), distance between

juncture of basal phalanx with the adjacent digit and the tip

of the digit, excluding the claw; snout-vent length (SVL)

distance between tip of snout and anterior margin of vent;

trunk length (TRL), distance between axilla and groin;

trunk width (TW), maximum width of body; trunk depth

(TD), maximum depth of body; femur length (FEL), dis-

tance between groin and knee; tibia length (TBL), distance

from knee to ankle with heel flexed; heel length (HEL),

distance between ankle (tarsus) and tip of longest toe (ex-

cluding the claw) with both foot and tibia flexed: length of

pedal digits I-V (DLP), distance between juncture of basal

phalanx with the adjacent digit and the digit tip, excluding

the claw; tail length (TAL), distance between anterior mar-

gin of vent and tail tip; tail base depth (TBD), maximum

height of tail base; tail base width (TBW), widest point of

tail base.

Meristic characters: Twenty-nine characters were ob-

served on scales and other structures using a Leica Wild

M3Z dissecting microscope on both left and right sides of

the body (reported as L/R): number of supralabials (SUP)

and infralabials (INF) between first labial scale and corner

of mouth; number of interorbital scales (INOS), between

left and right supraciliary scale rows; number of postmen-

tals (PM) bounded by chin scales, 1* infralabial on the

left and right, and mental; number of chin scales (CHS)

touching medial edge of infralabials and mental between

juncture of 1% and 2™ infralabials on left and right; num-

ber of supranasal (SUN) scales between nares; presence

of postnasal (PON) scales posterior to naris; presence of

internasal (INT) scale between supranasals; number of

supraciliary scales (SUS) above eye; number of scales

between eye and tympanum (BET) from posterior-most

point of orbit to anterior-most point of tympanum; number

of canthal scales (CAS), number of scales from posterior-

most point of naris to anterior most point of orbit; total

lamellae on manus I-V (SLM) counted from first proximal

enlarged scansor greater than twice the width of the largest

palm scale, to most distal lamella at tip of digits; num-

ber of dorsal paravertebral granules (PG) between pelvic

and pectoral limb insertion points along a straight line im-

mediately left of vertebral column; number of midbody

scales (MBS) from center of mid-dorsal row diagonally

towards the ventral scales; number of midventral scales

(MVS) from first scale posterior to the mental to last scale

anterior to the vent; number of belly scales (BLS) across

venter between lowest rows of granular dorsal scales; total

lamellae on pes I-V (SLP), counted from first proximal

enlarged scansor greater than twice the width of largest

heel scale, to distalmost lamella at tip of digits; number

of femoral pores (FP) present on femur; number of non-

pored postfemoral scales (PFS) counted from distal end of

femoral pore row to knee; tnterfemoral scales (IFS), num-

ber of non-pored scales between innermost femoral pores

June 2019 | Volume 13 | Number 1 | e177

A new species of Cnemaspis from Sri Lanka

300 - 600m

800 - 1000m,

1200 - 1500m

Arid zone forest

Dry zone forest

Y Intermediate zone forest

Wet zone forest

Montane forest

Fig. 1. Currently known distribution of Cnemaspis godagedarai

sp. nov. (Ensalwatte, star), and related species: C. gemunu (Haggala,

circle), C. phillipsi (Gammaduwa, triangle), and C. scalpensis (Gannoruwa, square) in Sri Lanka. Photos: Suranjan Karunarathna.

on both femurs. In addition, the texture (smooth or keeled)

of ventral scales, the texture (homogeneous or heteroge-

neous) of dorsal scales, the number of spinous scales on

the flanks (FLSP), and characteristics such as appearance

of the caudal scales (except in specimens with regener-

ated tails) were also evaluated. Coloration was determined

from digital images of living specimens and from direct

observations in the field.

Habitat and ecology: New species described herein were

collected on field surveys conducted in various habitats of

Sri Lanka (Fig. 1). During this survey, behavior and other

aspects of natural history of the focal species were ob-

served through opportunistic and non-systematic means.

Such observations were done at a minimum distance of

1—2 m from the focal animals while taking precautions to

avoid disturbance. The ambient and substrate tempera-

tures were measured using a standard thermometer and a

N19 Q1370 infrared thermometer (Dick Smith Electron-

ics, Shanghai, China), respectively. The relative humidity

and light intensity were measured with a QM 1594 mul-

tifunction environment meter (Digitek Instruments Co.,

Ltd., Hong Kong, China). Recording elevation and geore-

ferencing species locations, used an eTrex® 10 GPS (Gar-

min, Johannesburg, South Africa). Sex was determined by

the presence (male) or absence (female) of precloacal and

femoral pores. The conservation status of the species was

Amphib. Reptile Conserv.

evaluated using the 2001 IUCN Red List Categories and

Criteria version 3.1 (IUCN 2012).

Taxonomy

Cnemaspis godagedarai sp. nov.

urn:Isid:zoobank.org:act:36C54FD1-01F8-4BC0-9283-991C75804899

Godagedaras’ Day Gecko (English)

Godagedarage Diva-seri Hoona (Sinhala)

Godagedaravin Pahalpalli (Tamil)

Figs. 2-3; Table 1.

Holotype: NMSL 2019.09.01, adult male, 35.5 mm

SVL (Fig. 2), collected from agranite wall in Ensalwatte,

Dentyaya, Matara District, Southern Province, Sri Lanka

(6.388767°N, 80.599781°E, WGS1984; elevation 858 m)

on 6 October 2018 by Suranjan Karunarathna and Anslem

de Silva.

Paratypes: ADS 232, adult female, 34.2 mm SVL, and

ADS 233, adult female, 34.7 mm SVL, collected from

agranite wall in Ensalwatte, Dentyaya, Matara District,

Southern Province, Sri Lanka (6.389128°N, 80.605767°E,

WGS1984; elevation 831 m) on 6 October 2018 by Suran-

jan Karunarathna and Anslem de Silva.

June 2019 | Volume 13 | Number 1 | e177

de Silva et al.

Fig. 2. Close-ups of Cnemaspis godagedarai sp. nov. male holotype (NMSL 2019.09.01.NH). (A) Dorsal head; (B) Lateral head;

(C) Ventral head; (D) Homogeneous dorsal scales; (E) Scales on lateral surface of trunk; (F) Smooth ventral scales; (G) Cloacal

characters with preclocal pores and femoral pores; (H) Subdigital lamellae on manus; (I) Subdigital lamellae on pes; (J) Dorsal

scalation of tail; (IX) Lateral side of tail; (L) Oval shaped subcaudals. Photos: Suranjan Karunarathna.

Diagnosis: Cnemaspis godagedarai sp. nov., can be read-

ily distinguished from its Sri Lankan congeners by the fol-

lowing combination of morphological and meristic char-

acteristics and color pattern: maximum SVL 35.5 mm;

dorsum with homogeneous, smooth, granular scales; 2/2

supranasals; one internasal and 1/1 postnasal present; 26—

28 interorbital scales present; 14-15 supraciliaries, 11-12

canthal scales, 24—26 eye to tympanum scales; three en-

larged postmentals; postmentals bounded by 5-6 chin

Amphib. Reptile Conserv.

scales; chin and gular scales smooth, juxtaposed granules;

pectoral and abdominal scales smooth, subimbricate; 21—

23 belly scales across venter; 5-6 weakly developed tu-

bercles on posterior flank; 101—106 paravertebral granules

linearly arranged; precloacal pores absant in males, 12—13

femoral pores in males and 8-9 unpored interfemoral

scales in males; 133-137 ventral scales; 98-102 midbody

scales; subcaudals smooth, large subhexagonal, subequal

in width, in a regular series forming a median row; 7-8

June 2019 | Volume 13 | Number 1 | e177

A new species of Cnemaspis from Sri Lanka

Fig. 3. Cnemaspis godagedarai sp. nov. male holotype (NMSL 2019.09.01.NH) in life in situ. (A) Dorsolateral view of the full body

displaying color pattern. (B) Lateral view with temporal coloration. Photos: Chen Lee.

supralabials; 7-8 infralabials; 17-18 subdigital lamellae

on digit IV of manus, and 20-21 subdigital lamellae on

digit IV of pes.

Comparisons with other species: The new species is a

member of the C. podihuna clade sensu Agarwal et al.

(2017) based on the presence of large subhexagonal sub-

caudals scales. However, it differs from the all the other

members of the clade as follows: from C. kandambyi

Batuwita and Udugampala, 2017, C. molligodai Wick-

ramasinghe and Munindradasa, 2007 and C. podihuna

Deraniyagala, 1944 by the absence (versus presence) of

precloacal pores; from C. alwisi Wickramasinghe and

Munindradasa, 2007, C. nilgala Karunarathna et al., 2019,

C. punctata Manamendra-Arachchi et al., 2007 and C.

rajakarunai Wickramasinghe et al., 2016 by presence

of more midbody scales (98-102 versus 71—78, 71-78,

71-78 and 69-74), respectively and by fewer femoro-

precloacal scales (eight versus 18-19, 14-15, 25-27 and

Amphib. Reptile Conserv.

20-22), respectively; from C. gemunu Bauer et al., 2007

and C. phillipsi Manamendra-Arachchi et al., 2007 by

presence of more midbody scales (98-102 versus 74—87

and 76-91), respectively and more paravertebral granules

(101-106 versus 79-93 and 86-93), respectively; from C.

rammalensis Vidanapathirana et al., 2014 by fewer ventral

scales (133-137 versus 186-207) and by fewer femoro-

precloacal scales (eight versus 19-24); from C. scalpen-

sis (Ferguson, 1877) by more belly scales (21—23 versus

17-19), by more midbody scales (98-102 versus 81-89)

and by fewer flank spines (5—6 versus 9-11).

Among species of the C. kandiana clade sensu Agarwal

et al. (2017) C. godagedarai sp. nov. differs in the pre-

sense (versus absence) of clearly enlarged, hexagonal or

subhexagonal subcaudal scales, and absence of precloa-

cal pores (versus presence) from the following species: C.

amith Manamendra-Arachchi et al., 2007, C. ingerorum

Batuwita et al., 2019, C. kallima Manamendra-Arachchi et

al., 2007, C. kandiana (Kelaart, 1852), C. kumarasinghei

June 2019 | Volume 13 | Number 1 | e177

de Silva et al.

Table 1. Morphometric and meristic data of male holotype and two female paratypes of Cnemaspis godagedarai sp. nov. from Ensalwatte,

Matara District, Sri Lanka. Abbreviations: L - left, R - right, M - male, F - female.

ists ADS 232 ADS 233 er ADS 232 ADS 233

Measurement 2019.09.01 Counts 2019.09.01

Holotype (M) Paratype(F) Paratype (F) Holotype (M) Paratype(F) Paratype (F)

SVL 3515 34.2 34.7 FLSP (L/R) 6/5 5/5 6/5

ED 2.0 2.0 1.9 SUP (L/R) 8/8 8/8 8/7

OD 3.4 3.4 oe INF (L/R) 7/8 8/8 7/7

EN Die) 2.8 2.8 INOS 28 26 28

ES Ba Sf. 3.6 PM 3 s 3

SN 1.6 1.6 1.4 CHS 6 6 5

NW 0.3 0.3 0.2 SUN (L/R) 2/2 2/2 2/2

EE oe?) Dy 2 PON (L/R) 1/1 1/1 1/1

SA 15.8 1Ss7. |e Rae: INT l l 1

EL 0.8 0.8 0.6 SUS (L/R) 15/14 15/15 14/14

IO 3.8 3.8 3.6 BET (L/R) 26/24 25/26 26/26

IE 3.9. 3.8 She CAS (L/R) 11/11 12/11 11/11

HL 9.7 9.6 9.6 TLM (i) (L/R) 10/10 11/10 10/10

HW Sx 5.3 5.1 TLM (ii) (L/R) 14/15 14/14 14/14

HD 2.8 2.6 2.3 TLM (iit) (L/R) 16/16 15/16 16/16

JL 6.4 6.3 6.1 TLM (iv) (L/R) 18/18 17/18 18/18

IN 1.8 1.8 1.6 TLM (v) (L/R) 13/12 12/12 12/12

SED 8.8 8.7 8.6 PG 105 106 101

UAL 5.9 5.8 5.7 MBS 102 98 99

LAL 57 5.6 5.4 MVS 134 137 133

PAL 3.6 Ps 3.5 BLS 21 21 23

DLM (i) 1.6 LS 1.4 TLP (i) (L/R) 11/10 10/10 11/10

DLM (ii) 1.8 1.7 1.7 TLP (it) (L/R) 14/14 13/14 13/14

DLM (iii) 2.4 ease pa | TLP (iii) (L/R) 16/16 15/15 16/15

DLM (iv) S71 3.1 29 TLP (iv) (L/R) 20/21 21/21 21/21

DLM (v) 25 2.4 2.2 TLP (v) (L/R) 16/16 15/16 16/16

TRL 12.8 12.6 12:2 FP (L/R) 12/13 - -

TW 5.4 5.6 5.4 PFS 4/3 - -

TD 3.4 3.4 3.3 IFS 8 - -

FEL 6.0 5.9 5.8

TBL 5.8 5.7 5.6

HEL a3 5.4 3

DLP (1) 1.6 1.5 1.4

DLP (ii) a 3.1 |

DLP (ili) 3.7 3.6 3.4

DLP (iv) 4.4 43 4.1

DLP (v) 3.9 34 3.5

TAL 38.9 SES 37.6

TBW 3.3 31 a8)

TBD 29 ome 287

Amphib. Reptile Conserv. 203 June 2019 | Volume 13 | Number 1 | e177

A new species of Cnemaspis from Sri Lanka

Wickramasinghe and Munindradasa, 2007, C. latha Man-

amendra-Arachchi et al., 2007, C. menikay Manamendra-

Arachchi et al., 2007, C. pava Manamendra-Arachchi et

al., 2007, C. pulchra Manamendra-Arachchi et al., 2007,

C. retigalensis Wickramasinghe and Munindradasa, 2007,

C. samanalensis Wickramasinghe and Munindradasa,

2007, C. silvula Manamendra-Arachchi et al., 2007, C.

tropidogaster (Boulenger, 1885), and C. upendrai Mana-

mendra-Arachchi et al., 2007.

Description of holotype: An adult male, 35.5 mm SVL.

Body slender, relatively short (TRL 36.0% of SVL). Head

relatively small (HL 27.3% of SVL, HL 75.8% of TRL),

narrow (HW 15.4% of SVL, HW 56.4% of HL), depressed

(HD 8.0% of SVL, HD 29.3% of HL) and distinct from

neck. Snout relatively long (ES 72.1% of HW, ES 40.6%

of HL), slightly less than twice eye diameter (ED 51.9%

of ES), more than half length of jaw (ES 61.0% of JL),

snout slightly concave in lateral view; eye relatively small

(ED 21.1% of HL), larger than ear (EL 39.7% of ED),

pupil rounded; orbit length more than eye to ear distance

(OD 114.3% of EE) and greater than length of IV digit of

manus (OD 109.8% of DLM IV); supraocular ridges not

prominent; ear opening very small (EL 8.4% of HL), deep,

taller than wide, larger than nostrils, smaller than eyes; one

row of scales separate orbit from supralabials; interorbital

distance is broad (IO 96.4% of ES), smaller than snout to

nostril length (IO 39.2% of HL); eye to nostril distance

subequal to eye to ear distance (EN 99.0% of EE).

Dorsal surface of trunk with smooth, small homoge-

neous granules; 105 paravertebral granules; 134 smooth

midventral scales; 102 midbody scales; 6/5 weakly devel-

oped tubercles on flanks; ventrolateral scales not enlarged;

granules on snout smooth and fairly raised, larger than

those on interorbital and occipital regions; canthus rostra-

lis nearly absent, 11/11 smoothly round scales from eye

to nostril; scales of interorbital region oval and smooth;

tubercles absent on sides of neck and around ear; ear

opening vertically oval, slanting from anterodorsal to

posteroventral, 26/24 scales between anterior margin of

ear opening and posterior margin of the eye. Supralabi-

als 8/8, infralabials 7/8, becoming smaller towards gape.

Rostral scale wider than long, partially divided (75%) by

median groove, in contact with first supralabial. Nostrils

separated by 2/2 enlarged supranasals with one internasal;

no enlarged scales behind supranasals. Nostrils oval, dor-

solaterally orientated, not in contact with first supralabials;

1/1 postnasal, smooth, equal to nostrils in size, partially in

contact with first supralabial.

Mental sub-triangular, as wide as long, posteriorly in

contact with four enlarged postmentals (smaller than men-

tal, and lager than posterior postmentals); postmentals in

contact and bordered posteriorly with six smooth, poste-

rior postmental scales (smaller than nostrils), in contact

with the 1* infralabials; ventral scales slightly smaller than

posterior postmentals. Smooth, rounded, juxtaposed scales

present on chin and gular region; pectoral and abdominal

Amphib. Reptile Conserv.

scales smooth, subimbricate towards precloacal region,

abdominal scales larger than dorsals; 21 belly scales

across venter; scales around vent and base of tail smooth,

subimbricate; precloacal pores absent in males, 12/13

femoral pores in males; original tail of holotype slightly

longer than snout-vent length (TAL 109.6% of SVL); tail

base greatly swollen (TBW 3.2 mm), homogeneous round

scales on dorsal aspect of the tail, no spine-like tubercles

at base of tail; tail with 4—5 enlarged, round and flattened

obtuse scales forming whorls; no post-cloacal spur on tail

base; subcaudals enlarged, hexagonal, smooth, arranged

median series.

Forelimbs moderately short, slender (LAL 16.0% of

SVL, UAL 16.8% of SVL); hind limbs long, tibia quite

a bit shorter than femur (TBL 16.3% of SVL, FEL 17.0%

of SVL). Dorsal, anterior, ventral and posterior surfaces of

upper arm and lower arm with smooth scales. Scales on

dorsal surface of femur smooth, granular, less imbricate

scales on anterior, posterior and ventral surfaces, scales on

the anterior surface twice the size of those on other surfac-

es. Dorsal, anterior, posterior and ventral surfaces of tibia

with smooth scales, both anterior and posterior surfaces

of tibia with smooth granules, scales of ventral surface

twice as large as those of other parts. Dorsum and venter

of manus and pes smooth, granular; dorsal surfaces of dig-

its also with granular scales. Digits elongate and slender

with inflected joints, all bearing slightly recurved claws.

Subdigital lamellae entire (except divided at first interpha-

langial joint), unnotched; subdigital lamellae on manus

(left/right): digit I, 10/10, digit II, 14/15, digit II, 16/16,

digit IV, 18/18, digit V, 13/12; subdigital lamellae on pes

(left/right): digit I, 11/10, digit II, 14/14, digit III, 16/16,

digit IV, 20/21, digit V, 16/16; interdigital webbing absent;

length of digits of manus (left): I (1.6 mm), II (1.8 mm),

II (2.4 mm), V (2.5 mm), IV (3.1 mm); length of digits

of pes (left): I (1.6 mm), II (3.2 mm), III (3.7 mm), V (3.9

mm), IV (4.4 mm).

Variation of the type series: SVL of adult specimens in

type series (n = 3) ranges from 34.2—35.5 mm, TAL ranges

from 37.5—38.9 mm, and TRL ranges from 12.2—12.8 mm;

number of supralabials 7-8, and infralabials 7—8; spines

on flank 5-6; interorbital scales 26-28; supraciliaries

14-15; canthal scales 11-12; scales from eye to tympa-

num 24—26; total lamellae on digits of manus (L/R): digit

I (10-11), digit I (14-15), digit II (15-16), digit IV (17—

18), digit V (12-13); total lamellae on digits of the pes

(L/R): digit I (10-11), digit I (13-14), digit III (15—16),

digit IV (20-21), digit V (15-16); ventral scales 133-137;

midbody scales 98-102; paravertebral granules 101—106;

belly scales 21-23; femoral pores 12-13, unpored inter-

femoral scales 8—9, and unpored postfemoral scales 3-4

respectively in males.

Color of living specimen: Dorsum of head, body and

limbs generally grey brown; three large irregular cinna-

mon brown blotches along vertebral line; a distinct narrow,

June 2019 | Volume 13 | Number 1 | e177

de Silva et al.

Fig. 4. General habitat of Cnemaspis godagedarai sp. nov. at Ensalwatte forest, Matara District, Sri Lanka. (A) Complete view of

the forest hill. (B) Dense forest with cool and shady habitat. Photos: Suranjan Karunarathna.

short longitudinal black line on occipital area. Tail cinnamon

brown dorsally, with 12 faded black cross-bands (Fig. 3); pu-

pil is circular and black with surrounding yellow and light

brown margins, supraciliaries being brownish; two postor-

bital stripes are present on each side, the upper white and the

lower black; a light and dark interorbital stripe present; su-

pralabials and infralabaials brown and with dirty white spots:

chin and gular scales light yellow, without dark spots; pec-

toral, abdominal, cloacal and subcaudal scales immaculate

cream; long transverse black line on posterior thigh; dorsum

of limbs with faded brown patches; manus and pes with black

and cream white crossed stripe arrangement.

Color of preserved specimen: Dorsally body 1s pale grey,

colored with dark distinct irregular brown blotches; su-

pralabials and infralabaials are brown, black and white in

color; chin and gular scales becoming dirty white; ventral

surface uniformly dirty white tn color with some scales on

thigh, tail base and arms with dark brown margins.

Amphib. Reptile Conserv.

Etymology: The specific epithet is an eponym Latinized

(godagedarai) in the masculine genitive singular, honor-

ing Sri Lankan warrior Godagedara Rate Adhikaram for

his valiant feats in the Great Rebellion of 1817-1818,

which was initiated in Uva-Wellassa.

Habitat and ecology: The Ensalwatte forest area is a

Mesua-Doona-dominated tropical evergreen rainforest

~150 ha in size (Gunatileke and Gunatileke 1990), locat-

ed in the lowland wet zone of southern Sri Lanka (near

Deniyaya, Mathara District, Southern Province). The

area lies between 6.381864—6.397750°N and 80.589167—

80.617853°E, with an elevation range of 620-860 m (Fig.

4). The mean annual rainfall is 2,500—3,500 mm, received

mostly during the southwest monsoon (May-September)

season. The mean annual temperature is 22—25 °C. Only

SIX specimens (four females and two males) were found

on scattered boulders. The new species was sympatric (at

both local habitat and microsite scales) with several other

June 2019 | Volume 13 | Number 1 | e177

A new species of Cnemaspis from Sri Lanka

micro-endemic geckos (Cnemaspis pulchra and Cyrtodac-

tylus subsolanus). No eggs were found in the same habi-

tat. These microhabitats were well-shaded (light intensity

0-587 Lux), relatively moist (relative humidity 73-88%)

and cool (rock-surface temperature 25.3—27.5 °C).

Conservation status: Application of the IUCN Red List

criteria indicates that C. godagedarai sp. nov. 1s Critically

Endangered, due to having an area of occupancy (AOO)

<10 km? (three locations, 0.8 km? in total assuming a 500

m radius around the georeferenced location) and an extent

of occurrence (EOO) <100 km? (3.9 km?) in the second

peneplain of Southern Province. The applicable criterion

is B2-b (iil).

Remarks: Of its insular congeners, Cnemaspis godage-

darai sp. nov. most closely resembles C. gemunu, C. phil-

lipsi, and C. scalpensis. The type localities of these species

are separated by ~60 km (Hakgala in Central highland),

~95 km (Kandy in Central highland), and ~100 km (Gam-

maduwa in Knuckles highland) airline distances, respec-

tively, from Ensalwatte in Dentyaya (Fig. 1). See also

Comparison with other species above.

Discussion

The addition of another endemic gecko species to the Sri

Lankan reptile list further highlights the island’s status as a

key center of reptilian diversity (see Batuwita et al. 2019;

Karunarathna et al. 2019). Cnemaspis now comprises

26 species in Sri Lanka, bringing the total number of

geckos to 48. Among Sri Lankan gekkonids, 38 (~80%)

species are endemic to the island, most of which are

restricted to the wet zone (>2,000 mm of annual average

precipitation). However, as this study demonstrates, Sri

Lanka’s Cnemaspis diversity is not limited to either the

southwestern lowlands or the central massif, but is spread

throughout the bio-climatic regions and floristic regions

of Sri Lanka, which suggests intricate biogeographic

patterns possibly due to multiple colonizations from the

Indian mainland rather than a singular colonization event

and an insular radiation (Agarwal et al. 2017). These

findings suggest the possible speciation of Cnemaspis

in geographically isolated mountains with granite caves,

rock outcrops, and favorable environmental conditions,

and the number of species is predicted to increase to

more than 50 (e.g., Agarwal et al. 2017; Batuwita et al.

2019; Karunarathna et al. 2019b). Thus, continuation of

faunal surveys and detailed examination of morphological

diagnostic features, as well as genetic analyses, are

critical for revealing the true Cnemaspis diversity in Sri

Lanka. We strongly recommend that such studies focus

on isolated hills, smaller forests, rock outcrops, and

granite caves, including underground tunnel systems.

Habitats of Cnemaspis species, including the type

localities, are undergoing extensive habitat conversion,

and are threatened by localized human disturbances such

Amphib. Reptile Conserv.

as encroachments for tea plantations and other crops,

and human settlements. Habitat conservation of isolated

mountains 1s absolutely crucial for the conservation of the

unique fauna of the island.

Acknowledgements.—We thank the Director Gen-

erals, Laxman Peiris, the research committee, and the

field staff of the Department of Wildlife Conservation

(WL/3/2/1/14/12, and WL/3/2/42/18) and the Forest

Department (FRC/5, and FRC/6) for granting permis-

sion and providing help during the field surveys. Nanda

Wickramasinghe, Sanuja Kasthuriarachchi, Lankani So-

maratne, Manori Nandasena, Chandrika Munasinghe,

Rasika Dasanayake, Ravindra Wickramanayake, and P.

Gunasiri at NMSL assisted while we were examining

collections under their care. Dinesh Gabadage, Mayintha

Madawala, Gayan Edirisinghe, Niranjan Karunarathna,

and Chen Lee (for Fig. 3), are acknowledging for their

valuable assistance. This work was mainly supported by

the Nagao Natural Environment Foundation (2018-20),

grants to SK, and United States National Science Foun-

dation grants DEB 1555968 and EF 1241885 (subaward

13-0632) to AMB. Finally, we would like to thank Kan-

ishka Ukuwela, Thilina Surasinghe, Thasun Amaras-

inghe, and Zeeshan Mirza for support and constructive

criticism of an earlier draft, and anonymous reviewers

for useful comments.

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maratna L, Madawala M, Gabadage D, Botejue M,

Henkanaththegedara S, Ukuwela K. 2019a. Descrip-

tion of a new species of the genus Cnemaspis Strauch,

1887 (Reptilia: Squamata: Gekkonidae) from the

Nilgala Savannah forest, Uva Province of Sri Lanka.

Zootaxa 4545: 389-407.

Karunarathna S, Poyarkov NA, de Silva A, Madawala

M, Botejue M, Gorin VA, Surasinghe T, Gabadage D,

Ukuwela K, Bauer AM. 2019b. Integrative taxonomy

reveals six new species of day geckos of the genus

Cnemaspis Strauch, 1887 (Reptilia: Squamata: Gek-

konidae) from geographically isolated hill forests in

Sri Lanka. Vertebrate Zoology, In Press.

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2007. A taxonomic revision of the Sri Lankan day

geckos (Reptilia: Gekkonidae: Cnemaspis), with de-

scription of new species from Sri Lanka and Southern

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Vidanapathirana DR, Rajeev MDG, Wickramasinghe

N, Fernando SS, Wickramasinghe LJM. 2014. Cne-

maspis rammalensis sp. nov., Sri Lanka’s largest day-

gecko (Sauria: Gekkonidae: Cnemaspis) from Ram-

malakanda Man and Biosphere Reserve in southern

Sri Lanka. Zootaxa 3755: 273-286.

Wickramasinghe LJM, Munindradasa DAI. 2007. Re-

view of the genus Cnemaspis Strauch, 1887 (Sauria:

Gekkonidae) in Sri Lanka with the description of five

new species. Zootaxa 1490: 1-63.

Wickramasinghe LJM, Vidanapathirana DR, Rathnayake

RMGBP. 2016. Cnemaspis rajakarunai sp. nov., a rock

dwelling day-gecko (Sauria: Gekkonidae: Cnemas-

pis) from Salgala, an unprotected lowland rainforest

in Sri Lanka. Zootaxa 4168: 92-108.

Wood PL Jr, Grismer LL, Aowphol A, Aguilar CA, Cota

M, Grismer MS, Murdoch ML, Sites JW Jr. 2017.

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and the phylogenetic placement of C. punctatonuch-

alis and C. vandeventeri. Peer J 5: 01-46 (e2884).

Anslem de Silva, M.Sc., D.Sc. (University of Peradeniya), started keeping reptiles at the early age of seven. Anselm has

taught herpetology at the Rajarata University of Sri Lanka and final year veterinary students at University of Peradeniya.

Anselm has conducted herpetofaunal surveys in most of the important ecosystems in Sri Lanka and has published more

than 400 papers, of which nearly 60 are books or chapters in books. Anslem had done yeoman service to the country and

the region for more than 50 years, and he is the Regional Chairman of the Crocodile Specialist Group for South Asia and

Tran, Co-Chair of the Amphibian Specialists Group IUCN/SSC Sri Lanka.

Aaron Bauer grew up collecting reptiles and amphibians in his native New York. Aaron has been studying reptiles,

especially geckos, for more than 35 years, and is the Gerald M. Lemole Endowed Professor of Integrative Biology at

Villanova University. From his extensive work in Sri Lanka, India, southern Africa, Australia, and the South Pacific, Aaron

has described nearly 200 species of reptiles and written more than 750 publications. Aaron is a former Secretary General

of the World Congress of Herpetology, President of the Society for the Study of Amphibians and Reptiles, President of the

Herpetologists’ League, and Chairman of the Herpetological Association of Africa.

Madhava Botejue is a wildlife biologist, and has been conducting research on the biodiversity, ecology, conservation,

behavior and taxonomy of Sri Lankan fauna for the past fourteen years. Madhava has contributed to environmental

conservation through many community-based awareness programs. Madhava earned his B.Sc. degree in Natural

Sciences from The Open University of Sri Lanka in 2009, and currently serves as an Environmental Officer at the Central

Environmental Authority, Sri Lanka and also as the Secretary of the Biodiversity Conservation Society, Sri Lanka.

Madhava is a member of IUCN/SSC, Crocodile Specialist Group and an expert committee member of the National Red

Suranjan Karunarathna obtained a Masters in Environmental Management from the University of Colombo, Sri Lanka.

Suranjan’s scientific exploration of biodiversity began with the Young Zoologists’ Association of Sri Lanka (YZA) in

early 2000, and he was president of YZA in 2007. As a wildlife researcher, Suranjan studies herpetofaunal ecology and

taxonomy, promotes the scientific basis of conservation awareness and the importance of biodiversity and its conservation

among the Sri Lankan community. Suranjan is an active member of many specialist groups in the IUCN/SSC, and an

expert committee member of Herpetofauna in the National Red List development programs of Sri Lanka since 2004.

June 2019 | Volume 13 | Number 1 | e177

A new species of Cnemaspis from Sri Lanka

Appendix 1

Comparative material:

Cnemaspis alwisi: NMSL 2004.9.1 (holotype), NMSL 2004.9.2 (paratype), NMSL 2004.9.3 (paratype), WHT 5918, WHT 6518,

WHT 6519, WHT 7336, WHT 7337, WHT 7338, WHT 7343, WHT 7344, WHT 7345, WHT 7346.

C. amith: BMNH 63.3.19.1066A (holotype), BMNH 63.3.19.1066B (paratype), BMNH 63.3.19.1066C (paratype).

C. gemunu: AMB 7495 (holotype), AMB 7507 (paratype), WHT 7221, WHT 7347, WHT 7348, NMSL 2006.11.01, NMSL

2006.11.02, NMSL 2006.11.03, NMSL 2006.11.04.

C. ingerorum: WHT 7332 (holotype), WHT 7330 (paratype) WHT 7331 (paratype).

C. kallima: WHT 7245 (holotype), WHT 7222 (paratype), WHT 7227 (paratype), WHT 7228 (paratype), WHT 7229 (paratype),

WHT 7230 (paratype), WHT 7239 (paratype), WHT 7249 (paratype), WHT 7251 (paratype), WHT 7252 (paratype), WHT 7253

(paratype), WHT 7254 (paratype), WHT 7255 (paratype).

C. kandambyi: WHT 9466 (holotype), WHT 9467 (paratype).

C. kandiana: BMNH 53.4.1.1 (lectotype), BMNH 80.2.2.119A (paralectotype), BMNH 80.2.2.119B (paralectotype), BMNH

80.2.2.119C (paralectotype), WHT 7212, WHT 7213, WHT 7267, WHT 7305, WHT 7307, WHT 7308, WHT 7310, WHT 7313,

WHT 7319, WHT 7322.

C. kumarasinghei: NMSL 20061301 (holotype), NMSL 20061302 (paratype).

C. latha: WHT 7214 (holotype).

C. menikay: WHT 7219 (holotype), WHT 7218 (paratype), WHT 7349 (paratype).

C. molligodai: NMSL 2006.14.01 (holotype), NMSL 2006.14.02-5 (paratype), NMSL 2006.14.03 (paratype), NMSL 2006.14.04

(paratype), NMSL 2006.14.05 (paratype).

C. nilgala: 2018.07.01.NH (holotype), 2018.06.01.NH (paratype), 2018.06.02.NH (paratype), 2018.06.03.NH (paratype).

C. pava: WHT 7286 (holotype), WHT 7281 (paratype), WHT 7282 (paratype), WHT 7283 (paratype), WHT 7285 (paratype),

WHT 7288 (paratype), WHT 7289 (paratype), WHT 7290 (paratype), WHT 7291 (paratype), WHT 7292 (paratype), WHT 7293

(paratype), WHT 7294 (paratype), WHT 7295 (paratype), WHT 7296 (paratype), WHT 7297 (paratype), WHT 7298 (paratype),

WHT 7299 (paratype), WHT 7300 (paratype), WHT 7301 (paratype), WHT 7302 (paratype).

C. phillipsi: WHT 7248 (holotype), WHT 7236 (paratype); WHT 7237 (paratype); WHT 7238 (paratype).

C. podihuna: BMNH 1946.8.1.20 (holotype), NMSL 20061002, NMSL 20061003, NMSL 20061004.

C. pulchra: WHT 7023 (holotype), WHT 1573a (paratype), WHT 7011 (paratype), WHT 7021 (paratype), WHT 7022 (paratype).

C. punctata: WHT 7256 (holotype), WHT 7223 (paratype), WHT 7226 (paratype), WHT 7243 (paratype), WHT 7244 (paratype).

C. rajakarunai: NMSL 2016.07.01 (holotype), DWC 2016.05.01 (paratype), DWC 2016.05.02 (paratype).

C. rammalensis: NMSL 2013.25.01 (holotype), DWC 2013.05.001.

C. retigalensis: NMSL 20061201 (holotype), NMSL 20061202 (paratype), NMSL 20061203 (paratype), NMSL 20061204 (paratype).

C. samanalensis: NMSL 2006.15.01 (holotype), NMSL 2006.15.02 (paratype), NMSL 2006.15.03 (paratype), NMSL 2006.15.04

(paratype), NMSL 2006.15.05 (paratype).

C. scalpensis: NMSL 2004.1.1 (neotype), NMSL 2004.2.1, NMSL 2004.3.1, NMSL 2004.4.1, WHT 7265, WHT 7268, WHT 7269,

WHT 7274, WHT 7275, WHT 7276, WHT 7320.

C. silvula: WHT 7208 (holotype), WHT 7206 (paratype), WHT 7207 (paratype), WHT 7209 (paratype), WHT 7210 (paratype), WHT

7216 (paratype), WHT 7217 (paratype), WHT 7018, WHT 7027, WHT 7202, WHT 7203, WHT 7220, WHT 7354, WHT 7333.

C. tropidogater: BMNH 71.12.14.49 (lectotype), NMSL 5152, NMSL 5151, NMSL 5159, NMSL 5157, NMSL 5970, NMSL 5974.

C. upendrai: WHT 7189 (holotype), WHT 7184 (paratype), WHT 7187 (paratype), WHT 7188 (paratype), WHT 7181 (paratype), WHT

7182 (paratype), WHT 7183 (paratype), WHT 7185 (paratype), WHT 7190 (paratype), WHT 7191 (paratype), WHT 7192 (paratype),

WHT 7193 (paratype), WHT 7194 (paratype), WHT 7195 (paratype), WHT 7196 (paratype), WHT 7197 (paratype), WHT 72.

Amphib. Reptile Conserv. 208 June 2019 | Volume 13 | Number 1 | e177

Amphibian & Reptile Conservation

13(1) [General Section]: 209-229 (e178).

Official journal website:

amphibian-reptile-conservation.org

Reptiles of Ecuador: a resource-rich online portal, with

dynamic checklists and photographic guides

‘Omar Torres-Carvajal, ?Gustavo Pazmiho-Otamendi, and *David Salazar-Valenzuela

'?Museo de Zoologia, Escuela de Ciencias Biolégicas, Pontificia Universidad Catolica del Ecuador, Avenida 12 de Octubre y Roca, Apartado 17-

01-2184, Quito, ECUADOR ?Centro de Investigacion de la Biodiversidad y Cambio Climdatico (BioCamb) e Ingenieria en Biodiversidad y Recursos

Genéticos, Facultad de Ciencias de Medio Ambiente, Universidad Tecnologica Indoamérica, Machala y Sabanilla EC170301, Quito, ECUADOR

Abstract.—With 477 species of non-avian reptiles within an area of 283,561 km?, Ecuador has the highest density

of reptile species richness among megadiverse countries in the world. This richness is represented by 35

species of turtles, five crocodilians, and 437 squamates including three amphisbaenians, 197 lizards, and 237

snakes. Of these, 45 species are endemic to the Galapagos Islands and 111 are mainland endemics. The high

rate of species descriptions during recent decades, along with frequent taxonomic changes, has prevented

printed checklists and books from maintaining a reasonably updated record of the species of reptiles from

Ecuador. Here we present Reptiles de! Ecuador (http://bioweb.bio/faunaweb/reptiliaweb), a free, resource-rich

online portal with updated information on Ecuadorian reptiles. This interactive portal includes encyclopedic

information on all species, multimedia presentations, distribution maps, habitat suitability models, and

dynamic PDF guides. We also include an updated checklist with information on distribution, endemism, and

conservation status, as well as a photographic guide to the reptiles from Ecuador.

Keywords. Biodiversity, Bioweb, distribution maps, PDF guide, Reptilia, South America

Abstract.—Con 477 especies de reptiles (excluyendo aves) en un area de 283,561 km?, Ecuador tiene la

mayor densidad de riqueza de especies de reptiles de los paises megadiversos del mundo. Esta riqueza esta

representada por 35 especies de tortugas, cinco crocodilios y 437 escamosos, incluyendo tres anfisbeéenidos,

197 lagartijas y 237 serpientes. De estas, 45 especies son endémicas de las Islas Galapagos y 111 son endémicas

del Ecuador continental. La alta tasa de descripciones de especies durante las ultimas deéecadas, junto con los

frecuentes cambios taxonomicos, han impedido que las listas de especies y los libros impresos mantengan un

registro razonablemente actualizado de las especies de reptiles del Ecuador. Aqui presentamos Reptiles del

Ecuador (http://bioweb.bio/faunaweb/reptiliaweb), un portal en linea gratuito y rico en recursos con informacion

actualizada sobre los reptiles ecuatorianos. Este portal interactivo incluye informacion enciclopédica sobre

todas las especies, presentaciones multimedia, mapas de distribucion, modelos de aptitud de habitat y guias

dinamicas en PDF. Incluimos ademas una lista de especies actualizada, con informacion sobre distribucion,

endemismo y estado de conservacion, asi como una guia fotografica de los reptiles del Ecuador.

Palabras clave. América del Sur, biodiversidad, Bioweb, guia PDF, mapas de distribucion, Reptilia

Citation: Torres-Carvajal O, Pazmifo-Otamendi G, Salazar-Valenzuela D. 2019. Reptiles of Ecuador: a resource-rich portal, with a dynamic checklist

and photographic guides. Amphibian & Reptile Conservation 13(1): [General Section]: 209-229 (e178).

tribution 4.0 International (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in

any medium, provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced,

are as follows: official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org.

Received: 31 January 2019; Accepted: 15 April 2019; Published: 18 May 2019

Introduction

477 non-avian reptiles (Torres-Carvajal et al. 2019) have

With an area of 283,561 km? including that of the Gala-

pagos Islands, Ecuador is one of both the smallest and

the most biodiverse countries in South America and the

world (Joppa et al. 2011; Myers et al. 2000; Ulloa Ulloa

et al. 2017). Among tetrapod vertebrates, 609 species of

amphibians (Ron et al. 2019), 1,690 birds (Freile and

Poveda 2019), 432 mammals (Brito et al. 2019), and

been recorded in this country to date. For all these taxa,

Ecuador also has the highest density of species richness

(i.e., number of species per area unit) among megadi-

verse countries in the world, as well as a remarkable pro-

portion of endemic species, even if Galapagos taxa are

excluded (Brito et al. 2019; Freile and Poveda 2019; Ron

et al. 2019; Torres-Carvajal et al. 2019).

Correspondence. 'omartorcar@gmail.com (Corresponding author), *gustavopo85(@gmail.com, *davidsalazarv@gmail.com

Amphib. Reptile Conserv.

May 2019 | Volume 13 | Number 1 | e178

Torres-Carvajal et al.

a 50

— 45

ox 37

ep)

O 30 27 a

—_ 25

Q 20 19 i 19

= 14 iS 14

= ES

nd 11 410 11

Z 8 8

6 5

3 3

% oS OS DS © —S pS GS & S S © % Bw po BS B SG pAB® —& DGD HY

ae oes oe Me Se ee SS FS ee ol of of ol of ol ol? of :

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Decade

Fig. 1. Histogram showing the description of reptile species present in Ecuador through time, from Linnaeus’ 10" edition of Systema

Naturae to the end of 2018. The number of species described per decade is presented above each bar.

Ecuador’s remarkable non-avian reptile richness com-

bined with major knowledge gaps have challenged the

production of a single comprehensive review of the rep-

tiles from this country. Starting with the classic works

of James A. Peters “Zhe Snakes of Ecuador: a check

list and key” (Peters 1960) and “The Lizards of Ecua-

dor: a check list and key” (Peters 1967), several check-

lists (Almendariz 1992; Miyata 1982; Torres-Carvajal

2001, 2011) and, more recently, books and field guides,

on one or more Ecuadorian regions or taxa have been

published (e.g., Arteaga et al. 2013; MECN 2010; Valen-

cia et al. 2008, 2016; see also the Field Museum’s field

guides at http://fieldguides.fieldmuseum.org). A com-

mon problem with these publications, however, is that

they quickly become outdated in both numbers of spe-

cies and taxonomy (Torres-Carvajal 2011). Only in this

century (2000-2018), 67 species of reptiles occurring in

Ecuador have been described; moreover, the last decade

(2009-2018) is the second most productive of all times

in number of species described, being outnumbered only

by 1859-1868 (Fig. 1). This development is positive be-

cause it indicates that the interest in the systematic study

of the reptiles of Ecuador has increased in recent years, in

agreement with the Systematics-Agenda-2020’s mission

one: “To discover and document past and present life on

earth” (Daly et al. 2012).

As a response to the rapidly changing nature of the

knowledge on Ecuadorian reptiles, in 2000 the Museum

of Zoology at Pontificia Universidad Catolica del Ec-

uador (QCAZ) published an on-line checklist (URL no

longer exists) including scientific names, authors, year of

description, and general altitudinal distribution data. In

2009, however, this online checklist was improved dra-

Amphib. Reptile Conserv.

210

matically into an interactive website, with the aim of in-

cluding more detailed information (e.g., species diagno-

ses, natural history, distribution, conservation status) and

multimedia for all species of the reptiles of Ecuador. This

platform, Reptiles del Ecuador (http://bioweb.bio/fau-

naweb/reptiliaweb), is part of a larger portal named BIO-

WEB (http://bioweb.bio) that also includes information

on other taxonomic groups. The main goal of this portal,

currently available only in Spanish, is to make biologi-

cal information on the biodiversity of Ecuador available

to researchers, students (at any level), educators, policy

makers, and the general public. In this sense, Reptiles del

Ecuador also supports Systematics-Agenda-2020’s mis-

sion four: “To communicate and apply this knowledge

to science and society” (Daly et al. 2012). BIOWEB is

publishing, for the first time, databases from Ecuadorian

biocollections with nearly half a million specimens avail-

able online (as of March 2019).

After a decade of development, here we describe the

most important features and resources currently available

at Reptiles del Ecuador, and present an updated checklist

of the reptiles from Ecuador (Table 1), with information

on distribution across major biogeographic regions (Fig.

2), and a sample photographic guide (Supplementary file

1).

Methods

Taxonomy

Following the hierarchical nature of phylogenetic trees,

species names in Reptiles del Ecuador are taxonomically

organized by major clades, which have traditionally been

May 2019 | Volume 13 | Number 1 | e178

Reptiles of Ecuador portal

-84.0 -83.0 -82.0 -81.0 -80.0

0.0

-2.0

Biogeographic Regions

|__| Dry Coastal Shrub

-3.0 = Deciduous Coastal Forest

= Chocoan Tropical Rainforest

| Western Foothill Forest

=] Western Montane Forest

= Andean Shrub

HM Paramo

[Eastern Montane Forest

=) Eastern Foothill Forest

= Amazonian Tropical Rainforest

-5.0 i Galapagos

-4.0

-84.0 -83.0 -82.0 -81.0 -80.0

-79.0 -78.0 -77.0 -76.0 -75.0

COLOMBIA A 7

-2.0

-3.0

-4.0

-5.0

-79.0 -78.0 -77.0 -76.0 -75.0

Fig. 2. Biogeographic regions of Ecuador. Source: https://bioweb.bio (modified from Sierra et al. 1999).

ranked as orders, families, and/or subfamilies. As an excep-

tion, the popular division of Squamates: amphisbaenians

(Amphisbaena), lizards (Sauria, paraphyletic), and snakes

(Serpentes) has been adopted. In general, the taxonomy of

Reptiles del Ecuador is based on information available in

scientific publications, but the decision to adopt or reject

a particular taxonomic arrangement is the responsibility of

the editors. Following articles 8.1 and 11.1 of the Inter-

national Code of Zoological Nomenclature (1999), addi-

tional scientific names and nomenclatural acts available in

unpublished theses and dissertations are excluded from the

list; as is the use of the subspecies category, following the

logic adopted at the Mesoamerican Herpetology website

(http://mesoamericanherpetology.com; accessed 2 Febru-

ary 2019; see also de Queiroz 2005).

Species accounts and images

For each species, Reptiles del Ecuador includes informa-

tion on authorship, type specimens, type locality, syn-

onyms, etymology, identification (1.e., morphological

characterization), coloration, natural history, distribution,

conservation status, systematics, and bibliography. Be-

cause this information is mostly taken from the literature,

the accounts of poorly known species are incomplete. In

addition, each account is linked to an up-to-date list of

specimens housed at the Museum of Zoology, QCAZ, the

largest reptile collection in Ecuador (>17,000 specimens).

Amphib. Reptile Conserv.

Species accounts are also associated with image gal-

leries, which include relevant figures from the literature,

maps, and photographs (if available). As of March 2019,

Reptiles del Ecuador includes over 60,000 photographs of

the majority of species. For rare or extinct species (e.g.,

Holcosus orcesi or Chelonoidis niger), scientific illustra-

tions are provided. In most cases, photographs have been

taken in a studio with a white background; however, the

photographic collection of Reptiles del Ecuador has ben-

efited greatly from many donors. While only a subset of

the best shots is professionally edited and included in a

separate album for each species, free access is provided

to most available photographs under the CC BY-NC-ND

4.0 license.

Distribution maps and habitat suitability models, pres-

ent and future

Based on locality data from the QCAZ reptile collection,

as well as literature records, Reptiles del Ecuador includes

interactive Google maps for the distribution of each spe-

cies, in which the user can obtain general information

(voucher number and locality) for each record. These maps

are automatically updated as new specimens are entered in

the collection’s database. Data from GBIF, VertNet, and

iNaturalist can also be displayed on each map.

Moreover, based on locality data from the QCAZ col-

lection, every two weeks Reptiles del Ecuador automati-

May 2019 | Volume 13 | Number 1 | e178

Torres-Carvajal et al.

Table 1. List of reptile species of Ecuador as of March 2019. Scientific name, authorship, biogeographic region, and information on

endemism (EN; x = endemic to Ecuador) and conservation status (CS! = assessment of Carrillo et al. 2005; CS” = assessment from

IUCN Red List 2018) is presented for each species. Number of species within each taxon is presented in parentheses. Biogeographic

regions (see Fig. 2): 1=Dry Coastal Shrub, 2=Deciduous Coastal Forest, 3=Chocoan Tropical Rainforest, 4=Western Foothill Forest,

5=Western Montane Forest, 6=Andean Shrub, 7=Paramo, 8=Eastern Montane Forest, 9=Eastern Foothill Forest, 10=Amazonian

Tropical Rainforest, and 11=Galapagos. The website Reptiles del Ecuador (https://bioweb.bio/faunaweb/reptiliaweb) should be

checked in perpetuity for updates.

ae reer Rallies tees

es 1 }2t3}4}sfol7}s}ofiofin

Alligatoridae (4)

[Caiman crocodins _[uimaows758 —- | ‘| Ix] | | ||] ]*| | | |

[Melanosuchusniger | Spnts05———S—S=«d«<S«dY|«d| «| | | | | | | (|e

[Paleosuchus palpebroms [cwiertso7 +t ‘| | | |||) tx] | |oo[ic

[Paleosuchas rigonanus [Sehmeider 1801 | | | || | 11 Ixlx} | [tc [ee

Crocodylidae (1)

[Crocodviusacumus fewer? ————s«dY~dx)] TL TT PTT 3p

SQUAMATA: AMPHISBAENIA (3)

Amphisbaenidae (3)

[Anphisbaenacaiba____[uimaews758 | ||| |] 1] ]*[x] | [1c [ cc.

[Anphisbaena basi _[vanaoini2002 | | || ||| |«f«[x| | [ic | ne

[Anphisboona vara _[tawens.v768 | | leleteE PE TE LP tbe

SQUAMATA: SAURIA (197)

Alopoglossidae (9)

Linnaeus 1758

Duellman 1973

O’Shaughnessy 1881

Boulenger 1885

Peracca 1904

Torres-Carvajal and Lobos 2014

Boulenger 1890

Boulenger 1912

Harris 1994

Ptychoglossus bilineatus'

Ptychoglossus brevifrontalis

Ptychoglossus gorgonae

[Peyhogtossus iineatus"

[Peyhoglossus brevifontls

[Peychoglossusgorgonae

[Duinértandrwibron 1636 x][x [=| | | |] [=]*|=| [Ne [oc

Paorean de Jonnés 1818 —Tx| | | 111.1 | | |_[e [xe

PDusnérit and-wibron 1636 | [x[xtxt [| LL. xl=L [ve [xe

Betancourt et al. 2018

Oftedal 1974

Gunther 1859

Sanchez-Pacheco et al. 2012

Sanchez-Pacheco et al. 2011

Boulenger 1902

Sanchez-Pacheco et al. 201

O’Shaughnessy 1879

Kizirian 1996

Amphib. Reptile Conserv. 212 May 2019 | Volume 13 | Number 1 | e178

Reptiles of Ecuador portal

Table 1 (continued). List of reptile species of Ecuador as of March 2019. Scientific name, authorship, biogeographic region,

and information on endemism (EN; x = endemic to Ecuador) and conservation status (CS' = assessment of Carrillo et al. 2005;

CS? = assessment from IUCN Red List 2018) is presented for each species. Number of species within each taxon is presented in

parentheses. Biogeographic regions (see Fig. 2): 1=Dry Coastal Shrub, 2=Deciduous Coastal Forest, 3=Chocoan Tropical Rainforest,

4=Western Foothill Forest, 5=Western Montane Forest, 6=Andean Shrub, 7=Paramo, 8=Eastern Montane Forest, 9=Eastern Foothill

Forest, 10=Amazonian Tropical Rainforest, and 11=Galapagos. The website Reptiles del Ecuador (https://bioweb.bio/faunaweb/

reptiliaweb) should be checked in perpetuity for updates.

Kizirian 1996

O’Shaughnessy 1881

Boulenger 1885

Kohler et al. 2004

Boulenger 1890

Fritts and Smith 1969

Fritts et al. 2002

O’Shaughnessy 1881

Seid ep onsite ame acu RAS

Burt and Burt 1931

Altamirano-Benavides et al. 201

Torres-Carvajal et al. 201

Montanucci 1973

his:

Pholidobolus vertebralis O’Shaughnessy 1879

lam

fan

lan

Potamites strangulatus Cope 1868

Kizirian 1996

Kizirian and Coloma 1991

[Riama colomaromani |

Riama colomaromani Kizirian 1996

Kizirian 1996

Boulenger 1885

Kizirian 1995

Kizirian 1996

O’Shaughnessy 1879

Kizirian 1996

Aguirre-Pefiafiel et al. 201

ée

ia trisanale

osaura argula

osaura manicata

nosaura keyi

a elegans

Pholidobolus prefrontalis Montanucci 1973

mites ecpleopus

a anatoloros

a balneator

na cashcaensis

va unicolor

Spe) el el SY am ASB Pf | ea ag] PS aE |

ie [ing JA she ep I [Cif SUE SU aS ta] THY Yi

Amphib. Reptile Conserv. 213 May 2019 | Volume 13 | Number 1 | e178

Torres-Carvajal et al.

Table 1 (continued). List of reptile species of Ecuador as of March 2019. Scientific name, authorship, biogeographic region,

and information on endemism (EN; x = endemic to Ecuador) and conservation status (CS' = assessment of Carrillo et al. 2005;

CS? = assessment from IUCN Red List 2018) is presented for each species. Number of species within each taxon is presented in

parentheses. Biogeographic regions (see Fig. 2): 1=Dry Coastal Shrub, 2=Deciduous Coastal Forest, 3=Chocoan Tropical Rainforest,

4=Western Foothill Forest, 5=Western Montane Forest, 6=Andean Shrub, 7=Paramo, 8=Eastern Montane Forest, 9=Eastern Foothill

Forest, 10=Amazonian Tropical Rainforest, and 11=Galapagos. The website Reptiles del Ecuador (https://bioweb.bio/faunaweb/

reptiliaweb) should be checked in perpetuity for updates.

ae AT ar (ee ee

1 }2t3}4}sfo]7}s}ofiojin

Iguanidae: Corytophaninae (1)

[Iguanidae: Corytophaninae()

[Basitscus gaterins __ [Duméritand Dumert 851 | [x]x[x[x. |] LL 11 ele,

[Anolis aequaroriis [weer | | | [x[x] |] |] | | [St[sE

[Anolis anchicayae __[Poeeta.2009 | | [x}xlxl || || | | [ve [ Ne

[Anolis bioratus | Perrsisea___[x}x|x}x]x} || || | | [oo[ ne,

[Anolis biectus [Cope 86a __[x}x|x}x]x} || |] | |= [1c [ne

[Anolis bombiceps ____|oove ss | | 1111111 1«

Boulenger 1898

Yanez-Mufioz et al. 2018

Boulenger 1885

Peracca 1904

Williams and Duellman 1984

Ginther 1859

D’Orbigny 1837

O’Shaughnessy 1875

Boulenger 1898

Duméril and Duméril 1851

Anolis dracula

Anolis festae

Anolis fitchi

Torres-Carvajal et al. 201

Boulenger 1898

Torres-Carvajal et al. 2017

Miyata 1985

Poe et al. 2009

Boulenger 1898

Williams 1965

Lazell 1969

Cope 1868

Ayala-Varela and Velasco 2010

Williams 1975

Williams 1966

Boulenger 1898

Ayala-Varela and Torres-Carvajal 2010

Ayala- Varela et al. 2014

Boulenger 1902

Peters and Orcés 1956

Daudin 1802

Cope 1899

Anolis lososi

Anolis lynchi

Anolis lyra

Anolis orcesi

Anolis poei

Anolis princeps

[Anotsfraseri [Ginter 1659

[Anolis gommosss | 0'Shmghoessy 1875

[Anolis non | Cope 1868

[Anolis peraccae | Boulenger 1898

Seals

pt tT TNE | NE

pt tt txt xt | dtc | Ne |

Amphib. Reptile Conserv. 214 May 2019 | Volume 13 | Number 1 | e178

Reptiles of Ecuador portal

Table 1 (continued). List of reptile species of Ecuador as of March 2019. Scientific name, authorship, biogeographic region,

and information on endemism (EN; x = endemic to Ecuador) and conservation status (CS' = assessment of Carrillo et al. 2005;

CS? = assessment from IUCN Red List 2018) is presented for each species. Number of species within each taxon is presented in

parentheses. Biogeographic regions (see Fig. 2): 1=Dry Coastal Shrub, 2=Deciduous Coastal Forest, 3=Chocoan Tropical Rainforest,

4=Western Foothill Forest, 5=Western Montane Forest, 6=Andean Shrub, 7=Paramo, 8=Eastern Montane Forest, 9=Eastern Foothill

Forest, 10=Amazonian Tropical Rainforest, and 11=Galapagos. The website Reptiles del Ecuador (https://bioweb.bio/faunaweb/

reptiliaweb) should be checked in perpetuity for updates.

io eee sisal ieee how

Z i }2{3/4]sio]7]3}ofioli

[Anotissomni [Pecan Vater wirnea 2008 | | | | [ [xl [x[al | [ [Ne | ne

[Anolis rackyderma [cores | | 11111 |«felx] | [ic | ne

[Anolis ransversals ___[Dumértand umes 1951 | | || [1-11 [x«f | [ac | ne

[Anolis vancolin [Wiliams eta 1996 | ||| [|| [«[ | [|= [en | ne

Anolis venrimecidans [Boutenger912_| || |x[xP 11 [1 || [xe [ar

Anolis wittamsmivermeeroran [Pocmavateziewcaror7 | | || | | |x| | | | [Ne | Ne.

Riguanidue: Hoploeercimaec) | t{{{1l{[,_11 1 1 _

Enyalioides altotambo Torres-Carvajal et al. 2015

Enyalioides anisolepis Torres-Carvajal et al. 2015

Duellman 1973

Bocourt 1874

Guichenot 1855

O’Shaughnessy 1881

Boulenger 1881

O’Shaughnessy 1881

ve!

basi

[al

Torres-Carvajal et al. 2009 =

[si

[md

x |

Enyalioides cofanorum

Enyalioides heterolepis

Enyalioides laticeps

Enyalioides microlepis

Enyalioides oshaughnessyi

Enyalioides praestabilis

Enyalioides rubrigularis

Enyalioides touzeti Torres-Carvajal et al. 2008

Morunasaurus annularis O’Shaughnessy 1881

Iguanidae: Iguaninae (5)

Bell 1825

Gentile and Snell 2009

Heller 1903

Gray 1831

Linnaeus 1758

Conolophus marthae

Conolophus pallidus

Conolophus subcristatus

Iguanidae: Polychrotinae (5) ‘fe

Werner 1910

Berthold 1846 al

Boulenger 1908 [ia

Linnaeus 1758 ‘ee

Noble 1924

Polychrus femoralis

Polychrus gutturosus

Polychrus liogaster

Polychrus marmoratus

Amblyrhynchus cristatus

Polychrus peruvianus

Iguanidae: Tropidurinae (32)

Baur 1890

Baur 1892

Peters 1871

[mal

Baur 1890 |

Microlophus albemarlensis

Microlophus barringtonensis

Microlophus bivittatus

Microlophus delanonis

Baur 1890

Bell 1843

Microlophus duncanensis

Microlophus grayii

Microlophus habelii Steindachner 1876

[crotophusjocobi —__[pateo2 | | 11 {> L111 bel [ee [cc

Amphib. Reptile Conserv. 215 May 2019 | Volume 13 | Number 1 | e178

Torres-Carvajal et al.

Table 1 (continued). List of reptile species of Ecuador as of March 2019. Scientific name, authorship, biogeographic region,

and information on endemism (EN; x = endemic to Ecuador) and conservation status (CS' = assessment of Carrillo et al. 2005;

CS? = assessment from IUCN Red List 2018) is presented for each species. Number of species within each taxon is presented in

parentheses. Biogeographic regions (see Fig. 2): 1=Dry Coastal Shrub, 2=Deciduous Coastal Forest, 3=Chocoan Tropical Rainforest,

4=Western Foothill Forest, 5=Western Montane Forest, 6=Andean Shrub, 7=Paramo, 8=Eastern Montane Forest, 9=Eastern Foothill

Forest, 10=Amazonian Tropical Rainforest, and 11=Galapagos. The website Reptiles del Ecuador (https://bioweb.bio/faunaweb/

reptiliaweb) should be checked in perpetuity for updates.

Ean ire | Biogeographic Region | | Gg | og

‘ 1 }2{3/4]sio]7]s}ofiolin

‘nirolophus ccpuas _[rewsien _IxIx| || | [| 1] | | [clec

[Mcrolophus paojious | Steindactner 376 | | || | 11111 [x] [ar {cc

[Mcrolophus perwiams —[tessonisx0__Ix} || |} 11111 | [cle

[Plcaplica——____-[Linaewsi756 | |||. Pt] xp | [ac| ne.

[Pica umbea ______[tinaersr758 | |||. 11 lele[e> | [ac [ne

stenocereusaedeanas [O'Shaughnessy 1979 | | || 1111 [«{ || [aclec

Stenocereus angel —____[Tores-Carval 2000 | | | | [x}xlx|a

Stenocercus angulifer Werner 1901 | x |

Stenocercus cadlei Torres-Carvajal and Mafla-Endara 2013

Fal

ee i

Stenocercus carrioni Parker 1934

Fea

Stenocercus chota Torres-Carvajal 2000

Stenocercus festae Peracca 1897 x

Stenocercus guentheri Boulenger 1885

Led

Stenocercus haenschi Werner 1901

Stenocercus humeralis Gunther 1859 x

Stenocercus iridescens Gunther 1859

Rime

Stenocercus limitaris Cadle 1998

Stenocercus ornatus Gray 1845

Stenocercus puyango Torres-Carvajal 2005

Stenocercus rhodomelas Boulenger 1899

Stenocercus simonsii Boulenger 1899

Stenocercus varius Boulenger 1885

BERS SERRE

Fea fl

a a

DEAD: Fee Re ee oe eee

Uracentron flaviceps Guichenot 1855

Phyllodactylidae (14)

Phyllodactylus barringtonensis | Van Denburgh 1912

Phyllodactylus baurii Garman 1892

Phyllodactylus darwini Taylor 1942

Phyllodactylus duncanensis Van Denburgh 1912

PAE

S| Sa GY ad esa el | | a

Phyllodactylus galapagensis | Peters 1869

Phyllodactylus gilberti Heller 1903

Phyllodactylus gorii Lanza 1973

Phyllodactylus kofordi Dixon and Huey 1970

Phyllodactylus leei Cope 1889

Phyllodactylus leoni Torres-Carvajal et al. 2013

Piplodactus punitis [Dixon and tuey 197 x} | |x| | ||| [|_| x [op | op

Piplodacpus reissi ___[rewsieea______Ix}x|xlx[x} || || [a] [aclec

rhecadatyusrapicanda —[Hioutuyn 792 | [x|x[x{ ||] 1 | | [ac [ne

rhecadactyiussfimoensis [Bergmann ana Rusctt2007 | T | | LL > 1 (etx L [ve [ne

fseineige@

Scincidae (2)

[Mabioa altanazonica __[Mialteseeal.2006 | | 111111... Dele

Amphib. Reptile Conserv. 216 May 2019 | Volume 13 | Number 1 | e178

ied

Reptiles of Ecuador portal

Table 1 (continued). List of reptile species of Ecuador as of March 2019. Scientific name, authorship, biogeographic region,

and information on endemism (EN; x = endemic to Ecuador) and conservation status (CS' = assessment of Carrillo et al. 2005;

CS? = assessment from IUCN Red List 2018) is presented for each species. Number of species within each taxon is presented in

parentheses. Biogeographic regions (see Fig. 2): 1=Dry Coastal Shrub, 2=Deciduous Coastal Forest, 3=Chocoan Tropical Rainforest,

4=Western Foothill Forest, 5=Western Montane Forest, 6=Andean Shrub, 7=Paramo, 8=Eastern Montane Forest, 9=Eastern Foothill

Forest, 10=Amazonian Tropical Rainforest, and 11=Galapagos. The website Reptiles del Ecuador (https://bioweb.bio/faunaweb/

reptiliaweb) should be checked in perpetuity for updates.

roa, oe aNeleeet'| ae

: i }2}3t4}s}o}7]s] 9/10} i

nie Cd dE

Sphaerodactylidae (11)

Giiher 1859 Gee P PEE, ee, Pe. pene

Gonatodes caudiscutatus

O’Shaughnessy 1881

Guichenot 1855

Werner 1910

Avila-Pires 2001

Peracca 1897

Miyata 1985

Boulenger 1914

Parker 1926

Parker 1935

Boulenger 1902

Gonatodes concinnatus

Gonatodes humeralis

Lepidoblepharis buchwaldi

Lepidoblepharis conolepis

Lepidoblepharis festae

Lepidoblepharis grandis

Lepidoblepharis intermedius

Lepidoblepharis ruthveni

Pseudogonatodes guianensis

Sphaerodactylus scapularis

—~

—

Linnaeus 1758

Duméril and Bibron 1839

Dumeéril and Bibron 1839

Daudin 1802

Cope 1868

Peters 1964

Dumeéril and Duméril 185

Cope 1875

Cope 1868

Bocourt 1874

Murphy et al. 2016

SQUAMATA: SERPENTES (237)

Callopistes flavipunctatus

Dicrodon guttulatum

Dracaena guianensis

Holcosus bridgesii

Holcosus orcesi

Holcosus septemlineatus

Kentropyx altamazonica

Kentropyx pelviceps

Medopheos edracanthus

Tupinambis cuzcoensis

Linnaeus 1758

Amphib. Reptile Conserv. 217 May 2019 | Volume 13 | Number 1 | e178

Torres-Carvajal et al.

Table 1 (continued). List of reptile species of Ecuador as of March 2019. Scientific name, authorship, biogeographic region,

and information on endemism (EN; x = endemic to Ecuador) and conservation status (CS' = assessment of Carrillo et al. 2005;

CS? = assessment from IUCN Red List 2018) is presented for each species. Number of species within each taxon is presented in

parentheses. Biogeographic regions (see Fig. 2): 1=Dry Coastal Shrub, 2=Deciduous Coastal Forest, 3=Chocoan Tropical Rainforest,

4=Western Foothill Forest, 5=Western Montane Forest, 6=Andean Shrub, 7=Paramo, 8=Eastern Montane Forest, 9=Eastern Foothill

Forest, 10=Amazonian Tropical Rainforest, and 11=Galapagos. The website Reptiles del Ecuador (https://bioweb.bio/faunaweb/

reptiliaweb) should be checked in perpetuity for updates.

7 te | ___Biogeographie Region | | og, | gs

sd at 1 }2]st4atstoi7}s}o fof

Linnaeus 175 oe) dele pe, [2c ne

Werner 1909 all

Linnaeus 1758

Peters 1869

Schmidt and Walker 1943

Duméril et al. 1854

Giinther 1858

Schlegel 1837

Dumeéril et al. 1854

Giinther 1860

Peters 1863

Linnaeus 1758

Cope 1868

Barbour and Amaral 1924

Andersson 1916

Spilotes megalolepis ___| Giimther 1865

Linnaeus 1758

Berthold 1846

Boulenger 1903

Wilson and Mena 1980 | x | cR | DD_

Cope 1875 CECE [oo fic

Amphib. Reptile Conserv. 218 May 2019 | Volume 13 | Number 1 | e178

a] Te YT] ay [eae ee al eict

Hed] Tales] YA hep ae ae eal | li ee | ea CSPI Te be 2 er a a

ay 5s PP a] | EL ef el] LY i 0a a a 4 P

BERR eee

SERRE bese Seas Rene

Linnaeus 1758 x

FWemer1909 | x

Peters 1869 | x |

Rove 1952 x |

Giinther 1858 x | x |

Dunn i933 |

Cadle2012 | |

/Dumériletal 1854 || x

[Cope 1860 x

Linnaeus 1758 x | x

Despax 1910 | x |

[Cope 1875 x |

Cope 1868 x | x |

Peters 1863 x | x |

Wager 1824 | x | x |

Copetsor | x |

Giinther 1865 x

Berthold 1846 | x | x

[Cope 1875 x x |

Reptiles of Ecuador portal

Table 1 (continued). List of reptile species of Ecuador as of March 2019. Scientific name, authorship, biogeographic region,

and information on endemism (EN; x = endemic to Ecuador) and conservation status (CS' = assessment of Carrillo et al. 2005;

CS? = assessment from IUCN Red List 2018) is presented for each species. Number of species within each taxon is presented in

parentheses. Biogeographic regions (see Fig. 2): 1=Dry Coastal Shrub, 2=Deciduous Coastal Forest, 3=Chocoan Tropical Rainforest,

4=Western Foothill Forest, 5=Western Montane Forest, 6=Andean Shrub, 7=Paramo, 8=Eastern Montane Forest, 9=Eastern Foothill

Forest, 10=Amazonian Tropical Rainforest, and 11=Galapagos. The website Reptiles del Ecuador (https://bioweb.bio/faunaweb/

reptiliaweb) should be checked in perpetuity for updates.

oe eee ree ere] pe

: 1 }2{3}4]siol7]3}ofiolin

Tanita nsitanontna __[Wisonmawenatoao | | || [xf [| [1 | [x [alo

rena melanocephaia _[Linnaensi758 | [x|xlx[x} | [x[x[x] | [ac [ne

rant miata | Wisonandkigt9e7 | | [x| | | 1] |. | [x [op] op

Tantilla petersi Wilson 1979 ee

Peters 1863 Sesrirs

Tantilla supracincta

Colubridae: Dipsadinae (136)

Passos et al. 2018

Parker 1930

Arteaga et al. 2017

Peracca 1897

Boulenger 1880

Savage 1955

Giinther 1858

Arteaga et al. 2017

Savage 1955

Myers and Schargel 2006

Peracca 1896

Boettger 1898

Boulenger 1894

Cope 1868 baal

= ia

ie Ea

[j=

[ad

fel

Boulenger 1894 i x

[ad

a El

i [nl

et tt yt fete] [ex [tc [ic

x | x | [Zan

Atractus gigas ee | RX.

Jan 1865 en

Savage 1955

x | x |

Jan 1862

Savage 1955

Despax 1910 x | x

Arteaga et al. 2017

Peracca 1897

Despax 1910

Salazar-Valenzuela et al. 2014

Atractus savagei

Cunha and Nascimento 1983

Dumeéril et al. 1854

Schargel et al. 2013

Passos et al. 2009

Daudin 1803

Zetl

Clelia clelia

Clelia equatoriana

Pp txtxtx] | [tc | Ne |

Amaral 1924 (Se ct DD ae

Peters 1863 xtxtx} |] | tt tT | ft vo]

xt tt tt tf fp {ic |

pt | tt | | NE] NE |

Amphib. Reptile Conserv. 219 May 2019 | Volume 13 | Number 1 | e178

ial

Caracas atts —_____‘[Passoserah 2018 i

[Atractus carrion) [Parker 1930

[Atractus cerberus | Arteaga etal. 2017 Ec

[Atractus colts | Peneea 1897 id

[Atractu dbo | Boulenger 1880 iz

[Atrctus duit | Savage 1955 i

Atrctu euadorensis ___| Savage 1955 Ei

[Atrctu laps | Ginter 1658 ci

[Atrctus esepe | Arteaga etal. 2017 Ei

[Atrciu gigene | Savage 1955 on

[Atractus gigas | Myers and Schagel 2006 a

[Atrcta iidescons | Pencoas96 in

[Atractus tehmarni | Boitger 1898

[Atrctes major Boulenger 1894 il

Atractes microvhyachas [Cope 868 wD

Atractus modests | Boulenger 1894 a

Aractus mdtcincas [an 1865 is

Aractus occidentalis | Savage 1955 oF

Aractas ociptoatbus [Jan 1862 Ri

[arractu ovcesi | Savage 1955 x

[Atractu paucidens | Despax 910 x

[Atrctus proni_____| Arteaga etal. 2017 ia

[Atroctu resplendens | Pencca 897 a

[Atrctus rouiei____| Despax 910

[Atractu savages | Salaae- Valeria 2014 Ei

[Atracts snethlageae | Cua and Nascimento 1983 a

[Atctatoruates __[Dunérletal 1654 ol

[Atracta toes | Sehargel etal. 2013 x

[Atracts phon | Passos eal. 2009 FI

[Cie cieia | Dawdin 1803 Ei

[Cie quatorina [Amaral 924

[Coniophanes dromicornis [Petes 1863 is

Coniophanes dromiciformis

Torres-Carvajal et al.

Table 1 (continued). List of reptile species of Ecuador as of March 2019. Scientific name, authorship, biogeographic region,

and information on endemism (EN; x = endemic to Ecuador) and conservation status (CS' = assessment of Carrillo et al. 2005;

CS? = assessment from IUCN Red List 2018) is presented for each species. Number of species within each taxon is presented in

parentheses. Biogeographic regions (see Fig. 2): 1=Dry Coastal Shrub, 2=Deciduous Coastal Forest, 3=Chocoan Tropical Rainforest,

4=Western Foothill Forest, 5=Western Montane Forest, 6=Andean Shrub, 7=Paramo, 8=Eastern Montane Forest, 9=Eastern Foothill

Forest, 10=Amazonian Tropical Rainforest, and 11=Galapagos. The website Reptiles del Ecuador (https://bioweb.bio/faunaweb/

reptiliaweb) should be checked in perpetuity for updates.

ie pone | __—Biogeographie Region | | gs | gs

: i }2]}3t4atstel7} sto] rola

jan 1863 CT leet OL, [iar iae

Boulenger 1896 SS

Arteaga et al. 2018 L)

Sentzen 1796 i

Boulenger 1896 (=

Boulenger 1898 i

Arteaga et al. 2018 |

Boulenger 1902 fe

Laurenti 1768 [=

Orcés and Almendariz 198 fal

Arteaga et al. 2018 ‘|

Orcés and Almendariz 198 |

Cope 1868 [i

Arteaga et al. 2018

Boulenger 1912 iw

Schlegel 1837 oa

Werner 1909 [a

Duméril et al. 1854 C

Peters 1960 im

Jan 1863 [=

Wied-Neuwied 1824 fw

Fritts and Smith 1969 iG

Boulenger 1898 fal

[mi

[al

Linnaeus 1758

Cope 1860

Peracca 1897

Linnaeus 1758

Cope 1868

Linnaeus 1758

Erythrolamprus subocularis Boulenger 1902

Erythrolamprus taeniogaster | Jan 1863

Linnaeus 1758

Dixon 2000

Myers and McDowell 201

Schlegel 1837 CTT [oo [ne

Shreve 1934 ee

Amphib. Reptile Conserv. 220 May 2019 | Volume 13 | Number 1 | e178

hd a] 3] = sap ae | AT t f_L e S L

ee MS] A a a aE Tall | ae ad

es UE aT] Sap ae Tan Se aE TP (es TS

Reptiles of Ecuador portal

Table 1 (continued). List of reptile species of Ecuador as of March 2019. Scientific name, authorship, biogeographic region,

and information on endemism (EN; x = endemic to Ecuador) and conservation status (CS' = assessment of Carrillo et al. 2005;

CS? = assessment from IUCN Red List 2018) is presented for each species. Number of species within each taxon is presented in

parentheses. Biogeographic regions (see Fig. 2): 1=Dry Coastal Shrub, 2=Deciduous Coastal Forest, 3=Chocoan Tropical Rainforest,

4=Western Foothill Forest, 5=Western Montane Forest, 6=Andean Shrub, 7=Paramo, 8=Eastern Montane Forest, 9=Eastern Foothill

Forest, 10=Amazonian Tropical Rainforest, and 11=Galapagos. The website Reptiles del Ecuador (https://bioweb.bio/faunaweb/

reptiliaweb) should be checked in perpetuity for updates.

cP Bl[slo[7[s [pope

ielicops pours ____[Rowmanrorg | | || [111 [xx] [x [tian

tvirops marti ______[Weeleriee | 111.111] i«P) [clic

tips triangular __[Waelerieaa | || 11111] Ix] | [1c [ve

Imantodes cenchoa Linnaeus 1758 x

Torres-Carvajal et al. 2012

Boulenger 1896

(in

ial

Cope 1894 [mal

Imantodes chocoensis

Imantodes inornatus

Imantodes lentiferus

Linnaeus 1758

Kennicott 1859

Leptodeira annulata

Leptodeira septentrionalis x x

fa)

x |

Linnaeus 1758

Hallowell 1845

fe)

Parker 1940 (i

[ui

Lygophis lineatus re

Angarita-Sierra and Lynch 2017

Nothopsis rugosus Cope 1871

Oxyrhopus fitzingeri Tschudi 1845 x

Werner 1916

Tschudi 1845

Wagler 1824

Linnaeus 1758

Lynch 2009

Zaher et al. 2014

Daudin 1803

Boulenger 1900

Linnaeus 1758

Cope 1862

Ginther 1860

[pal

[Si

[ial

Zaher et al. 2018 =

[Ss

[—

[ual

fa}

[al

Oxyrhopus leucomelas i

Oxyrhopus melanogenys i

Oxyrhopus occipitalis fad

Oxyrhopus petolarius

Oxyrhopus vanidicus

Philodryas argentea

Philodryas simonsii ie

Philodryas viridissima a

Pseudalsophis biserialis

Pseudalsophis darwini

Steindachner 1876

Tschudi 1845

Zaher et al. 2018

Van Denburgh 1912

Zaher et al. 2018

Schneider 1801

Linnaeus 1758

Pseudalsophis dorsalis

Pseudalsophis hephaestus

Pseudalsophis hoodensis

Pseudalsophis occidentalis

Pseudalsophis slevini

Pseudalsophis steindachneri

Pseudalsophis thomasi

Pseudoboa coronata

Pseudoeryx plicatilis

[Nini ctrata | Hallowell 1845

[Nini hudsoni [Parker 1940

[Nina teresitae | neers yooh 2077

[Plccercuseuryzonus [Cope 1862

Rhadinaea decorata Gunther 1858

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Saphenophis bourse [nies | | 1 telxleE | >> LL lw

Amphib. Reptile Conserv. 221 May 2019 | Volume 13 | Number 1 | e178

Torres-Carvajal et al.

Table 1 (continued). List of reptile species of Ecuador as of March 2019. Scientific name, authorship, biogeographic region,

and information on endemism (EN; x = endemic to Ecuador) and conservation status (CS' = assessment of Carrillo et al. 2005;

CS? = assessment from IUCN Red List 2018) is presented for each species. Number of species within each taxon is presented in

parentheses. Biogeographic regions (see Fig. 2): 1=Dry Coastal Shrub, 2=Deciduous Coastal Forest, 3=Chocoan Tropical Rainforest,

4=Western Foothill Forest, 5=Western Montane Forest, 6=Andean Shrub, 7=Paramo, 8=Eastern Montane Forest, 9=Eastern Foothill

Forest, 10=Amazonian Tropical Rainforest, and 11=Galapagos. The website Reptiles del Ecuador (https://bioweb.bio/faunaweb/

reptiliaweb) should be checked in perpetuity for updates.

oP Plls[o]7[s [pope

Shon armada fommerem | |iet T| LI || pete

Arteaga et al. 2018 [imal

Peters 1957 =

Linnaeus 1758 (-]

Sheehy et al. 2014 lal

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Daudin 1803 [mal

Peracca 1896 |

Torres-Carvajal et al. 2015 ie

Hillis 1990 =

Nicéforo-Maria 1950 ie)

| Pyron et al. 2015 [ial

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Torres-Carvajal et al. 2015

zea Ya

Peters 1863

Linnaeus 1758

Bocourt 1891

Boulenger 1903

Cope 1886

Berthold 1859

Wied-Neuwied 1824

Linnaeus 1758

Wucherer 1861

Taeniophallus brevirostris

Thamnodynastes pallidus

Tretanorhinus mocquardi

Tretanorhinus taeniatus

Urotheca fulviceps

Urotheca lateristriga

Xenodon severus

Reeniophalis breasts

rharmedyacites plus

[netanorinns mocguand

[netanorinnsteniatas

[wrothecafubiceps

[wrote letersiga

[xenodon severis

Linnaeus 1766

Micrurus ancoralis Jan 1872 iC

Jan 1872

Jan 1858

Wagler 1824

Linnaeus 1758

Schmidt 1936

Dumeéril et al. 1854

Rendahl and Vestergren 1940

hes

[mt

Jan 1863 iS

Micrurus dumerilii

Micrurus hemprichii

Micrurus lemniscatus

Micrurus mertensi

Micrurus mipartitus

Micrurus multiscutatus

Jan 1858

Schmidt 1936

Roze 1967

Cope 1870

Wagler 1824

Werner 1901

Micrurus ornatissimus

Micrurus peruvianus

Micrurus scutiventris

Safe ae eA emi ead] |S] 5 Oe |

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Amphib. Reptile Conserv. 222 May 2019 | Volume 13 | Number 1 | e178

Reptiles of Ecuador portal

Table 1 (continued). List of reptile species of Ecuador as of March 2019. Scientific name, authorship, biogeographic region,

and information on endemism (EN; x = endemic to Ecuador) and conservation status (CS' = assessment of Carrillo et al. 2005;

CS? = assessment from IUCN Red List 2018) is presented for each species. Number of species within each taxon is presented in

parentheses. Biogeographic regions (see Fig. 2): 1=Dry Coastal Shrub, 2=Deciduous Coastal Forest, 3=Chocoan Tropical Rainforest,

4=Western Foothill Forest, 5=Western Montane Forest, 6=Andean Shrub, 7=Paramo, 8=Eastern Montane Forest, 9=Eastern Foothill

Forest, 10=Amazonian Tropical Rainforest, and 11=Galapagos. The website Reptiles del Ecuador (https://bioweb.bio/faunaweb/

reptiliaweb) should be checked in perpetuity for updates.

— Rae me ee ee

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[nicraas surnamensis [ower | ||| [1111 1x] | [cle

[aicrarusischudi [mess | Txt | LPeE LLL LP expe

Leptotyphlopidae (6)

Jan 1861

Klauber 1939

Bailey 1946

Orejas-Miranda and Peters 1970

Peters 1857

Salazar- Valenzuela et al. 2015

Epictia subcrotilla

Trilepida anthracina

Trilepida guayaquilensis

Trilepida macrolepis

Peery

Trilepida pastusa

Tropidophiidae (4)

Trachyboa boulengeri Peracca 1910

Peters 1860

Laurent 1949

Steindachner 1880

Trachyboa gularis

Tropidophis battersbyi

Tropidophis taczanowskyi

| Lt pxtxt tx} | | | Ten] Ne

Typhlopidae (1)

[Ameronphlopsreentans [uimaowsi758 | | | 1 1.11.1. | [ele

Berthold 1846

Freire-Lascano 1991

Amaral 1935

Cope 1875

Garman 1884

Linnaeus 1758

Wied-Neuwied 1821

Hoge 1954

Parker 1930

Freire-Lascano 1991

Peters 1862

Garcia 1896

Wagler 1824

Garcia 1896

Linnaeus 1766

Schatti and Kramer 1993

Bocourt 1868

a=)

io)

-—

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Bothriechis schlegelii

Bothrocophias campbelli

Bothrocophias hyoprora

Bothrocophias microphthalmus

Bothrops bilineatus

Bothrops brazili

Bothrops punctatus

xtx{x| || LC | NE |

Bothrops taeniatus

Lachesis acrochorda

Porthidium arcosae

[Bothriecis shiegehi _[Bonold 1846

[Botiocophias campheli | Feeire-Lascano 1991

[Bothrocophias oprora [Amaral 1935 iam

[Borhrocaphias micophhans | Cope 1875 rae

[Bothrops sper [Garman 1884 eile

[Bothrops atrox [Linnaeus 1758 ial

[Bothrops bitineatas | Wied-Newwied 182 Airs

[Bothrops racih [Hoge 1958 Pin

[Bothrops tejamas [Parker 1930 =I

[Botiops asbomei | Feere-Lascano 1991 me

[Bothrops puicher [Petes 1862 nie

[Botts punctatas | Gurcia 1696 ne

[Bothrops tacniatus | Wagler 824 an

[Lachess acrochorda | Gmrcia 1696 na

[tachesis mata [Linnaeus 1765 n=

[Portion cosas | Sehit and Kramer 1953 ol

[Portion nation [Bovour 1868 ale

Porthidium nasutum Tee = Saree

Schneider 1783

Schweigger 1812

McCord et al. 2001

Amphib. Reptile Conserv. 223 May 2019 | Volume 13 | Number 1 | e178

Torres-Carvajal et al.

Table 1 (continued). List of reptile species of Ecuador as of March 2019. Scientific name, authorship, biogeographic region,

and information on endemism (EN; x = endemic to Ecuador) and conservation status (CS' = assessment of Carrillo et al. 2005;

CS? = assessment from IUCN Red List 2018) is presented for each species. Number of species within each taxon is presented in

parentheses. Biogeographic regions (see Fig. 2): 1=Dry Coastal Shrub, 2=Deciduous Coastal Forest, 3=Chocoan Tropical Rainforest,

4=Western Foothill Forest, 5=Western Montane Forest, 6=Andean Shrub, 7=Paramo, 8=Eastern Montane Forest, 9=Eastern Foothill

Forest, 10=Amazonian Tropical Rainforest, and 11=Galapagos. The website Reptiles del Ecuador (https://bioweb.bio/faunaweb/

reptiliaweb) should be checked in perpetuity for updates.

ory rere pallies tees

aa 1 }2{3}4]s[ol]7]8}ofiolin

Cheloniidae (4)

[carenacareta +. sd TT TTT]

Eretmocheiss imbricata [Linnaeus 1766 _|x{x} | | | | | | | [x |op[ cr

[Lepidochels otvacea _[sehsehoww 1829 | Ixt || | ||. 1 tx, [oolw

Chelydridae (1)

[Chebyiraacuirosis [oes | | [ele] 1} 111111. [wpe

Dermochelyidae (1)

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Geoemydidae (3)

ete) thee I) iL ses nr)

ee eee ewes

jt txtet Tt tt | tt EN [ty

Troschel 1848

Chelonoidis abingdonii Giinther 1877

Chelonoidis becki Rothschild 1901

Chelonoidis chathamensis Van Denburgh 1907

Chelonoidis darwini Van Denburgh 1907

Chelonoidis denticulatus Linnaeus 1766

Chelonoidis donfaustoi Poulakakis et al. 2015

Chelonoidis duncanensis Pritchard 1996

Chelonoidis guntheri Baur 1889

Chelonoidis hoodensis Van Denburgh 1907

Ginther 1874

Quoy and Gaimard 1824

Van Denburgh 1907

Rotschild 1903

De Sola 1930

Ginther 1875

'The distribution of Ptychoglossus bilineatus is uncertain because the only known specimen is the holotype, which lacks locality

data other than simply “Ecuador.”

Amphib. Reptile Conserv. 224 May 2019 | Volume 13 | Number 1 | e178

Reptiles of Ecuador portal

Table 2. Degree of endemism of the Ecuadorian reptile fauna at the species level, arranged by clades traditionally recognized as

families.

Clade “Family” Total Number of Species

CROCODYLIA

Alligatoridae 4

Crocodylidae

Subtotals 5

SQUAMATA

Amphisbaenidae 3

Alopoglossidae 9

Anguidae

Gekkonidae 3

Gymnophthalmidae 49

Iguanidae OF

Phyllodactylidae 14

Scincidae 2

Sphaerodactylidae 1]

Teiidae 10

Aniltidae

Anomalepididae

Boidae

Colubridae 180

Elapidae 19

Leptotyphlopidae

Tropidophiidae

Typhlopidae

Viperidae 17

Subtotals 437

TESTUDINES

Chelidae 6

Cheloniidae 4

Chelydridae 1

Dermochelyidae 1

Geoemydidae a

Kinosternidae 2

Podocnemidae 3

Testudinidae 15

Subtotals 35

Totals A477

cally runs habitat suitability models (HSMs) for each spe-

cies with more than four locality records (Austin 2002).

These models are constructed under two approaches, BIO-

CLIM (Busby 1991) and MAXENT (Phillips et al. 2006),

depending on the number of localities, 5—9 and >10, re-

spectively. Additionally, projections on future WorldClim

climatic layers (2030, 2050, and 2070) are periodically

calculated under four carbon dioxide emission scenarios

(RCP 2.6, RCP 4.5, RCP 6.0, and RCP 8.5) using the Had-

ley Centre Global Environment Model version 2 Earth

Amphib. Reptile Conserv.

Number of Endemic Species

Percentage of Endemism

0 0

2 iM i

0 0

28 57.1

40 41.2

10 71.4

0 0

3 24D

1 1

0 0

1 50

0 0

50 WHEE

1 Dee

2 50

0 0

1 Sid

142 a2.

0 0

14 D323

14 40

156 <=

System (Collins et al. 2011; Martin et al. 2011). More de-

tails on these analyses are available at the website.

Dynamic species guides in PDF format

Species guides are available in two formats. One is a full

PDF guide with maps, photographs, and the species ac-

count information described above. The other format is a

photographic guide, with two photographs per species, if

available. A sample of the photographic guide is included

May 2019 | Volume 13 | Number 1 | e178

Torres-Carvajal et al.

Table 3. IUCN Red List categories for reptiles from Ecuador. CA (grey columns): 2005 Red List by Carrillo et al.; IU (white

columns): 2018 IUCN Red List.

Taxon

CROCODYLIA

Alligatoridae

Crocodylidae

SQUAMATA: AMPHISBAENIA

Amphisbaenidae

SQUAMATA: SAURIA

Alopoglossidae

Anguidae

Gekkonidae

Gymnophthalmidae

Iguanidae: Corytophaninae

Iguanidae: Dactyloinae

Iguanidae: Hoplocercinae

Iguanidae: Iguaninae

Iguanidae: Polychrotinae

Iguanidae: Tropidurinae

Phyllodactylidae

Scincidae

Sphaerodactylidae

Teiidae

SQUAMATA: SERPENTES

Aniliidae

Anomalepididae

Boidae

Colubridae: Colubrinae

Colubridae: Dipsadinae

Elapidae

Leptotyphlopidae

Tropidophiidae

Typhlopidae

Viperidae

TESTUDINES

Chelidae

Cheloniidae

Chelydridae

Dermochelyidae

Geoemydidae

Kinosternidae

Podocnemididae

Testudinidae

TOTALS

477

NE DD

CA IU CA

0 0 1

0 0 0

0 2 0

1 4 4

0 0

3 2 0

2 178

0 0 0

14 34 5

4 1

1 0 0

1 2 0

8 10 fe

5 + 1

1 2 0

0 3 1

1 6 0

0 1 0

1 0 1

0 4 0

4 17 oe

40 60 36

2 10 fie

3 2

0 1

0 0 0

0 15 1

0 6 1

1 1 2

0 1 0

0 0 1

0 1 0

0 Z, 0

0 0 1

3 2 0

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IU CA

0 2

0 0

0 2

1 2

0 0

5 6

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1 15

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0 1

0 Z

0 i

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0 4

0 8

0 0

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12 fee

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28 «114

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0 0

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4 1

1 1

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11 8

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6 6

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*Chelonoidis abingdonii was listed by Carrillo et al. (2005) as extinct in the wild (EW) because the last known individual of this

species, Lonesome George, was still alive. Lonesome George died in 2012.

Amphib. Reptile Conserv.

226

May 2019 | Volume 13 | Number 1 | e178

Reptiles of Ecuador portal

with this article (Supplementary file 1). Although several

guides are available as downloadable PDF files (e.g., by

protected area or biogeographic region), users are allowed

to generate their own guides by searching the database for

specific criteria, which include taxonomy, biogeographic

region, protected area, elevation, province, and conserva-

tion status. In addition, checklists can be generated by se-

lecting any location on a map and defining a search area,

the output is a list of species of reptiles occurring within

the defined area. These dynamic checklists can be down-

loaded freely as either full or photographic PDF guides.

Results and Discussion

Reptiles del Ecuador is currently available in Spanish

at https://bioweb.bio/faunaweb/reptiliaweb, and it docu-

ments the uniquely rich reptile fauna of Ecuador. Among

countries with the highest richness of reptiles, Ecuador

has one of the largest number-of-species/area ratios in

the world (8.4 species/5,000 km’). To date, 477 species of

reptiles—35 turtles, five crocodilians, and 437 squamates

(three amphisbaenians, 197 lizards, and 237 snakes)—are

known to occur in Ecuador (Table 1). Of these, two spe-

cies of Galapagos giant tortoises are extinct due to over-

hunting; Chelonoidis niger disappeared in the mid-1850s

(Broom 1929), whereas C. abingdonii went extinct with

the death of Lonesome George in 2012. Among main-

land species (430), over one-fourth (111) are endemic to

Ecuador, whereas all terrestrial species in the Galapagos

are endemic to the archipelago, except for the introduced

geckos Gonatodes caudiscutatus, Phyllodactylus reis-

sii, Hemidactylus frenatus, and Lepidodactylus lugubris

(Torres-Carvajal and Tapia 2011). Mostly due to Galapa-

gos endemics, Testudinidae and Phyllodactylidae are the

clades with by far the highest percentages of endemism

(93.3% and 71.4%, respectively), followed by the lizard

clade Gymnophthalmidae, in which 57.1% of the species

are Andean endemics (Table 2).

In agreement with a recent study on diversity and con-

servation of Ecuadorian reptiles (Reyes-Puig et al. 2017),

the richest biogeographic areas (Fig. 2) are the Amazonian

Tropical Rainforest (154 species, ~36% of mainland spe-

cies), Western Foothill Forest (139, ~32%), and Western

Montane Forest (131, ~30%). Other areas with over a hun-

dred species are the Eastern Montane Forest (127, ~29%),

Eastern Foothill Forest (126, ~29%), Chocoan Tropical

Rainforest (121, ~28%), and Deciduous Coastal Forest

(119, ~27%). With only 15 species (~3.5%), the Paramo

is the poorest area for reptiles. Overall, the reptilian fauna

of Ecuador is remarkably dominated by dipsadine snakes

(136 species), followed by iguanid lizards (97), gymnoph-

thalmid lizards (49), and colubrine snakes (44). The most

speciose genus is Anolis, with 43 species, followed by

Atractus (29 species).

Carrillo et al. (2005) published the first red list of the

reptiles from Ecuador based on IUCN criteria. Of the 377

evaluated species, ~30% (114) were categorized as Least

Amphib. Reptile Conserv.

Concern, and ~28% (105) as Threatened with Extinction.

Over a decade later, only 54% (259) of the species of

reptiles from Ecuador have been evaluated by the IUCN

(2018). Of these, ~55% (142) are categorized as Least

Concern, and ~24% (61) as threatened with extinction

(Table 3).

Given the uniquely rich diversity and conservation sta-

tus of reptiles in Ecuador, Reptiles del Ecuador is an im-

portant and evolving resource, which can serve as a model

for the development of similar resources dedicated to the

herpetofauna of other countries.

Acknowledgements.—Reptiles del Ecuador (for-

merly called ReptiliaWebEcuador from 2010-2017) is

part of Bioweb (http://bioweb.bio), a portal to the biodi-

versity of Ecuador. For coordinating Bioweb, we thank

its general manager, Santiago R. Ron. For developing

Bioweb, we thank lead programmers Jorge Orozco and

David Zuniga. For helping to compile information on Ec-

uadorian reptiles from the literature or providing relevant

information, we thank Maria Belén Andrango, Vanessa

Aguirre, Fernando Ayala, Andrés Calero, Amaranta Car-

vajal-Campos, Denisse Galarza, Estefany Guerra, Andrés

Marmol, Diego Ortiz, Andrea Rodriguez, Cecilia Tobar,

and Jorge Valencia. Most photographs were taken and

edited by Museo de Zoologia QCAZ staff, while others

were generously donated by many photographers, who

are acknowledged in the website’s gallery. Special thanks

to Andrés Merino-Viteri for programming the HSMs, and

to Belén Baus, Sebastian Davalos, Melissa Rodriguez,

Pilar Rodriguez, Diego Paucar, and Carlos Zambrano for

help with georeferencing. This manuscript was greatly

improved thanks to critical reviews by Larry D. Wilson

and Santiago Ron. Bioweb was funded by Secretaria de

Educacioén Superior, Ciencia, Tecnologia e Innovacion

del Ecuador (SENESCYT), JRS Biodiversity Founda-

tion, and Pontificia Universidad Catolica del Ecuador.

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Amphib. Reptile Conserv.

Reptiles of Ecuador portal

Omar Torres-Carvajal graduated in Biological Sciences from Pontificia Universidad Catolica del Ecuador

(PUCE) in 1998, and in 2001 received a Master’s degree in Ecology and Evolutionary Biology from the

University of Kansas under the supervision of Dr. Linda Trueb. In 2005, Omar received a Ph.D. degree from

the same institution with the thesis entitled Phylogenetic systematics of South American lizards of the genus

Stenocercus (Squamata: Iguania). Between 2006-2008 Omar was a postdoctoral fellow at the Smithsonian

Institution, National Museum of Natural History, Washington DC, USA, working under the supervision of Dr.

Kevin de Queiroz. Omar is currently Curator of Reptiles at the Zoology Museum QCAZ of PUCE and Full

Professor in the Department of Biology, PUCE. Omar has published more than 60 peer-reviewed scientific

papers on taxonomy, systematics, and biogeography of South American reptiles, with an emphasis on lizards.

He is mainly interested in the theory and practice of phylogenetic systematics, particularly as they relate to

the evolutionary biology of lizards.

Gustavo Pazmifio-Otamendi graduated in Biological Sciences from Pontificia Universidad Catolica del

Ecuador (PUCE) in 2012, and in 2015 received a Master’s degree in Ecology, Management, and Restoration

of the Natural Environment at Universidad Autonoma de Barcelona, Spain, under the supervision of Dr. Bernat

Claramunt. For both degrees, Gustavo developed projects which analyzed animal behavior and vocalization.

Since 2012, Gustavo has been involved in the development of Reptiles del Ecuador, a comprehensive, online

encyclopedia of the reptiles of Ecuador launched by the Zoology Museum (QCAZ) of PUCE. He is mainly

interested in animal behavior, vocalization and their evolutionary aspects, as well as the taxonomy and

systematics of reptiles, particularly snakes.

David Salazar-Valenzuela graduated in Biological Sciences from Pontificia Universidad Catolica del

Ecuador in 2007. In 2016, David received a Ph.D. in Evolution, Ecology, and Organismal Biology from The

Ohio State University with a thesis entitled Diversification in the Neotropics: Insights form demographic and

phylogenetic patterns of lancehead pitvipers (Bothrops spp.), and in that year became a postdoctoral fellow at

the same institution. David is currently Curator of Reptiles at the Zoology Museum (MZUTI) of Universidad

Tecnologica Indoamérica, Quito, Ecuador, and a professor in the Department of Environmental Sciences

in the same institution. He is mainly interested in the evolution, ecology, and toxinology of Neotropical

amphibians and reptiles.

229 May 2019 | Volume 13 | Number 1 | e178

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

13(1) [General Section]: 230-234 (e179).

A new state record of the Mandarin Rat Snake Euprepiophis

mandarinus (Cantor, 1842) (Squamata: Colubridae:

Coronellini) from Mizoram, India

'2Khan Ashaharraza, *V. Rangasamy, ‘Hmar Tlawmte Lalremsanga, °Lalbiakzuala,

SJenny Sailo, and ‘Tom Charlton

'Department of Wildlife and Biodiversity Conservation, North Orissa University, Baripada 757003, Odisha, INDIA *Indian Herpetological Society,

Pune 411009, Maharashtra, INDIA *Forest Survey of India, MOEFCC, Government of India, Eastern Zone, Kolkata 700601, West Bengal, INDIA

+°Department of Zoology, Mizoram University, Aizawl 796004, Mizoram, INDIA °Deputy Conservator of Forests, Khawzawl Wildlife Division,

Department of Environment, Forests and Climate Change, Champhai, Mizoram, INDIA ’Eco Animal Encounters, 33 Brisbane, Stonehouse,

Gloucestershire-GL10 2PX, UNITED KINGDOM

Abstract.—The Mandarin Rat Snake Euprepiophis mandarinus (Cantor, 1842) is a widely distributed species

in the Indo-Chinese region with western-most limits in North East India. However, it is known from very few

localities in the extreme north-eastern states in India. Herein, we report E. mandarinus based upon a roadkill

specimen collected from the Champhai in Mizoram. The record of Champhai extends the known distribution of

the species by about 292 km SW from the localities previously recorded. Morphological variation of the Indian

population is discussed.

Keywords. Champhai, montane forests, morphological variation, Palearctic, range extension, roadkill

Citation: Ashaharraza K, Rangasamy V, Lalremsanga HT, Lalbiakzuala, Sailo J, Charlton T. 2019. A new state record of the Mandarin Rat Snake

Euprepiophis mandarinus (Cantor, 1842) (Squamata: Colubridae: Coronellini) from Mizoram, India. Amphibian & Reptile Conservation 13(1): [General

Section]: 230-234 (e179).

tion 4.0 International (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in any

medium, provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced, are

as follows: official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org.

Received: 19 August 2018; Accepted: 14 April 2019; Published: 6 June 2019

Introduction

The Mandarin Rat Snake Euprepiophis mandarinus

(Cantor, 1842) is a relatively rarely encountered spe-

cies of Old World ratsnake found in Eastern Asia (Ji et

al. 2012; Wallach et al. 2014). The type specimen was

collected from Chusan (=Zhoushan) Island, China, by T.

Cantor (Boulenger 1894). Gunther (1864) provided ex-

cellent head illustrations of Euprepiophis mandarinus.

The species is distributed in Myanmar (Prater 1919),

Vietnam (Nguyen et al. 2009), China (SFAC 2009), and

possibly Laos (Das 2012). It occurs in forest habitats,

both tropical and temperate forests, but can also be found

in grasslands and shrublands (Schulz 1996). In Vietnam,

the species is known from Na Hang, Tam Dao, and found

in karst areas in the northern mountain ranges at around

1,400-1,500 m asl (Orlov et al. 2000; Quyet and Ziegler

2004; Nguyen et al. 2009). Among the known distribu-

tion localities of this species, the nearest to India lies

within the Chin Hills, Myanmar (Prater 1919).

Euprepiophis mandarinus is a secretive species, ac-

tive in the morning and early evening (Whitaker and

Captain 2004). Based on phylogenetic studies, the pres-

ent taxonomic status of this species 1s placed under the

subfamily Colubrinae (Utiger et al. 2005). More recent

broad-scale phylogenetic studies of snakes also sup-

port this relationship (Figueroa et al. 2016). A molecular

study of Euprepiophis mandarinus suggests that these

ratsnakes originated in tropical Asia in the late Eocene,

and subsequently dispersed to the Western and Eastern

Palearctic during early Oligocene (Burbrink and Law-

son 2007). The genus Euprepiophis is subtended by the

basal node, separating them from the remainder of the

ratsnakes lineages (Chen et al. 2017). The populations of

Euprepiophis mandarinus are declining, and Wang and

Xie (2009) indicated that this species has declined by

30% over the previous decade in China.

In India, Euprepiophis mandarinus has been recorded

from only three localities and reports are separated by

long time intervals. The first specimen was collected by

Annandale (1912) from Upper Rotung Valley (=East

Siang District), Arunachal Pradesh, and was described

as Ablabes pavo, which is presently regarded as a ju-

nior synonym of Euprepiophis mandarinus (Das et al.

1998; Wallach et al. 2014). After a gap of seventy-five

years, the species was recorded again from Namdapha

Correspondence. '?ashaharrazakhan@gmail.com (corresponding author), *rangasamymu@gmail.com, *htlrsa@yahoo.co. in,

*bzachawngthu123@gmail.com, ‘jensailo@gmail.com, ‘tom@ecoanimalencounters.co.uk

Amphib. Reptile Conserv.

June 2019 | Volume 13 | Number 1 | e179

Ashaharraza et al.

by V. Rangasamy.

NP-Gandhigram road, Changlang District of Arunachal

Pradesh, and this species was regarded as rare at the site

(Sanyal and Gayen 2006). More recently it has been re-

ported from the Zunheboto District of Nagaland (Mathew

2005; Sen and Mathew 2008; Lele et al. 2018). However,

the species was never reported from Mizoram by earlier

researchers (Mathew 2007; Lalremsanga et al. 2011; Lal-

tanpuia et al. 2008).

Methods and Materials

A dead specimen, possibly killed with a sharp blade, was

collected from the roadside (Fig. 1), 14 km north of Cham-

phai town (23°36’27.1”N, 93°21713.2”E), a district capital

of Champhai in Mizoram. The specimen was fixed in 4%

formalin, preserved in 70% ethanol, and deposited in the

departmental museum of Zoology, Mizoram University,

Aizawl (MZMU-1135). The specimen was photographed

with a digital Canon Powershot SX50 HS camera. Mea-

surements were taken with a Mitutoyo™ digital caliper

to the nearest 0.01 mm, except SVL and TL, which were

measured to the nearest mm with a measuring tape. Ventral

scales were counted after Dowling (1951), and standard

external morphological characters are given in Table 1.

Additional data on distribution was obtained from the col-

lections of BNHS, Mumbai, and ZSI, Kolkata. Geographic

co-ordinates were mapped with Garmin GPSMAP® 62s.

Abbreviations

BNHS: Bombay Natural History Society, Mumbai, India;

MZMU: Museum of Zoology department, Mizoram Uni-

versity, Aizawl, India; SVL: snout to vent length; TL: tail-

length; HL: head-length; EYED: horizontal diameter of

the eye; EYEN: distance from center of the eye to posteri-

or border of the nostril; WSNT: width of the snout; VENT:

number of ventrals; SUBC: number of subcaudals and

terminal scute excluded; DOR1: number of dorsal scale

rows at one head-length behind the head; DOR2: number

of dorsal scale rows at the position of the middle ventral;

Amphib. Reptile Conserv.

Fig. 1. Dorsal and ventral view of roadkill Euprepiophis mandarinus (MZMU-1135) collected from Champhai, Mizoram. Photo

DOR3: number of dorsal scale rows at one head-length

before the tail; TEMP: number of temporals (L+R); SLI:

number of supralabials (L+R); SL2: number of supralabi-

als touching the eyes (L+R); INFR: number of infralabials

(L+R); LOR: number of loreals (L+R); POC: number of

post-oculars (L+R); NP: National Park, CHK Rd: Cham-

phai-Hnahlan-Khuangphah Road; FSI: Forest Survey of

Table 1. Detailed meristic and pholidosis data of Euprepiophis

mandarinus (measurements in mm).

Collection Voucher Number | MZMU-1135

Mizoram, India

WSNT

VENT

June 2019 | Volume 13 | Number 1 | e179

Mandarin Rat Snake in Mizoram, India

Legends

@ Previous records

4\ New locality

— State Border

— National Border

90°E 92°E 94°E 96°E

Fig. 2. Map depicting Champhai (triangle) Mizoram, a new

locality for E. mandarinus in India with previously known

localities (filled circles). Inset map shows relative position of

all localities within Republic of India.

India; DST-SERB: Department of Science and Technol-

ogy, Science and Engineering Research Board, Govern-

ment of India; EMR: Extramural Research funding.

Results and Discussion

The snake has large, elliptical yellow spots, edged with

broad black circles on dorsum and tail, lateral scales are

dull grey, mixed with yellow dorsals; the forehead has a

a 2.

~- .

= >

“ = — oe

s7.

Fig. 3. A distinct form of Euprepiophis

Amphib. Reptile Conserv.

232

black V-shaped pattern with a yellow spot between pari-

etals; the belly is yellowish-green with wide black trans-

verse blotches, which turn into entire bands on the sub-

caudal scales. The body is robust; the head is short and

slightly distinct from neck; the snout is obtuse; the tail is

short and stout; the eyes are small with a rounded pupil;

the dorsal scales are smooth; the anal is divided. This de-

scription matches with Smith (1943) and Das (2012).

The collection of the Mandarin Rat Snake Euprepio-

phis mandarinus from Champhai is the first record for

Mizoram State and the fourth record of this species from

India (Fig. 2). Champhai is located at 292 km radial dis-

tance south-east of the Zunheboto District of Nagaland,

the closest previously known locality.

The collected specimen is distinct from the individu-

als we examined from Republic of China (Fig. 3) by the

absence of loreal scales and by the absence of olive-grey

and red lateral blotches on the dorsum. Although vehicu-

lar traffic is a known threat to this species, our study has

added manual killing as another evident threat that is po-

tentially responsible for population decline of Euprepio-

phis mandarinus.

Acknowledgements.—We are __ thankful to

Vanlalchualiova (Chief Conservator of Forests, Kolasib

District of Mizoram) for encouraging the publication of

these observations. VR is grateful to Kailash Chandra

(Director, Zoological Survey of India) for material

examination and support of the study. We are thankful to

Rahul Khot for his curatorial help at BNHS. We extend

Our sincere gratitude to Rosamma Mathew for the help

with the published literature. We acknowledge Ashok

Captain for his comments and personal communication

during the writing of the manuscript. Deepest regards to

mandarinus with red-brown dorsals from Republic of China. Photo by Tom Charlton.

June 2019 | Volume 13 | Number 1 | e179

Ashaharraza et al.

Anil Khaire (Chairman, Indian Herpetological Society)

for his guidance. Additional thanks are due to Suchetana

Sen, Mufaddal Shakir, Vishal Varma, Prasad Gond, Ravi

Pawar, Madhao Vaidya, and Abhijeet Dani for assisting

in the Indo-Burmese reptile survey. We are thankful to

Gernot Vogel, Deepak V., and an anonymous reviewer

for comments on an earlier version of this paper. At North

Orissa University, KA is thankful to Cuckoo Mahapatra,

Rabindra Kumar Mishra, and Shrustidhar Rout for their

input during manuscript construction. VR acknowledges

Dr. C. Vidhyasagar (DFO, Lunglei, Mizoram, India) and

Mahalaxmi (Annamalaiputhur, Tamil Nadu, India) for

continuous support and help. The collection of the present

specimen from Champhai was possible under permission

No. A.33011/2/99-CWLW/225 issued by Chief Wildlife

Warden, Environment, Forest and Climate Change

Department, Government of Mizoram, India. HTL and

LBZ extend their gratitude to DST-SERB, New Delhi

for providing laboratory facilities under EMR number

EMR/2016/002391. At FSI, our humble thanks to Ajaya

Kumar Nayak and Ashok Kumar Biswal for support and

encouragement.

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Amphib. Reptile Conserv.

Khan Ashaharraza, fondly known as Krishna Khan, is a central Indian herpetologist. Ashaharraza has received

many grants and accolades, and is a reputed member of Indian Herpetological Society. With a Master’s degree in

Wildlife and Biodiversity Conservation, he mentors the research of M.Sc. students as well as amateurs. Ashaharraza

is one of the pioneers of herpetological research in Vidarbha, and has published scientific communications from all

four biodiversity hotspots in India. His research projects now include the Indo-Burmese Reptile Survey, Natural

History and Biogeography of Indian Reptiles, and Avifauna of Central India.

V. Rangasamy is a government servant in the Forest Survey of India, under the Ministry of Environment, Forests

and Climate Change, Government of India. V. Rangasamy is also a Ph.D. scholar at the Zoological Survey of India

(ZSI), affiliated with Pondicherry University, focusing on the natural history and phylogenetics of Polypedates

insularis, an endangered tree frog of Great Nicobar Island, India. He worked for ZSI as a JRF and SRF in the

Andaman and Nicobar Islands under the supervision of Dr. Raghunathan (Scientist-E, ZSI, ANRC, Port Blair) and

Dr. Sivaperuman (Scientist-D, ZSI, ANRC, Port Blair). His research interests include the taxonomy, distribution,

and behavioral study of amphibians, reptiles and, plants of the north eastern states of India.

H.T. Lalremsanga is a northeast Indian zoologist, whose Ph.D. work in amphibian biology was at North Eastern

Hill University in 2011. He is working as Assistant Professor in the department of Zoology, Mizoram University, and

has described new species of frogs, eels, and snakes, and a new genus of snakes. He is interested in the systematics

and biology of amphibians and reptiles, and has established the Developmental Biology and Herpetology Lab in

which he guides his research scholars and M.Sc. students.

Lalbiakzuala is currently working on his M.Phil. degree under the supervision of Dr. H.T. Lalremsanga in the

Department of Zoology, Mizoram University, India. Lalbiakzuala completed his Master’s in Zoology from Mizoram

University, with a thesis on Preliminary Study on Helminth Parasites of Snakes in Mizoram, Northeast India.

Jenny Sailo is a member of the Mizoram Forest Service (2009 batch). Jenny is currently working as Divisional

Forest Officer, Khawzawl Wildlife Division, in the Environment, Forest and Climate Change Department,

Government of Mizoram, India. Jenny graduated from North Eastern Regional Institute of Science and Technology

(NERIST) in Nirjuli, Arunachal Pradesh, and obtained a post-graduate degree in Forestry from Mizoram University,

Mizoram. Her fields of interest include wildlife management, forest conservation, and the sustainable conservation

of Biodiversity.

Tom Charlton is a British herpetologist currently working as a coordinator in a snakebite mitigation program in

Central Province, Papua New Guinea. Back home in the United Kingdom, Tom works with several expedition and

wilderness medicine training providers to develop and present specialist lectures on snakebite first aid and mitigation

strategies for medics working overseas. Tom’s interests lie largely with the herpetofauna of Southeast Asia, and

its snakes in particular, and he has authored the book King Cobra: Natural History and Captive Management,

published in January 2018.

234 June 2019 | Volume 13 | Number 1 | e179

Mannophryne vulcano, Male carrying tadpoles. El Avila (Parque Nacional Guairarepano), Distrito Federal. Photo: Jose Vieira.

We want to dedicate this work to some outstanding individuals who encouraged

us, directly or indirectly, and are no longer with us. They were colleagues and close

friends, and their friendship will remain for years to come.

—

Erik Arrieta Marquez (1978-2008)

Jose Ayarzaguiena Sanz (1952-2011) Saul Gutiérrez Eljuri (1960—201 2)

Fy T .

a! | :

Juan Rivero (1923-2014) Luis Scott (1948-2011)

Marco Natera Mumaw (1972-2010)

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

13(1) [Special Section]: 1-198 (e180).

Catalogue of the amphibians of Venezuela:

lllustrated and annotated species list,

distribution, and conservation

‘2Cesar L. Barrio-Amoros, *4Fernando J. M. Rojas-Runjaic, and °J. Celsa Sefaris

'Fundacién AndigenA, Apartado Postal 210, Mérida, VENEZUELA *Current address: Doc Frog Expeditions, Uvita de Osa, COSTA RICA

3Fundacion La Salle de Ciencias Naturales, Museo de Historia Natural La Salle, Apartado Postal 1930, Caracas 1010-A, VENEZUELA *Current

address: Pontificia Universidade Catélica do Rio Grande do Sul (PUCRS), Laboratorio de Sistemdtica de Vertebrados, Av. Ipiranga 6681, Porto

Alegre, RS 90619-900, BRAZIL *Instituto Venezolano de Investigaciones Cientificas, Altos de Pipe, apartado 20632, Caracas 1020, VENEZUELA

Abstract.—Presented is an annotated checklist of the amphibians of Venezuela, current as of December 2018.

The last comprehensive list (Barrio-Amoros 2009c) included a total of 333 species, while the current catalogue

lists 387 species (370 anurans, 10 caecilians, and seven salamanders), including 28 species not yet described

or properly identified. Fifty species and four genera are added to the previous list, 25 species are deleted, and

47 experienced nomenclatural changes. Eleutherodactylus terraebolivaris Rivero, 1961 is synonymized with

Hylodes incertus Lutz, 1927 as Pristimantis incertus. Oreophrynella dendronastes Lathrop and MacCulloch,

2007, is considered a junior synonym of O. macconnelli (Boulenger 1895). Centrolene Jiménez de la Espada,

1872, is a feminine genus, so all species in the genus are amended. Centrolenella pulidoi Rivero, 1968 is

considered a junior synonym of Hyla benitezi Rivero, 1961, as Boana benitezi. Centrolenella estevesi Rivero,

1968 is considered a junior synonym of Hyla jahni Rivero, 1961, as Hyloscirtus jahni. \llustrated herein are 300

species (77.5% of the total). Lastly, the distributions for all species are revised, species that possibly occur

within Venezuela are suggested, and comments are provided on nomenclature and conservation issues.

Keywords. Biogeography, checklist, Anura, Urodela, Gymnophiona, management

Resumen.—Se presenta una lista anotada de los anfibios de Venezuela, actualizada hasta diciembre de 2018.

La ultima lista comprensiva (Barrio-Amoros 2019c) incluyo un total de 333 especies, mientras que la lista actual

contiene 387 especies (370 anuros, 10 cecilias y siete salamandras), incluyendo 28 especies aun no descritas

o identificadas propiamente. 50 especies y cuatro generos se anaden a la lista previa, 25 especies se eliminan

y 47 de ellas han experimentado cambios nomenclaturales. Eleutherodactylus terraebolivaris Rivero, 1961 se

sinonimiza con Hylodes incertus Lutz, 1927, como Pristimantis incertus. Oreophrynella dendronastes Lathrop

and MacCulloch, 2007, se considera sinonimo de O. macconnelli (Boulenger, 1895). Centrolene Jiménez de la

Espada, 1872, es un género femenino, asi que se emendan todos los nombres acorde. Centrolenella pulidoi

Rivero, 1968 es considerado sinonimo de Hyla benitezi Rivero, 1961, como Boana benitezi. Centrolenella

estevesi Rivero, 1968, se considera sinonimo de Hyla jahni Rivero, 1961, como Hyloscirtus jahni. Se presentan

fotografias de 300 especies (77.5% del total). Por ultimo, la distribucion de todas las especies es revisada, se

sugieren especies que podrian estar presentes en Venezuela y se presentan comentarios sobre nomenclatura

y conservacion.

Palabras clave. Anura, biogeografia, Gymnophiona, lista, manejo, Urodela

Citation: Barrio-Amorés CL, Rojas-Runjaic FUM, Sefaris JC. 2019. Catalogue of the amphibians of Venezuela: Illustrated and annotated species list,

distribution, and conservation. Amphibian & Reptile Conservation 13(1) [Special Section]: 1-198 (e180).

tribution 4.0 International (CC BY 4.0): https://creativecommons.org/licenses/by/4.0/], which permits unrestricted use, distribution, and reproduction in

any medium, provided the original author and source are credited. The official and authorized publication credit sources, which will be duly enforced,

are as follows: official journal title Amphibian & Reptile Conservation; official journal website: amphibian-reptile-conservation.org.

Received: 27 August 2016; Accepted: 26 April 2018; Published: 14 July 2019

Introduction amphibian diversity, after Brazil (1,026 species; http://

br.herpeto.org/anfibios/), Colombia (813; — https://

In consideration of the findings presented herein, | www batrachia.com), Peru (657; http://research.amnh.

Venezuela (387 species) will be the seventh- org/vz/herpetology/amphibia/index.php//content/

most diverse country in the world with respect to search?taxon=&subtree=&subtree_id=&english na

Correspondence. '?cesarlba@yahoo.com, *4rojas_runjaic@yahoo.com, *celsisenaris@gmail.com

Amphib. Reptile Conserv. 1 July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

400

o Anura

= J

350 B Urodela

O Gymnophiona

300 w Total

LF]

2 250

ro)

“~ 200

—

es !

o 150

Z 100

‘5 I i f |

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Fig. 1. Number of amphibian species known in Venezuela since 1927.

me=&author=&year=&country=167), Ecuador (594;

https://bioweb.bio/faunaweb/amphibiaweb/), Indonesia

(457; http://research.amnh.org/vz/herpetology/amphibia/

index.php//content/search?taxon=&subtree=&subtree_

id=&english_name=&author=&year=&country=1 66),

and Mexico (392; Parra-Olea et al. 2014; https://

amphibiaweb.org/amphibian/newspecies. html).

Several checklists for Venezuela, with varying

levels of detail, have appeared since the 1950s (Fig.

1), and clearly the number of known species has grown

with time and increased inventory activity, from 74

frog species in 1959 to 370 in 2019. The most data-

intensive and continuously updated listing of the world’s

amphibian species is currently Amphibian Species of

the World 6.0 (http://research.amnh.org/vz/herpetology/

amphibia/index.php), an Online Reference whose book

predecessor was published by Darrel Frost in the mid-

1980s (Frost 1985). Another important online source

is the AmphibiaWeb site (http://www.amphibiaweb.

org). Although both online databases are quite useful

as references, they lack precision in their ability to

specifically identify local species, and should not be

taken as the final taxonomic authority (Dubois 2017). It

is also unwise to rely on a third major online database,

the IUCN Red List of Threatened Species (UCN 2015),

since entries often lag behind current research, and with

few exceptions, corrections and updates are irregular.

Ten years after the last published official checklist

(Barrio-Amorés 2009c), the panorama has changed

again in many ways. Several new species have been

described, others reported as new for the country, and

Amphib. Reptile Conserv.

many were deleted from the list due to synonymy or

misidentification. The suprageneric systematics are still

in flux and subject to numerous, and sometimes shocking,

changes as well as errors (see Heinicke et al. 2009; Pyron

and Wiens 2011; De Sa et al. 2012, 2014; Padial et al.

2014; Duellman et al. 2016). Fora relatively stable listing

in the face of significant taxonomic changes, the reader is

advised to consult Frost (2019). This publication focuses

on concerns with taxonomic and nomenclatural issues

relevant to Venezuelan amphibians and not on major,

broad changes in species relationships or names.

Material and Methods

Several methods were used to clarify how the

determination of the taxonomic status was made for some

taxa. The majority of species are listed by their generic

and specific names, under the Family level. Not used are

unranked taxa uniting families (Arboranae, Terraranae),

which can facilitate classification uses in some cases

(Hedges et al. 2008; Duellman et al. 2016). Rather,

families are placed in alphabetical order. Where there are

doubts regarding specific identity, three disclaimers are

used. The first is the placement of aff (for affinis, from

the Latin for closely related or akin, but not identical to)

between the generic and specific names. This is used

to identify taxa for which evidence indicates existing

differences from a named, very similar taxon, but for

which this difference has not yet been further evaluated.

Thus, a species may be presented under a name, and a

second entry may be made for the species name with aff

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

inserted. A second disclaimer 1s cf (for confer, from the

Latin for compare with), used in cases where there is

insufficient evidence to connect the form to an existing

name, or to determine whether a close affinity exists. The

last disclaimer is the addition of the term “sp.” to only the

generic name, for forms with no current specific name

which may represent undescribed or uninvestigated

species.

This update is produced with a cut-off date of 1

January 2019, based on the earlier lists by Barrio-

Amoros (2009c, 2013). Six appendices complete the

information as follows: Appendix 1: Species known to

inhabit Venezuela, but not yet described or correctly

identified; Appendix 2: Taxonomic checklist of the

amphibians of Venezuela; Appendix 3: List of amphibian

species of Venezuela by biogeographic areas; Appendix

4: Additions of new species for Venezuela between 2009—

2018; Appendix 5: Species deleted here from previous

lists or reported after and deleted herein; and Appendix

6: Taxonomic changes at the generic and specific levels,

with emendations that occurred since Barrio-Amoros’

(2009c) last systematic list, including changes to new

species added in the previous list. For each species,

the holotype, type locality, overall distributions,

and Venezuelan distributions (as the biogeographic

regions and generic or specific localities are noted) are

referenced, but extensive locality lists are not provided.

New localities are added when determined to be valid and

useful. For a more complete listing of locality data see

Barrio-Amoros (1998) and Gorzula and Sefiaris (1998).

Taxonomic or biogeographic comments are added (under

Remarks). Relevant new localities or noteworthy shifts

in the reported distribution are noted in the Distribution

section of the accounts, listing vouchers when possible.

An asterisk after the species authority indicates that the

species is endemic to Venezuela. To avoid repeating the

work by Frost (2019), synonymies are not provided under

each species, but rather the reader can direct their attention

to an extensive online database (http://research.amnh.

org/vz/herpetology/amphibia/index.php). The current

conservation status of each species can be checked in

both the general (http://www.iucnredlist.org) and local

Red Lists (http://animalesamenazados.provita.org.ve/)

that are constantly being updated. In alphabetical order,

selected relevant literature dealing only with Venezuelan

taxa follows each account. References dealing with

the majority of species (e.g., La Marca 1992; Barrio-

Amoros 1998, 2004; Frost 2019) are not repeated in each

account. Acronyms follow Barrio-Amoros (1998, 2004),

Frost (1985, 2016), and Sabaj (2016) with the addition

of MBLUZ (Museo de Biologia, La Universidad del

Zulia, Maracaibo, Venezuela), IRSNB (Institut Royal

des Sciences Naturelles de Belgique, Brussels, Belgium),

and MUSM (Museo de Historia Natural Universidad

San Marcos, Lima, Peru). The International Code on

Zoological Nomenclature (ICZN) [http://www.iczn.org/

iczn/index.jsp]| is hereafter referred to as the Code.

Amphib. Reptile Conserv.

Biogeography

Rivero (1961) was the first to present a zoogeography

of Venezuelan amphibians. His approach was insightful,

especially considering the level of knowledge at the time

and lack of Internet resources, and it has only changed

in detail over time. Barrio-Amorés (1998) excluded

the Falcon Region as it has not been reported as having

biogeographic importance for amphibians (though it

is important for reptiles; see Rivas et al. 2012). Péfaur

and Rivero (2000) presented a biogeographic scenario

encompassing both amphibians and reptiles, and in so

doing included coastal and island ranges, and returned

to consider a Falcon-Lara xerophytic region. Sefiaris and

Rojas-Runyjaic (2009) analyzed and commented on the

distribution and biogeography of Venezuelan amphibians

without further changes.

Barrio-Amoros (1998, 2013) and Molina et al. (2009)

presented the biogeographic patterns of Venezuela

(mainly based on Rivero 1961) adapted for amphibians,

with seven major bioregions, including the Andes; the

Cordillera de la Costa (coastal mountain range); the Llanos

(the great plains); the Amazonian Region (including

the eastern versant of the Andes); the Venezuelan and

Guayanan including the Pantepui (the Venezuelan

part of the Guiana Shield); the Orinoco Delta; and the

Maracaibo Lake Basin. This arrangement is followed

herein with one significant change: the Orinoco Delta is

not considered a standalone region any longer, since it

actually represents an assembly fed by the herpetofauna

from the Llanos, the Amazonian Region, and the Guiana

Shield, with no significance for amphibians on its own.

Not a single species 1s known to inhabit only the Orinoco

Delta or to be endemic there (Barrio-Amoroés 2004).

Gorzula and Sefiaris (1998) also included the Delta of

the Orinoco as a part of the Venezuelan Guayana. The

Guayana Esequiba, a territory administered by Guyana

and historically claimed by Venezuela, is not considered

herein.

Venezuela is located on the northern coast of South

America, between the Equator and 12° North, and the

Western Meridians 60° and 74°. It is delimited in the

north by the Caribbean Sea and in the northeast by the

Atlantic Ocean; it borders Brazil to the south, Guyana

to the east, and Colombia to the west. It has a landmass

area of 916,445 km? and a coastline of 4.261 km (Barrio-

Amoros 1998) [Fig. 2].

Venezuela’s Bioregions

1. The Andes

The great Andean mountain range extends into Venezuela

for about 450 km. Within the borders of the country it has

a maximum width of about 100 km, and peaks reach a

maximum elevation of almost 5,000 m. Only two small

glaciers still exist, although their sizes have diminished

July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

Maracaibo"

lakebasin’

Western

Llanos

Eastern

Llanos

* 56 Lae

[evisnaishi leldy

Fig. 2. Map of Venezuela corresponding to the current geographic limits. The major bioregions mentioned in the text can

be distinguished along with the main physiographic features. Some noteworthy geographic elements are numbered. 1. Serrania

de Perija (Zulia state); 2. Sierra de San Luis (Falcon state); 3. Rancho Grande Biological station in Henri Pittier National Park

(Aragua state); 4. Puerto Ayacucho and surroundings (Amazonas state); 5. Cerro Autana (Amazonas state); 6. Serrania de la Neblina

(Amazonas state); 7. Source of the Orinoco River, between Serranias Tapirapeco and Parima (Amazonas state); 8. Cerro Duida,

Duida-Marahuaka National Park (Amazonas state); 9. Sarisarifiama-tepui, in Jaua-Sarisarifiama National Park (Bolivar state); 10.

Guaiquinima tepui (Bolivar state); 11. Auyan-tepui (Bolivar state); 12. Chimanta massif (Bolivar state); 13. Roraima-tepui (Bolivar

state); 14. Sierra de Lema (Bolivar state).

in recent years. Several habitats provide excellent

conditions for amphibians (Barrio-Amords 1998),

especially the sub-Andean forest, deciduous forest, and

cloud forest that collectively extend from 600-—3,000

m, and the paramos that range from about 2,400 m to

over 4,600 m (Rivero 1961). A dry-climate depression

in Tachira State (Depresion del Tachira) at elevations

as low as 600 m acts as a natural barrier to dispersal of

high-Andean species between the Cordillera Oriental

de Colombia (of which the only part in Venezuela is the

massif of Tama) and the Cordillera de Mérida.

The Sierra, or Serrania de Perijd, in the northwestern

corner of Venezuela, is also part of this montane

environment. It is about 240 km long and 35 km wide,

with a maximum elevation of 3,750 m. The Venezuelan

Andes lie entirely within the states of Barinas, Lara,

Merida, Tachira, Trujillo, and Zulia, with only a small

area in Apure State.

Currently, 128 species of amphibians are known from

the Venezuelan Andes (120 anurans, two caecilians,

Amphib. Reptile Conserv.

and six salamanders) that account for 32.9% of the

total number of species in Venezuela. Of these Andean

species, 78 (61%) are endemic to Venezuela with known

localities only in the Andes. This is the greatest amphibian

endemicity for any biogeographic region in Venezuela.

2. The Coastal Range

The furthest outreaches of the Andes form the Coastal

Range or “Cordillera de la Costa” (Rivero 1964a)

that runs west-to-east across the entire northern part

of the country, from Falcon and Yaracuy States to the

westernmost state of Sucre (and ultimately continuing

onto the continental-shelf island of Trinidad). The

Coastal Range has two distinct physiographic formations

separated by the valley of the Unare River, which cuts

across the mountain range and allows penetration of the

eastern Llanos ecosystem and herpetofauna towards the

Caribbean (Barrio-Amoroés 1998). The western part of

the Coastal Range lies in the states of Falcon, Yaracuy,

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

Carabobo, Aragua, Miranda, and Guarico, including

the Capital District around the capital city, Caracas.

The eastern part lies mainly in the states of Anzoategul,

Monagas, and Sucre, forming the mountains of the

Turimiquire Massif and the Serrania de Paria (Kaiser et

al. 2015). The typical vegetation is evergreen cloud forest

at elevations over 900 m. Maximum elevation is slightly

above 2,500 m (e.g., Naiguata 2,765 m; Turimiquire

2,630 m), although it never snows there. The amphibian

fauna includes 89 species (86 frogs, two caecilians, and

one salamander) and accounts for 23.2% of Venezuela’s

batrachofauna. Fifty-three (60%) of the Coastal Range

species are endemic to Venezuela and 61 (68.9%) occur

only in the Coastal Range.

3. The Llanos

The Llanos of Venezuela occupy an area of more than

180,000 km’. The main characteristic of the landscape

is the predominance of extensive savannahs irrigated

by large, seasonally-fluctuating rivers. The climate

is distinctive, encompassing two markedly different

seasons (dry and wet). The change from a dry season

in January-April to an extremely rainy one in May-

November markedly alters the living conditions of the

biota. The floods of the rainy season change the landscape

dramatically, from green savannah to a seemingly endless

lake. The temperature averages about 24 °C (ranging from

19-35 °C during the day) but commonly stays above 23

°C even at night. The biotopes are gallery forests fringing

the watercourses, savannahs of diverse vegetation with

temporary and/or permanent ponds, palm fields known as

morichales, as well as rivers and lagoons (Rivero-Blanco

and Dixon 1979).

Three major landscapes can be distinguished in

the Llanos. The upper Llanos situated to the West are

forested and connected with the Amazon biotope through

the Andean piedmont. The lower Llanos are endless

savannahs with scattered trees and gallery forest only

along rivers. The eastern Llanos are primarily covered

in shrubby vegetation. Included in the Llanos are the

savannahs located in the states of Apure, Anzoategul,

Barinas, Cojedes, Guarico, Monagas, northern Amazonas,

and northern Bolivar. The fauna of the Llanos is less-

diverse than that of the other Venezuelan bioregions,

although population densities of individual species

can be extremely high. In total, there are 30 species of

amphibians (29 frogs and one caecilian) constituting

7.8% of the total Venezuelan amphibian fauna. None of

them is endemic either to the Llanos or to Venezuela.

4. The Amazonian Region

The vast Amazon Basin extends into Venezuela as

lowlands (up to 200 m in elevation) through the Casiquiare

canal, a natural connection between the Amazon and

Orinoco Basins that flows southwards from the Orinoco

Amphib. Reptile Conserv.

and reaches as far north as the Sipapo and Autana Rivers.

The Amazonian Region (as a bioregion) must not be

confused with Amazonas State, which is a political unit

comprising the Amazonian Region, but also part of other

bioregions; the Amazon Region occupies a third of the

Amazonas State while the rest is of Orinoquian origin

(Gorzula and Sefiaris 1998). The region is characterized

by a perennially humid forest that is furrowed by a

complex net of cafios and tributaries to the Orinoco and

Rio Negro (Barrio-Amoros 1998).

As in the Llanos, two seasons are distinguishable;

the rainy season causes floods in the lowlands, and even

during the so-called dry season there are frequent rains.

The climate is hot and humid, with an annual mean

temperature varying from 24—27 °C (Lizot 1988). The

Amazonian Region 1s an extremely favorable biotope

for amphibians, though it does not constitute a center

of speciation in Venezuela. There are only 70 known

amphibian species recognized from the region (66

frogs and four caecilians) that constitute 18.4% of the

total Venezuelan amphibian fauna; none are endemic to

Venezuela. The Amazonian Region is a known corridor

for the expansion of amphibians into other biomes.

Amazonian herpetofaunal elements have penetrated

the eastern versant of the Venezuelan Andes (Barrio-

Amoros 1998) as indicated by the presence of a variety

of typical Amazonian species, such as Bolitoglossa aff.

altamazonica (but see Barrio-Amoros et al. (2015a)),

Boana boans, Scinax wandae, Lithodytes lineatus, and

Rhaebo glaberrimus (Barrio et al. 1999, 2002; Barrio-

Amoros 1999a, 2001b; Chacon et al. 2000; Barrio and

Chacon 2002; Schargel and Rivas 2003). Although

previous authors have used the term “Orinoco Delta”

to define a distinct bioregion, this concept 1s inadequate

since the fauna there is primarily composed of elements

from the Amazonian and the Llanos Regions, with some

Guiana Shield influences.

5. The Venezuelan Guayana

The Venezuelan part of the Guiana Shield is considered

here as following the concept developed by Hoogmoed

(1979), Barrio-Amoroés (1998), and Duellman (1999),

but it is not in complete agreement with the definition

used by Gorzula and Sefiaris (1998), who extended it

to the southern section of the state of Sucre. Herein, the

Venezuelan Guayana is delimited by the northern part

of the Orinoco Delta. This area is the most extensive

bioregion of Venezuela and the most ecologically

complex. It is one of the oldest geological regions on

Earth, dating back to the Precambrian (Aubrecht et

al. 2012). Geographically, it includes Delta Amacuro,

Bolivar, and Amazonas States, except for the savannahs

in the north that border the southern shore of the Orinoco

River and which are considered part of the Llanos, and

the lowland rainforest west of the Parima-Tapirapeco

mountain chain, which are typically Amazonian Region

July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

elements. The Gran Sabana, a savannah-covered plateau,

encompasses elevations from 800-1,400 m, and is

included in the Venezuelan Guayana as it is not linked

geographically to the similar savannahs in the north

of Bolivar and Amazonas States. Characteristic of the

Venezuelan Guayana are high tabletop mountains known

as tepuis, composed of Precambrian sandstone. Some

of the more famous tepuis include Cerro La Neblina

(elevation 3,014 m), Monte Duida (2,480 m), Roraima

(2,810 m), Sarisarifiama (2,100 m), and Auyantepui

(2,460 m). Approximately 100 tepuis occur in Venezuela

and there are additional tepuis in Guyana, Suriname,

and northern Brazil (Brewer-Carias 1988). Surrounding

these “insular” habitats 1s a wide variety of biotopes. The

most extensive ones are evergreen rainforest, similar to

the forests of the Amazonian Region, and cloud forest on

the slopes of each tepui (from 600—1,500 m elevation).

The climate is divided into two seasons, dry (December-

May) and rainy (June-November). Temperatures in

the lowlands oscillate between 24 °C and 27 °C. An

introduction to the region and an overview of the tepui

biota was provided by Aubrecht et al. (2012). Sefiaris

et al. (2014) provided a useful guide to the amphibians

of the Gran Sabana Region. The amphibian fauna of the

Venezuelan Guayana includes 163 species (156 frogs and

seven caecilians) that account for 42.8% of the amphibian

species in the country. Sixty-five species (17.1% of the

total and 39.8% of the bioregion) are endemic to the

tepuis.

6. The Maracaibo Lake Basin

The basin of Lake Maracaibo occupies parts of the states

of Zulia and Trujillo, and the northern parts of Mérida

and Tachira in northwestern Venezuela, as well as the

western versant of the Serrania de Siruma in Falcon

State. The basin contains xerophilous vegetation in the

coastal areas of the North (Barrio-Amoros 1998; Infante-

Rivero 2009). Toward the south this formation grades

into original pluvial forest and swamps along a gradient

of increasing humidity and more forested conditions.

However, the area is currently being deforested and

more pastures are being established, resembling those

established in the Llanos Region. In addition, extensive

plantations of banana and African Oil Palm now replace

part of the natural landscape. This region contrasts with

the Andes (with the Cordillera de Mérida to the east

and the Serrania de Perija to the west) that surround it

above an elevation of 250 m. Lake Maracaibo, the largest

natural lake in South America, 1s fed mainly by rivers

from the Sierra of Perija and from the Andes in the states

of Merida and Trujillo. There are 24 known species of

amphibians (23 frogs and one caecilian) that account for

6.3% of the amphibian species in the country, with no

endemics.

Amphib. Reptile Conserv.

SYSTEMATIC ACCOUNTS

Class AMPHIBIA Linnaeus, 1758

Order ANURA Fischer von Waldheim, 1813

Family Allophrynidae Gaige, 1926

Selected references: Savage 1986; Duellman 1993;

Barrio 1998; Frost et al. 2006; Guayasamin et al. 2009;

Castroviejo-Fisher et al. 2012; Caramashi et al. 2013.

Genus Allophryne Gaige, 1926

Type species: A/lophryne ruthveni Gaige, 1926, by

original designation.

Allophryne ruthveni Gaige, 1926

Holotype: UMMZ 63419.

Type locality: “Tukeit Hill, below Kaiteur Falls, British

Guiana.”

Distribution: Widely distributed in the Amazonian

Region of Venezuela, Guyana, and Suriname into central

Brazil. Regions 4 and 5. In Venezuela, known from

several localities south of the Orinoco River, in states

Amazonas (Surumoni) and Bolivar (upper Caura River,

upper Paragua River, Sierra de Imataca, San Martin de

Turumban, Cuyuni, upper Uey River). See Duellman

(1997); Barrio-Amorods (1998); Sefiaris and Rivas

(2008), and Sefiaris et al. (2009).

Remarks: Until recently, A//ophryne was considered

a monotypic genus, containing only A. ruthveni. Two

additional species have been described: A. resplendens

from the upper Amazon in Peru (Castroviejo-Fisher et

al. 2012) and A. relicta from the Mata Atlantica Forest in

Brazil (Caramaschi et al. 2013).

Selected references: Lynch and Freeman (1966);

Hoogmoed (1969); Rivero et al. (1986); Duellman

(1997); Caldwell and Hoogmoed (1998); Fabrezi and

Langone (2000); Sefiaris and Rivas (2008); Sefiaris et al.

(2009); Castroviejo-Fisher et al. (2012); Caramaschi et

al. (2013).

Family Bufonidae Gray, 1825

Remarks: Frost et al. (2006) proposed a monophyletic

taxonomy for several bufonid clades, such as Rhaebo for

the Bufo guttatus species group, Rhinella for the Bufo

margaritifer species group, and Chaunus for a number of

other South American species groups of Bufo. Chaparro

et al. (2007) embedded the genus Chaunus within

Rhinella. Bufo nasicus shown to be basal to Rhaebo by

implication; however, implication is not a resolution,

but merely implicates the need for elucidation and

further evidence. Barrio-Amoros (2009) supposed Bufo

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

nasicus to be a member of Rhinella based on external

characters and transferred it to that genus without further

clarification. Barrio-Amords and Castroviejo-Fisher

(2008a) considered Rhaebo anderssoni a synonym of

R. guttatus. Genus Dendrophryniscus Jiménez de la

Espada, 1870 was split by Fouquet et al. (2012a), and the

Amazonian-Guianan clade was then named Amazonella.

Fouquet et al. (2012b) realized that Amazonella was a

preoccupied unavailable name according to the Code and

proposed Amazophrynella as a replacement name for the

clade.

Genus Amazophrynella Fouquet, Recoder, Teixeira,

Cassimiro, Amaro, Camacho, Damasceno, Carnaval,

Moritz, and Rodrigues, 2012

Type species: Atelopus minutus Melin, 1941.

Amazophrynella is a replacement name (see above) for

Amazonella (Fouquet et al. 2012a).

Amazophrynella minuta (Melin 1941)

Holotype: BM 1968.940.

Type locality: Taracua, Uaupés, Brazil.

Distribution: Widely dispersed in Amazonia occurring

in Brazil, Ecuador, Colombia, Bolivia, the Guianas, and

Peru. In Venezuela, known only from near Raudal de

Danto, Cuao River, Amazonas State (Rojas-Runjaic et al.

2013).

Remarks: Amazophrynella minuta is an abundant

terrestrial species of the forest floor leaf litter throughout

its distribution. In Venezuela it is very rare. Only one

specimen found in many years of monitoring amphibians

in Amazonas State (Rojas-Runjaic et al. 2013). Fouquet et

al. (2012a) suggested a different species of the A. minuta

complex may exist in the Guianas and, thus, populations

currently considered to belong to species A. minuta need

to be better defined to delineate specification. Rojas et al.

(2016) further examined the A. minuta complex.

Selected references: Fouquet et al. (2012a,b); Rojas-

Runjaic et al. (2013); Rojas et al. (2014).

Genus Atelopus A.M.C. Dumeril and Bibron, 1841

Type species: Atelopus flavescens A. M C. Dumeril

and Bibron, 1841, by monotypy.

Atelopus carbonerensis Rivero, 1972*

Holotype: BM 1968.940.

Type locality: “La Carbonera (San Eusebio), 2,330 m,

Estado Mérida, Venezuela.”

Distribution: Region |. Paramo el Tambor; surroundings

of the type locality.

Remarks: Considered the most endangered species of the

genus in Venezuela, probably extinct (Rueda-Almonacid

et al. 2005). The last sighting of a live specimen was in

Amphib. Reptile Conserv.

1998 (Torres and Barrio-Amoroés 2001). La Carbonera

has been extensively deforested and used as pasture land,

and little original cloud forest remains in the Sierra de

la Culata National Park. Recent sightings (as recent as

2016) have not been proven, and must be verified.

Selected references: Rivero (1972, 1980); Dole and

Durant (1974); Durant and Dole (1974a,b); La Marca

(1984a, 1992); Rodriguez and Rojas-Suarez (1995,

1999); Lotters (1996); La Marca and Lotters (1997),

Torres and Barrio-Amords (2001); Barrio-Amoroés

(2001c, 2013); Rueda-Almonacid et al. (2005); Barrio-

Amoros and Torres (2010).

Atelopus chrysocorallus La Marca, 1994*

Holotype: ULABG 1814.

Type locality: Venezuela: Estado Trujillo: Distrito

Bocono: La Aguada, a stream flowing to Quebrada El

Molino (a tributary of rio Burate), nearly 2.5 km South

of Niquitao, 2,200 m.

Distribution: Region 1. Known only from type locality.

Remarks: Garcia-Pérez et al. (2013) reported several

hundred tadpoles in creeks in the type locality between

September 2010 and November 2011, but found no adult

specimens. A visit to the type locality in January 2016 by

FRR also failed to detect any adult individuals.

Selected references: La Marca (1994d); La Marca and

Lotters (1997); Barrio-Amorés (2001, 2013); Rueda-

Almonacid et al. (2005); Garcia-Pérez et al. (2013).

Atelopus cruciger

(Lichtenstein and von Martens 1856)*

Neotype: ZSM 93/1947/10

Type locality: Veragua, Panama (error); corrected to the

locality of the neotype: “vicinity of Rancho Grande on

the road from Maracay to Ocumare de la Costa (ca. 1,000

m above sea level), Estado Aragua, Venezuela” (Lotters

and La Marca 2001).

Distribution: Region 2. Known from at least 30 historical

localities in the Cordillera de la Costa, from Aragua,

Carabobo, Cojedes, Miranda, Vargas, and Yaracuy States,

as well as the Distrito Capital (Manzanilla and La Marca

2004). Currently, considered extinct at all localities

except for two populations in Aragua State (Rodriguez-

Contreras et al. 2008). A third surviving population was

mentioned by Rodriguez and Rojas-Suarez (2008); its

current status is unknown.

Remarks: The nomenclatural history of the species was

detailed by Lotters et al. (1998) and Lotters and La Marca

(2001). Lampo et al. (2011) provided data on the current

status of populations. Gonzalez et al. (2010) studied the

diet in museum specimens. Ate/opus cruciger 1s a species

complex (JCS, FRR, and CBA, unpub. data).

Selected references: Lichtenstein and von Martens

(1856); Gunther (1858); Lutz (1927); Muller (1935);

Sexton (1958); Ginés (1959); Rohl (1959); Rivero (1961,

July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

Images. The task of gathering photos of all possible species of amphibians from Venezuela for a project like this began some time

ago. We conducted an exhaustive search, asking many collaborators for permission to use their photos. Many species are already

extinct or believed to be (such as all Atelopus except A. cruciger); for some other species the photos presented herein are the only

published so far (for example, Metaphryniscus sosae, Oreophrynella huberi, Hyalinobatrachium guairarepanensis, Pristimantis

marahuaka, Dendropsophus yaracuyanus, Scinax baumgardneri), and for many other species they are the first reproduced in color.

For some species a picture taken in Venezuela could not be found, so a decision was made to either show the same species from

other close countries, or show specimens from Venezuela and also from other countries for comparison.

Finally, we illustrate with preserved material some species that remain problematic or were never photographed alive (such as

Atelopus vogli, Dendropsophus battersbyi, Dischidodactylus colonnelloi, Pristimantis longicorpus, Stefania breweri, and Boana

sp. cf. rufitela).

1A. Allophryne ruthveni. Female, Cuyuni, Bolivar. Photo: 1B. Allophryne ruthveni. Male, Sierra de Imataca, Delta

César Barrio-Amoros. Amacuro. Photo: César Barrio-Amoros.

Pa. ie Gee ™ Ge

b 0 rae sg 0 er Gi kt eS

2. Amazophrynella minuta. Raudal de Danto, rio Cuao, 3. Atelopus carbonerensis. Amplexus, La Carbonera, Mérida,

Amazonas. Photo: Fernando Rojas-Runjaic. Photo: Amelia Diaz de Pascual.

4A. Atelopus cruciger. Male, Cuyagua, a Photo: Gece 4B. Aelopas cruciger. Male, Quebrada Chacaito, Miranda.

Barrio-Amoros. Photo: Charles Brewer-Carias.

Amphib. Reptile Conserv. 8 July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

1964a); Tello (1968); La Marca et al. (1989); Cocroft et

al. (1990); La Marca (1992, 1995a,c); Rodriguez and

Rojas-Suarez (1995, 1999); Barrio (1996b); Lotters

(1996); La Marca and Lotters (1997); Lotters et al.

(1998, 2004); Rivas (1998); Barrio-Amoroés (2001,

2006c, 2013); Lotters and La Marca (2001); Bonaccorso

et al. (2003); Manzanilla and La Marca (2004); Rueda-

Almonacid et al. (2005); Rodriguez-Contreras et al.

(2008); Barrio-Amoros and Torres (2010); Gonzalez et

al. (2010); Lampo et al. (2011).

Atelopus mucubajiensis Rivero, 1972*

Holotype: BM 1971.763.

Type locality: “Region de Mucubaji, Santo Domingo,

3,100 m, Estado Mérida, Venezuela.”

Distribution: Region 1. Surroundings of the type

locality, paramo and subparamo, 2,800—3,500 m asl.

Remarks: In September 2004, the last known individual

was delivered to CBA (Barrio-Amoroés 2004). Soon

afterwards a project was begun to search for the species

(Barrio-Amoros 2009a), but unfortunately this effort

yielded no positive results despite an intensive search

effort. The female individual died a few weeks after being

received and tested weakly positive for the presence of

the fungus Batrachochytrium dendrobatidis (Bd; Lampo

et al. 2007).

Selected references: Rivero (1972, 1980); La Marca et

al. (1989); La Marca (1994, 1992, 1995a,c); Rodriguez

and Rojas-Suarez (1995, 1999); Barrio (1996b); Durant

and Diaz (1996); Lotters (1996); La Marca and Lotters

(1997); Barrio-Amorés (2001, 2004, 2009a); Rueda-

Almonacid et al. (2005); Lampo et al. (2007); Barrio-

Amoros and Torres (2010).

Atelopus oxyrhynchus Boulenger, 1903*

Lectotype: BM 1947.2.14.66. designated by Rivero

(1972).

Type locality (of the lectotype): Rio Albarregas, La

Culata, Sierra Nevada de Mérida, Venezuela (elevation

3,330 m).

Distribution: Region 1. Sierra de la Culata, Mérida

State.

Remarks: Apparently extinct, and not seen alive since

1994 (Lotters and La Marca 1997).

Selected references: Boulenger (1903); Ginés (1959);

Rivero (1961, 1963b, 1964d, 1972); La Marca et al.

(1989); Durant (1993); Pifiero and Durant (1993); La

Marca (1994, 1992, 1995a,c); Rodriguez and Rojas-

Suarez (1995, 1999): Lotters (1996); La Marca and

Lotters (1997); Barrio-Amords (2001, 2013); Rueda-

Almonacid et al. (2005).

Atelopus pinangoi Rivero, 1980*

Type: UPRM 5354.

Amphib. Reptile Conserv.

Type locality: “Pifiango, 2,920 m, Estado Mérida,

Venezuela.”

Distribution: Region 1. Surroundings of the type

locality.

Remarks: Last seen in situ in 1997 (Barrio-Amoros

2013). Garcia-Pérez et al. (2013) reported finding a

postmetamorph at the type locality in December 2008,

but several searches (2010-2013) found no additional

individuals.

Selected references: Rivero (1980); La Marca et al.

(1989); Rodriguez and Rojas-Suarez (1995, 1999);

Lotters (1996); La Marca and Lotters (1997); Barrio-

Amoros (2001, 2013); Rueda-Almonacid et al. (2005);

Garcia-Pérez et al. (2013).

Atelopus sorianoi La Marca, 1983*

Holotype: CVULA IV-2783.

Type locality: “Bosque nublado 10 km SSE de Tovar,

2,718 m, Estado Mérida, Venezuela.”

Distribution: Region 1. Initially thought restricted to the

type locality, but photographs of a specimen shown to

T.R. Kahn by E. La Marca in 2007 indicated a population

exists outside type locality. The type locality was

destroyed by a landslide (pers. comm., T.R. Kahn from

E. La Marca). Further work is required to determine if

this disjunct population is extant.

Remarks: Not seen in the type locality since 1990;

possibly extinct at the type locality, but also possibly

extant at a disjunct valley not far from the type locality.

Selected references: La Marca (1983, 1992, 1995a,c);

La Marca et al. (1989); Rodriguez and Rojas-Suarez

(1995, 1999); Barrio (1996b); Lotters (1996); La Marca

and Lotters (1997); Barrio-Amoros (2001, 2013); Rueda-

Almonacid et al. (2005).

Atelopus tamaense La Marca, Pérez and Renjifo, 1989*

Holotype: ULABG 1820.

Type locality: “Estado Apure, Distrito Paez, Cercanias

del “Boqueron del rio Oira,” Paramo de Tama, Parque

Nacional El Tama, aproximadamente 7°25’N, 72°23’ W,

elevation 2,950 m.”

Distribution: Region 1. Only known from the type

locality, in Venezuela and probably ranging into nearby

Colombia in Tama National Park.

Remarks: Population status unknown, but not seen since

its description.

Selected references: La Marca et al. (1989); La Marca

(1992, 1995c); Rodriguez and Rojas-Suarez (1995,

1999); Lotters (1996); La Marca and Lotters (1997),

Barrio-Amoros (2001, 2013); Rueda-Almonacid et al.

(2005).

Atelopus vogli Miller, 1934*

Holotype: ZSM 3/1933.

July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

5. Atelopus mucubajiensis. Female, Santo Domingo, Mérida.

Photo: César Barrio-Amoros.

7, Atelopus tamaense. Boqueron del rio Oira, paramo de Tama,

Apure. Photo: Juan M. Renjifo.

fh ak

9, Metaphryniscus sosae. Marahuaka sur Summit, Amazonas.

Photo: Charles Brewer-Carias.

Type locality: “Cascadas superiores del Rio Gtiey, en

la region llamada Las Pefias, cerca de la Hacienda de la

Trinidad, Maracay, 700 m,” Aragua State, Venezuela.

Distribution: Region 2. Known from two localities, the

type locality and a record reported by Barrio-Amoros and

Rojas-Runjaic (2009) in Montalban (Carabobo State);

last specimen known was collected in 1957.

Amphib. Reptile Conserv.

: : t oe Ce

H \ | i \ 4 1 ; L

! |

Garabobe Photo:

8. Atelopus vogli. MCNC 72, Montalban,

César Barrio-Amoros.

10. Oreophrynella cryptica. Summit of Auyan-tepui, Bolivar.

Photo: Josefa Celsa Sefiaris.

Remarks: Lotters et al. (2004) elevated Atelopus cruciger

vogii to a full species. Currently considered the only Venezuelan

amphibian verified as extinct (Rodriguez and Rojas-Suarez.

2008; La Marca and Sefiaris 2015).

Selected references: Muller (1935); Lotters et al.

(2004); Rueda-Almonacid et al. (2005); Rodriguez and

Rojas-Suarez (2008); Barrio-Amoros and Rojas-Runjaic

10 July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

(2009); Barrio-Amoros (2013); La Marca and Sefiaris

(2015).

Genus Metaphryniscus

Sefiaris, Ayarzagiiena and Gorzula, 1994

Type species: Metaphryniscus sosai Sefiaris et al., 1994,

by original designation.

Metaphryniscus sosae

Sefiaris, Ayarzagtiena and Gorzula, 1994*

Holotype: MHNLS 12347.

Type locality: “Tepuy Marahuaca-Sur, Estado Amazonas,

Venezuela (3°40’N, 65°27’ W). 2,600 m snm.”

Distribution: Region 5. Endemic to Marahuaka-tepui in

Amazonas State.

Remarks: Since its discovery and subsequent description,

no additional specimens of this genus and species have

been collected. Barrio-Amoros (1998) emended the name

sosai to sosae (as the species was dedicated to a woman),

without explanation. Naming here follows Article 31.1.2

of the Code in retaining the emendation of sosae, the -ae

ending.

Selected references: Sefiaris et al. (1994).

Genus Oreophrynella Boulenger, 1895

Type species: Oreophryne quelchii Boulenger, 1895, by

monotypy.

Oreophrynella cryptica Sefiaris, 1995 *

Holotype: EBRG 2956.

Type locality: Eastern sector of the Auyan-tepui summit,

Bolivar State, Venezuela (5°53’36’N, 62°29°12”W),

1,750 m.

Distribution: Region 5. Endemic to Auyan-tepui in

Bolivar State.

Remarks: A very rare species described from only two

specimens (Sefiaris 1995). Despite numerous expeditions

to the type locality, only five additional specimens have

been collected (in 1974) and deposited in the BMNH

(Myers and Donnelly 2008).

Selected references: Sefiaris (1995); Myers (1997);

Myers and Donnelly (2008); Sefiaris et al. (2014).

Oreophrynella huberi

Diego-Aransay and Gorzula, 1987*

Holotype: MHNLS 11148.

Type locality: Cerro El Sol, northeast of Auyan-tepui

(6°6’N, 62°32’W), Bolivar State, Venezuela, elevation

1,700 m.

Distribution: Region 5. Endemic to Cerro El Sol-tepui

in Bolivar State.

Selected references: Diego-Aransay and Gorzula

Amphib. Reptile Conserv.

ait

(1985); Sefiaris et al. (1994, 2014); Gorzula and Sefiaris

(1998).

Oreophrynella macconnelli (Boulenger 1895)

Holotype: BM 1947.2.14.49.

Type locality: “Base of Mount Roraima, 3,500 feet

(1,066 m),” Guyana.

Distribution: Region 5. Known from the base of Roraima

(type locality) and Maringma-tepui in Guyana (Kok

2009). It has been surmised to occur in the Venezuelan

foothills surrounding the base of Mt. Roraima (Rivero

1961; La Marca 1992: Barrio-Amoros 1998). To date

there are no vouchers from Venezuela in any known

scientific collection.

Remarks: It is not possible to designate the exact type

locality or determine whether it is in Venezuela or

Guyana, since Boulenger (1900) listed it as “the base of

Mt. Roraima, 3,500 ft (ca. 1,066 m),” which could be

in either Guyana, Venezuela or Brazil. Barrio-Amoros

(1998) considered Boulenger’s type locality to be in

Venezuela without explanation. He also incorrectly

stated that the type material in the BMNH was lost. Kok

(2009) assumed the type specimen was collected on the

Guyanan side of Roraima, as the title of Boulenger’s

article states, but provided no further information or

evidence. The journey made by F.V. McConnell and

J.J. Quelch in 1898, during which the specimen was

collected, originated in Guyana, but the only access to

the summit of Roraima by foot is on the Venezuelan

side. The exact route by which the two collectors walked

around the base of Roraima follows the Mazaruni-Cako-

Aruparu Rivers to reach the foot of Roraima. Here, only

one place at the Aruparu River seems to be at 1,066 m

(5°15°10.93”N, 60°42’28.57°W), while still seeing

the tepui from its base. Therefore, we restrict the type

locality to be in Guyana. But see a different conclusion

by Kok et al. (2018) for Pristimantis marmoratus.

We are unable to distinguish Oreophrynella

macconnelli from O. dendronastes Lathrop and

MacCulloch, 2007. The only listed difference is the shape

of the snout, reported as truncated for O. dendronastes

and pointed or acuminate for O. macconnelli (Lathrop

and MacCulloch 2007). Kok (2009) had examined a

larger set of specimens of O. macconnelli and noted

variation in this character, but snout shape alone may be

insufficient evidence to distinguish between these taxa.

The larger size of O. dendronastes, with a female of 37.3

mm SVL (as opposed to an SVL of 22.7 mm in males

of O. macconnelli, for which females are unknown)

seems significant, but could be attributed to maturity

and/or dimorphism. Furthermore, Kok et al. (2012)

found insufficient genetic differentiation to support the

recognition of O. dendronastes. We therefore suggest

considering O. dendronastes Lathrop and MacCulloch,

2007 as a junior synonym of O. macconnelli (Boulenger

1895). Barrio-Amoros (1998, 2004, 2009) incorrectly

July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

11. Oreophrynella huberi. Cerro el Sol summit, Bolivar. Photo:

Anonymous.

13. Oreophrynella nigra. Yuruani Summit, Bolivar. Photo:

César Barrio-Amoros.

! fs

4 = Be

‘-,” 4 _

14B. Oreophrynella quelchii. Summit of Mount Roraima,

Bolivar. Photo: Roger Manrique.

Amphib. Reptile Conserv.

12. Oreophrynella macconnelli. Maringma tepui, Guyana.

Photo: Philippe Kok.

= i a

14A. Oreophrynella quelchii. Summit of Mount Roraima,

Bolivar. Photo: Roger Manrique.

15. Oreophrynella vasquezi. Summit of Ilu-tepui, Bolivar.

Photo: Javier Mesa.

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

gave the species name as O. macconelli (lapsus calami).

Selected references: Boulenger (1900); Ginés (1959);

Rivero (1961, 1964b); McDiarmid (1971); Warren

(1971); Sefiaris et al. (1994, 2014); McDiarmid and

Donnelly (2005); Lathrop and MacCulloch (2007); Kok

(2009); Kok et al. (2018).

Oreophrynella nigra

Sefiaris, Ayarzagiiena and Gorzula, 1994

Holotype: MHNLS 10583.

Type locality: Kukenan-tepui I, Bolivar State, Venezuela

(9°51’N, 60°48’ W), 2,500 m asl.

Distribution: Region 5. Endemic to Kukenan and

Yuruani Tepuis in Bolivar State.

Remarks: Kok et al. (2012) implied that O. nigra could

be a synonym of O. quelchii due to the low genetic

divergence between these forms. However, inhabiting

different tepuis means reproductive isolation (allopatry),

and having a different color pattern could be sufficient

reason at the moment to maintain it as a valid species.

Selected references: McDiarmid and Gorzula (1989);

Solano (1989); Sefiaris et al. (1994, 2014); Gorzula and

Sefiaris (1998); McDiarmid and Donnelly (2005).

Oreophrynella quelchii (Boulenger 1895)

Syntypes: BM 95.4.19.1—5; 99.3.25.7-13; UK 126081-

82; ZFMK, MCZ 3500-02.

Type locality: “Summit of Mt. Roraima, between British

Guiana and Venezuela, at an altitude (elevation) of 8,500

feck:

Distribution: Region 5. Known from the summit of

Mt. Roraima, a tepui extending across three countries:

Venezuela, Guyana, and Brazil.

Selected references: Boulenger (1895a,b; 1900); Ginés

(1959); Rivero (1961, 1964b); McDiarmid (1971);

Hoogmoed (1979b); McDiarmid and Gorzula (1989): La

Marca (1992); Sefiaris et al. (1994, 2014); Gorzula and

Sefiaris (1998); McDiarmid and Donnelly (2005).

Oreophrynella vasquezi

Sefiaris, Ayarzagtiena and Gorzula, 1994*

Holotype: MHNLS 10244.

Type locality: [lu-tepui I, Bolivar State, Venezuela

(5°25’N, 60°58’ W); elevation 2,650 m.

Distribution: Region 5. Endemic to Iu, a tepui in

Bolivar State.

Selected references: Sefiaris et al. (1994, 2014); Gorzula

and Sefiaris (1998).

Genus Rhaebo Cope, 1862

Type species: Bufo haematiticus Cope,

monotypy.

1862, by

Amphib. Reptile Conserv.

Rhaebo glaberrimus (Gunther 1869)

Holotype: BM 1947.2.20.56. (formerly 68.3.4.9).

Type locality: “Bogota” (Bogota), Cundinamarca,

Colombia (in error: Mueses-Cisneros et al. (2012) regard

Bogota as the origin of shipment, not the locality where

the type specimen was collected).

Distribution: Regions 1, 4. Amazonian piedmont of

the Cordillera Oriental in Colombia, and southeastern

versant of the Venezuelan Andes. In Venezuela, known

from a single locality in the Doradas River Basin, Tachira

State (Chacon et al. 2000).

Selected references: Chacon et al. (2000, 2001);

Mueses-Cisneros et al. (2012).

Rhaebo guttatus (Schneider 1799)

Holotype: ZMB 3517.

Type locality: “India Orientali.” In error; Rivero (1961)

clarifies that the label of the holotype states the type

specimen was collected in Suriname.

Distribution: Regions 4, 5. Typical Guyano-Amazonian

element, present in Bolivia, Peru, Ecuador, Colombia,

Venezuela, Suriname, French Guiana, Guyana, and

Brazil. In Venezuela, widespread in Amazonas and

Bolivar States, also occurs in one disjunct area of Apure

State (Barrio et al. 2001).

Remarks: Bufo anderssoni Melin, 1941 was shown to

be a junior synonym of Rhaebo guttatus (Barrio-Amoros

and Castroviejo-Fisher 2008a).

Selected references: Ginés (1959): Rivero (1961, 1964b,

1967a); Hoogmoed and Gorzula (1979); Rivero et al.

(1986); La Marca (1992); Duellman (1997); Gorzula

and Sefiaris (1998); Barrio et al. (2001, 2011b); Barrio-

Amoros and Brewer-Carias (2008); Barrio-Amoros

and Castroviejo-Fisher (2008a); Barrio-Amordés and

Duellman (2009); Sefiaris et al. (2014).

Rhaebo haematiticus Cope, 1862

Syntypes: USNM 4844849.

Type locality: “Region of the Truando (Choco), New

Grenada (Colombia).”

Distribution: Region 1. From eastern Honduras into

southern Costa Rica throughout the western slopes of the

Cordillera Occidental and eastern versant of Cordillera

Central of Colombia, also occuring in the Chocoan

Region of northwestern Ecuador and northwestern

Venezuela. In Venezuela, known from scattered localities

within the Sierra de Perija, the northernmost point of

the Andes on the border with Colombia and Venezuela

(Barrio-Amoros 2001; Rojas-Runjaic et al. 2007; Vieira-

Fernandes et al. 2016). Another single record is from

Cordillera de Mérida (Vieira-Fernandes et al. 2016).

Apparently, R. haematiticus is likely a complex of cryptic

species (Mueses-Cisneros 2009). Venezuelan populations

could be represented by an undescribed species (Mueses-

July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

16. Rhaebo Baber Rio Frio, Tachira. Photo: César

Barrio-Amoros.

17B. Rhaebo guttatus. Santa Elena de Uairén, Bolivar. Photo:

César Barrio-Amoros.

ISB. Rhaebo haematiticus. Female. Ipika, Takeo river basin,

Serrania de Perija, Zulia. Photo: Fernando Rojas-Runjaic.

Cisneros 2009; Vieira-Fernandes et al. 2016).

Selected references: Barrio-Amorés (2001); Rojas-

Runjaic et al. (2007); Mueses-Cisneros (2009).

Rhaebo nasicus (Werner 1903)

Holotype: MRHN I.G. 9422 reg. 1015.

Amphib. Reptile Conserv.

17A. Rhaebo guttatus. pera Las Lajas, Siena de ena:

Bolivar. Photo: César Barrio-Amoros.

18A. Rhaebo haematiticus. Subadult. Ipika, Tukuko river

basin, Serrania de Perija, Zulia. Photo: Edwin Infante-Rivero.

19A. Rhaebo nasicus. Subadult. Kavanayen, Gran Sabaja,

Bolivar state. Photo: Edward Camargo.

Type locality: Type without locality data, restricted to

“South America, probably along the Atlantic drainage”

by Smith and Laurent (1950).

Distribution: Region 5. Known from eastern Venezuela

and northwestern Guyana. In Venezuela it was previously

known only from the La Escalera Region of Bolivar State

(Barrio-Amoroés 2009). Occurrence reported here from

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Barrio-Amoros et al.

7.1kmN Kavanayen (EBRG 5581, 5583, 5588, 5763),

and from the Churun River, near the base of Angel Falls,

on the northern side of Auyan-tepui (MHNLS 20429).

Remarks: See remarks under Rhaebo ceratophrys.

Barrio-Amoros (2009) and Barrio-Amoros and Duellman

(2009) were reluctant to recognize nasicus as a member

of Rhaebo, but thought it should be placed in Rhinella

based on external characteristics. Pramuk (2006) showed

the relationship of nasicus as being the most basal in the

Bufo guttatus species group (now Rhaebo).

Selected references: Hoogmoed (1977, 1979b, 1986,

1990a); Duellman (1997); Pramuk (2006); Kok and

Kalamandeen (2008); Barrio-Amoroés (2009); Barrio-

Amoros and Duellman (2009); MacCulloch and Lathrop

(2009); Sefiaris et al. (2014).

Genus Rhinella Fitzinger, 1826

Type species: Bufo proboscideus Spix,

monotypy.

1824, by

Rhinella beebei (Gallardo, 1965)

Holotype: AMNH 557774.

Type locality: Churchill-Roosevelt highway, Trinidad,

Trinidad and Tobago, 30 m.

Distribution: Regions 3, 5, 6. Widespread in open

areas throughout northern Venezuela and Trinidad.

In Venezuela species is found in lowland open areas,

north and south of the Orinoco River, and has also been

documented on Margarita Island (Ugueto and Rivas

2010).

Remarks: Narvaes and Rodrigues (2009) changed the

previous concept of the R. granulosa species group,

recognizing available subspecific names as full species

levels, and invalidating others. Bufo granulosus

barbouri from Isla Margarita and B. g. beebei from the

Orinoco Basin and northern Venezuela were considered

synonyms of R. humboldti. Pereyra et al. (2016) showed

two parapatric populations (one on lowlands around

Puerto Ayacucho, another in uplands of Gran Sabana)

as monophyletic under the name R. humboldti. Murphy

et al. (2017) revised the name beebei for populations in

Trinidad and northern Venezuela, noting differences in

genetics and vocalizations among R. humboldti (a trans-

Andean species) and R. beebei. Notably, no specimens

from the Lake Maracaibo Basin were analyzed. Further

work is required to determine if that population

represents R. humboldti, R. beebei, or R. centralis, and/or

a combination of these species living in sympatry.

Selected references: Some of these references are

based on populations of R. beebei under the names Bufo

granulosus or Rhinella humboldti. Boettger (1892); Lutz

(1927); Parker (1936); Ginés (1959); Rivero (1961,

1964a-c, 1967c); Roze (1964); Gallardo (1965); Tello

(1968); Cochran and Goin (1970); Staton and Dixon

(1977); Duellman (1979a, 1997); Hoogmoed (1979b,

Amphib. Reptile Conserv.

1990a); Hoogmoed and Gorzula (1979); Péfaur and Diaz

De Pascual (1982, 1987); Rivero et al. (1986); Ramo and

Busto (1989, 1990); La Marca (1992); Manzanilla et al.

(1995); Péfaur and Pérez (1995); Yustiz (1996); Murphy

(1997); Gorzula and Sefiaris (1998); Rivas and Barrio-

Amoros (2005); Narvaes and Rodrigues (2009): Tarano

(2010); Ugueto and Rivas (2010); Barrio-Amoros et al.

(2011b); Pereyra et al. (2016).

Rhinella ceratophrys (Boulenger 1882)

Holotype: BM 80.12.5.151.

Type locality: “Ecuador.”

Distribution: Region 5. Widely distributed in upper

Amazon Basin, from northeastern Peru, through eastern

Ecuador and southeastern Colombia, into southern

Venezuela. Its occurrence in Brazil was presumed

(Fenolio et al. 2012) and verified by Rojas-Runjaic et

al. (2017). These last authors reported new localities for

this species with specimens from Venezuela at Cerro

Marahuaca and at Cerro de la Neblina, Amazonas State.

Elevations 1,350—2,713 m asl.

Remarks: Miyares-Urrutia and Arends (2001) confused

a female specimen of Rhaebo nasicus from La Escalera

with Rhinella ceratophrys (CBA, personal observation).

Barrio-Amoros (1998) doubted the identity of Rivero’s

record (based on a single juvenile of 12 mm SVL).

However, Fenolio et al. (2012) recently verified the

Venezuelan specimen and confirmed its identity using

morphological and morphometric data. Recent reports

of this species from up- and highland western tepuis

solves its distribution in Venezuela. Further sampling and

analyses will determine if a species complex exists and

may further be elucidated (Rojas-Runyaic et al. 2017).

Fouquet et al. (2007a) removed this species from the

Rhinella margaritifera species group.

Selected references: Rivero (1961, 1964b); Cochran

and Goin (1970); Miyares-Urrutia and Arends (2001);

Fenolio et al. (2012); Rojas-Runjaic et al. (2017).

Rhinella horribilis (Wiegmann 1833)

Syntypes: ZMB 3479 (Misantla), ZMB 3480 (no locality

given), ZMB 3481 (Veracruz), ZMB 3493 (Mexico),

all from Mexico, based on the original description.

Lectotype: ZMB 3480, assigned by Fouquette and

Dubois (2014).

Type locality: “In der Umgegend von Vera Cruze,”

Mexico.

Distribution: Regions 1, 6. In Venezuela west of the

Andes, including the dry valley of Chama River (Mérida

State), up to 2,000 m asl (Barrio-Amoros 1998; Acevedo

et al. 2016); northern Colombia and Panama north

throughout Central America and into the southern United

States (Frost 2018).

Remarks: This taxon was recently removed from

synonymy with Rhinella marina by Acevedo et al. (2016).

July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

pe,

19B. Rhaebo nasicus. Female. Sierra de Lema, Bolivar. Photo: 20A. Rhinella beebei. Puerto Ayacucho, Amazonas. Photo:

Alan Highton. César Barrio-Amoros.

‘tons

Ee , ; ee

= eM

20B. Rhinella beebei. Barinas, Barinas. Photo: Fernando 20C. Rhinella beebei. La Guaquira, Yaracuy. Photo: Fernando

Rojas-Runjaic. Rojas-Runjaic.

‘

apie - : i

ella cf. beebei. Northern pie

20D. Rhin

Zulia. Photo: Fernando Rojas-Runjaic.

21A. Rhinella ceratophrys. Loreto, Peru. Photo: Danté Fenolio.

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Barrio-Amoros et al.

Populations west of the Andes formerly referred to as R.

marina are genetically and osteologically distinct from

populations east of the Andes. The form ranging from

western Venezuela through northwestern South America

north into Mexico and the southern United States should

be considered as R. horribilis.

Selected references: Mostly under the names Rhinella

marina or Bufo marinus. Aleman (1952), Péfaur and

Pérez (1995); Acevedo et al. (2016).

Rhinella margaritifera (Laurenti 1768)

Neotype: MNRJ 71538, assigned by Lavilla et al. (2013).

Type locality: “Brasilia.” The neotype is from

Municipality Humaita, Amazonas State, Brazil (Lavilla

et al. 2013).

Distribution: Regions 4, 5. Widespread in the Amazonian

Region. In Venezuela this species is present in rainforests

and cloud forests of the Cordillera de la Costa, the Andes,

and south of the Orinoco. It is absent from the Llanos

Region.

Remarks: Rhinella margaritifera represents a species

complex likely including several undescribed and/or

misidentified taxa. At least two distinct taxa are present

on the eastern versant of the Cordillera de Mérida. One

from Falcon State was removed from this complex

and redescribed as Rhinella sclerocephala by Mijares-

Urrutia and Arends (2001; see account). Rivero (1964)

considered specimens from Guatopo and Falcon as Bufo

typhonius alatus. La Marca (1997) followed Rivero’s

example, but the decision to use a/atus as the name for

populations in the complex north of the Orinoco River

was premature and speculative. Santos et al. (2015)

restricted the distribution of R. alata to Panama, Western

Colombia, and Ecuador. La Marca (1992, 1997) included

R. acutirostris in the amphibian fauna of Venezuela, but

no populations of this species have been documented

based on voucher specimens. In Venezuela south of the

Orinoco River, at least four distinct populations of the

Rhinella margaritifera species complex occur, one in

western Amazonas State (CBA, unpub. data), one in

the Sarisarifiama foothills (Barrio-Amoros and Brewer-

Carias 2008), one in El Pauji, southern Gran Sabana,

where females lack prominent crests, and one along the

northern versant of the Serrania de Lema (Barrio-Amoros

et al. 2011b). Pending formal identification, this last

population could correspond to Rhinella martyi (Fouquet

et al. 2007), a taxon so far not reported from Venezuela.

Selected references: Spix (1824); Ginés (1959); Rivero

(1961, 1964a,b,d, 1967a, 1971a); Yustiz (1976a, 1996);

Hoogmoed (1977, 1986, 1989, 1990); Péfaur and Diaz

De Pascual (1982, 1987); Hoogmoed and Gruber (1983);

Rivero et al. (1986); Hass et al. (1995); Velez (1995),

Gorzula and Sefiaris (1998); Rivas and Barrio-Amoros

(2005); Barrio-Amords and Brewer-Carias (2008);

Barrio-Amoros (2010a); Barrio-Amoros et al. (2011b);

Sefiaris et al. (2014); Santos et al. (2015).

Amphib. Reptile Conserv.

Rhinella marina (Linnaeus 1758)

Holotype: Not identified (originally in the collection of

Albertus Seba).

Type locality: “America.” Restricted to Suriname by

Muller and Hellmich (1936).

Distribution: Regions 3, 4, 5. After the partition of

Rhinella marina into two species (Acevedo et al. 2016),

the name marina 1s now applied to populations occurring

east of the Andes in Venezuela and Colombia, south to

central Brazil to Bolivia. This species has been introduced

onto several islands and into countries in the Caribbean,

Pacific, and Indian Oceans (Zug and Zug 1979), although

distinction between R. marina and R. horribilis invasive

populations has yet to be determined.

Remarks: All populations sampled west of the Andes

are currently recognized as Rhinella horribilis (Acevedo

et al. 2016). These authors, however, did not include in

their analysis samples of R. marina from the majority of

Venezuela. The current authors have observed important

morphological and acoustic differences in populations

east of the Andes, at least in Venezuela. A wider sample

should be analyzed to properly assess the complex

taxonomy and nomenclature of these species and/or

species complex.

Selected references: Lichtenstein and von Martens

(1856); Lutz (1927); Ginés (1959); Rohl (1959); Rivero

(1961, 1964a—d, 1967a); Heatwole et al. (1965); Tello

(1968); Staton and Dixon (1977); Hoogmoed and

Gorzula (1979); Rivero-Blanco and Dixon (1979); Zug

and Zug (1979); Péfaur and Diaz De Pascual (1982,

1987); Hoogmoed and Gruber (1983); Gremone et

al. (1986); Rivero et al. (1986); Hoogmoed (1989);

La Marca (1992); Manzanilla et al. (1995); Evans and

Lampo (1996); Duellman (1997); Gorzula and Sefiaris

(1998); Barrio-Amords and Brewer-Carias (2008);

Barrio-Amoros (2010a); Ugueto and Rivas-Fuenmayor

(2010); Barrio-Amoros et al. (2011b); Sefiaris et al.

(2014); Acevedo et al. (2016).

Rhinella merianae (Gallardo 1965)

Holotype: AMNH 46531.

Type locality: “Head falls of Essequebo River, British

Guiana (= Guyana).”

Distribution: Regions 5, 6. Brazil, Suriname, Guyana,

French Guiana, and Venezuela. In Venezuela, known

from the Orinoco River, Gran Sabana (Bolivar State), and

a disjunct record in Zulia State (Narvaes and Rodrigues

2009). Obviously, the last record is likely in error. It may

be R. beebei or another species within the granulosus

species group.

Remarks: A member of the Rhinella granulosa species

group. Even though Narvaes and Rodrigues (2009) stated

examining voucher specimens, the biogeography is not

evidential to the current authors. There are specimens

from Puerto Ayacucho, Gran Sabana, the Orinoco River,

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Catalogue of the amphibians of Venezuela

i em aie Sor i Pe ae

21B. Rhinella ceratophrys. Parque Nacional Pico da Neblina, 22. Rhinella horribilis. La Mucuy Baja, Mérida. Photo: César

Brazil. Photo: Vinicius T: de Carvalho. Barrio-Amoros.

& - 7 a

_ ira ~ a Aaa = ee pe sie =e wh a fra * aS =" F "oy Se ee

23A. Rhinella margaritifera. Male. Rio Autana, Amazonas. 23B. Rhinella margaritifera. Female. Rio Doradas, Tachira.

Photo: Javier Mesa. Photo: César Barrio-Amoros.

z es 2 _ : ; * " : i |

Pauji, Bolivar. Photo: 23D. Rhinella margaritifera. Male. Los Alcaravanes, Calderas,

César Barrio-Amoros. Barinas state. Photo: César Barrio-Amoros.

23C. Rhinella margaritifera. Female. El

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Barrio-Amoros et al.

and Zulia State. The authors consider particularly the

Zulia State specimen doubtful, and believe it is more

likely to be R. beebei or R. centralis (the last would be a

first record for Venezuela). However, if the distribution is

valid, then that of R. merianae would overlap with that

of R. beebei along the Orinoco River and with that of R.

nattereri in the Gran Sabana Region.

Selected references: Narvaes and Rodrigues (2009);

Sefiaris et al. (2014); Pereyra et al. (2016).

Rhinella nattereri (Bokermann 1967)

Holotype: MZUSP 73715.

Type locality: Cachoeira Uranduique, Mau _ River,

Roraima, Brazil.

Distribution: Region 5. Brazil, Guyana, and Venezuela.

In Venezuela this species is known only from the Gran

Sabana Region in Bolivar State (Narvaes and Rodrigues

2009).

Remarks: A member of the Rhinella granulosa species

group; little is known about this species in Venezuela.

Pereyra et al. (2016) could not obtain genetic samples

to test its phylogenetic relationships. The morphological

characteristics of this form should be more carefully

compared with R. beebei and R. merianae to determine

and support species-level validity.

Selected references: Narvaes and Rodrigues (2009);

Sefiaris et al. (2014); Pereyra et al. (2016).

Rhinella sclerocephala

(Mijares-Urrutia and Arends 2001)*

Holotype: EBRG 3415.

Type locality: “1.5 km (by road) from Curimagua to

Cerro Galicia, municipio Petit, Sierra de San Luis,

Estado Falcon, Venezuela (about 11°10’N, 69°41’W),

about 1,150 m.”

Distribution: Region 2. Apparently endemic to the cloud

forests at Sierra de San Luis, Falcon State. No additional

specimens of this species have been collected in the

last 25 or so years, according to material preserved in

Venezuelan museums. Other than the series by Miyares-

Urrutia and Arends (2001) no other specimens are known

in the scientific collections examined.

Selected references: Mijares-Urrutia and Arends (2001).

Rhinella sternosignata (Gunther 1858)

Syntypes: BM — 1947.2.21.68-69; = 1947.2.21.70;

1947.2.21.87; 1947.2.21.88.

Type locality: “Venezuela;”’ “Puerto Cabello;”

“Cordova,” “México;” restricted to Puerto Cabello by

Cochran and Goin (1970).

Distribution: Regions 1, 2. The eastern versant of

the Cordillera Oriental of Colombia and Venezuela.

In Venezuela the species is distributed in cloud and

rainforests of the Cordillera de la Costa and in the eastern

Amphib. Reptile Conserv.

Andean versant, or so-called Andean Piedmont.

Remarks: Pereyra et al. (2016) found Rhinella

sternosignata closely related to the R. margaritifera and

R. veraguensis species groups.

Selected references: Boulenger (1882); Boettger (1892);

Lutz (1927); Ginés (1959); Rivero (1961, 1964a,d);

Cochran and Goin (1970); Hoogmoed (1990); La Marca

(1992); Manzanilla et al. (1995); La Marca and Mijares

(1996); Yustiz (1996); La Marca and Manzanilla (1997),

Vélez (1999); Pereyra et al. (2016).

Family Centrolenidae Jiménez de la Espada, 1872

Remarks: We follow the taxonomic arrangement

proposed by Guayasamin et al. (2009).

Subfamily Centroleninae Taylor, 1951

Genus Centrolene Jiménez de la Espada, 1872

Type species: Centrolene geckoideum Jiménez de la

Espada, 1872, by monotypy.

Remarks: The generic name Centrolene is feminine,

not neuter as Myers and Donnelly (1997) interpreted.

Jimenez de la Espada (1872) described Centrolene

without indicating a gender and associated the name

with a neuter species name (geckoideum) in error.

Following the Code it must be corrected (M.A. Alonso

de Zarazaga, pers. comm.), as Centrolene is feminine

from the Greek nouns (kéntron -sting, spur- and oléné

-elbow-, thus feminine in Greek), so, the emendations

proposed by Myers and Donnelly (1997) require revision.

Following Article 30.1.2 of the Code, the species names

are amended to feminine when determined to require

correction according to the Code (ICZN 1999). The

amended Centrolene follows, as feminine: Centrolene

altitudinalis (Rivero 1968), Centrolene antioquensis

(Noble 1920), Centrolene bacata Wild, 1994, Centrolene

daidalea (Ruiz-Carranza and Lynch 1991), Centrolene

geckoidea Jiménez de la Espada, 1872, Centrolene

gemmata (Flores 1985), Centrolene hesperia (Cadle and

McDiarmid 1990), Centrolene huilensis Ruiz-Carranza

and Lynch, 1995, Centrolene lemniscata Duellman and

Schulte, 1993, Centrolene notosticta Ruiz-Carranza and

Lynch, 1991, Centrolene peristicta (Lynch and Duellman

1973), Centrolene pipilata (Lynch and Duellman 1973),

and Centrolene venezuelensis (Rivero 1968). The species

not mentioned remain as previously published.

Centrolene altitudinalis (Rivero 1968)*

Holotype: MCZ 72500.

Type locality: “Quebrada cerca de Rio Albarregas, 2,400

m. Estado Mérida, Venezuela.”

Distribution: Region 1. Andes of Mérida State. Known

from five localities, including (1) the type locality; (2)

Monte Zerpa, N of the city of Mérida; (3) La Joya, NE of

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Catalogue of the amphibians of Venezuela

23E. Rhinella margaritifera. Male. Triunfo, Bolivar. This population

could represent R. martyi. Photo: César Barrio-Amoros.

23G. Rhinella margaritifera. Female. Kanarakuni, base of

Sarisarifiama tepui, Bolivar. Photo: César Barrio-Amoros.

fe “al

24B. Rhinella marina. Female. Santa Maria de Erebato, Caura

river basin, Bolivar. Photo: César Barrio-Amoros.

Mérida; (4) Altos de San Luis and El Chorotal, towards

Merida to La Azulita; and (5) Altos de San Luis.

Selected references: Rivero (1968b); Sefiaris and

Ayarzagtiena (2005).

Centrolene daidalea (Ruiz-Carranza and Lynch 1991)

Holotype: ICN 18008.

Amphib. Reptile Conserv. 20

ments aha ee

23F. Rhinella margaritifera. Female. Yacambu, Lara state.

Photo: César Barrio-Amoros.

‘ a

Phe be ai =.

24A. Rhinella marina. Female. San Vicente, Apure. Photo:

César Barrio-Amoros.

Bolivar. Photo:

César Barrio-Amoros.

Type locality: Granja Infantil del Padre Luna, Alban

Municipality, Cundinamarca, Departmento, western

slopes of the Cordillera Oriental, Colombia (4°45’56’N,

74°26’, 2,060 m).

Distribution: Region 1. Colombia and Venezuela. In

Venezuela, known from eight localities on the eastern

versant of the Sierra de Perija, 800—1,832 m asl.

Selected references: Rojas-Runjaic et al. (2010).

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

Centrolene notosticta Ruiz-Carranza and Lynch, 1991

Holotype: ICN 12632.

Type locality: “Departamento de Santander, vertiente

occidental de la Cordillera Oriental, municipio de Charala,

Virolin (= Inspeccién de Policia de Cafiaverales), sitio

“El Encino” ca. cabeceras Rio Luisito, Latitud 6°13'N,

73°05'W de Greenwich, 1,750 m,” Colombia.

Distribution: Region 1. Colombia and Venezuela. In

Venezuela, known from only one locality (Campamento

Guacharaca, 1,660 m asl) in the eastern versant of the

Sierra de Perija, Zulia State (Rojas-Runjaic et al. 2012).

Selected references: Rojas-Runjaic et al. (2012).

Centrolene venezuelensis (Rivero 1968) *

Holotype: MCZ 77503.

Type locality: “Valle de la Culata, bosque a 2,700 m,

Estado Mérida, Venezuela.”

Distribution: Region 1. Occurrs throughout the

Venezuelan Andes, from Tachira to Mérida States, and

in the Sierra de Periyja (Zulia State), 2,100—3,050 m

asl. Centrolene venezuelensis is speculated to occur in

Colombia, yet has not been documented.

Remarks: Myers and Donnelly (1997) elevated

Centrolene buckleyi venezuelensis to species level, as

C. venezuelense, without explanation. This decision

was supported by Sefiaris and Ayarzagtiena (2005).

Guayasamin et al. (2009) validated this species elevation.

Selected references: Ginés (1959); Rivero (1961, 1963b,

1964a, 1968b); Péfaur and Diaz De Pascual (1982);

Gremone et al. (1986); La Marca (1991b “1994,” 1996c);

Ruiz-Carranza and Lynch (1991); Ayarzagtiena (1992);

Myers and Donnelly (1997); Sefiaris and Ayarzagtiena

(2005); Rojas-Runjaic et al. (2012).

Genus Cochranella Taylor, 1951

Type species: Centrolenella granulosa Taylor, 1949, by

original designation.

Remarks: The species riveroi and duidaeana were not

evaluated by Guayasamin et al. (2009) in their phylogeny

of Centrolenidae. They are left as taxa incertae sedis

within Cochranella sensu lato, until more data becomes

available.

Cochranella duidaeana (Ayarzagitiena 1992)*

Holotype: MHNLS 12000.

Type locality: Region 5. “Cumbre sur del Monte Duida.

Territorio Federal Amazonas. Venezuela. (3°19°N,

65°38’ W). 2,140 m snm.”

Distribution: Endemic to Cerro Duida, a tepui in

Amazonas State.

Selected references: Ayarzagtiena (1992); Ayarzagtiena

and Sefiaris (1996); Sefiaris and Ayarzagtiena (2005);

Amphib. Reptile Conserv.

Guayasamin et al. (2009).

Cochranella riveroi (Ayarzagiiena 1992)*

Holotype: MBUCV 6190.

Type locality: Cumbre Cerro Aracamuni, Amazonas

State, 1,600 m asl.

Distribution: Region 5. Endemic to Cerro Aracamuni, a

granitic mountain in Amazonas State.

Selected references: Ayarzagtiena (1992); Ayarzagtiena

and Sefiaris (1996); Sefiaris and Ayarzagtiena (2005);

Guayasamin et al. (2009).

Genus Espadarana Guayasamin, Castroviejo-Fisher,

Trueb, Ayarzagtiena, Rada and Vila, 2009

Type species: Centrolenella andina Rivero, 1968, by

original designation.

Espadarana andina (Rivero 1968)

Holotype: MCZ 72502.

Type locality: “La Azulita, 1,050 m, Estado Mérida,

Venezuela.”

Distribution: Region 1. Colombia and Venezuela.

Widely distributed throughout the Cordillera de Mérida

and the Sierra de Pera, in Mérida, Tachira, and Zulia

States, 505—2,200 m asl.

Selected references: Rivero (1968b, 1985); Mijares-

Urrutia (1990a); La Marca (1994c); Ruiz-Carranza and

Lynch (1995); Guayasamin and Barrio-Amoros (2005);

Sefiaris and Ayarzagtiena (2005); Guayasamin et al.

(2009); Rojas-Runjaic et al. (2012).

Genus Vitreorana Guayasamin, Castroviejo-Fisher,

Trueb, Ayarzagtiena, Rada and Vila, 2009

Type species: Centrolenella antisthenesi Goin, 1963, by

original designation.

Vitreorana antisthenesi (Goin 1963)*

Holotype: MBUCV 4033.

Type locality: “Parque Nacional de Rancho Grande,

Aragua, Venezuela.”

Distribution: Region 2. Cordillera de la Costa, 220-

1,200 m asl.

Selected references: Goin (1963); Rivero (1968b);

Cannatella and Lamar (1986); Manzanilla et al. (1995),

Sefiaris and Ayarzagtena (2005); Barrio-Amoros

(2006c); Guayasamin et al. (2009).

Vitreorana castroviejoi Ayarzaguena and Sefiaris, 1997 *

Holotype: MHNLS 13356.

Type locality: “Cerro El Humo, Peninsula de Paria, Estado

Sucre, Venezuela. (10°42’N-62°37’W). 750 m snm.”

July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

! r , S ‘

Photo:

Ae fre A Ri

26. Rhinella aff. proboscidea.

Javier Mesa.

La Escalera, Bolivar.

a he = 2 on ‘ =o ’ z : »

27B. Rhinella sternosignata. Ventral view. Campamento

Arassari, Acequias River, Barinas. Photo: César Barrio-Amoros.

Patgh ~ Par a a ene |

? a 4

29. Centrolene daidalea. Male. Cerro Las Antenas, Lajas river

basin, sierra de Perija, Zulia. Photo: Fernando Rojas-Runjaic.

Distribution: Region 2. Endemic on the Peninsula de

Paria, Sucre State, 750-800 m asl.

Selected references: Ayarzagiiena and Sefiaris (1997);

Sefiaris and Ayarzagtiena (2005); Guayasamin et al.

(2009).

Vitreorana gorzulae (Ayarzagtiena 1992)

Holotype: MHNLS 11221.

Amphib. Reptile Conserv. 22

27A. Rhinella sternosignata. Campamento Arassari, Acequias

River, Barinas. Photo: César Barrio-Amoros.

28. Centrolene altitudinalis. El Cedral, Altos de San Luis,

Merida. Photo: César Barrio-Amoros.

4 «= 4 5H ¥ 4 5 2 >.

30. Centrolene notosticta. Male. Campamento Guacharaca, Cafio

Tétari, Sierra de Perija, Zulia. Photo: Fernando Rojas-Runjaic.

Type locality: “Cerro Auyantepuy-Centro. Edo. Bolivar.

Venezuela. (5°56’N-62°34’W). 1,850 m snm.”

Distribution: Region 5. Widespread in the Guiana Shield

(Venezuela and Guyana), 430—1,850 m asl. In Venezuela,

known from several localities in the Sierra de Lema, the

Gran Sabana, and Auyan-tepui.

Remarks: Duellman and _ Sefiaris (2003) place

Centrolenella auyantepuiana_ (Ayarzaguiena 1992)

(reported as Hyalinobatrachium auyantepuianum by

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

Barrio-Amoros 1998), in synonymy with Centrolene

gorzulae based on its trilobate liver and a small

humeral spine not reported before in this taxon. Kok

and Castroviejo-Fisher (2008) synonymized Centrolene

papillahallicum Noonan and Harvey, 2000 with what

they called V. gorzulai. Later, using integrative taxonomy,

Castroviejo-Fisher et al. (2009) placed Centrolene lema

into synonymy with V. gorzulae. Though the name

gorzulae is a masculine patronym (honoring Stefan

Gorzula) that seems to be correctly emended to gorzulai,

this is not the case according to Article 31.1.1 of the Code

(ICZN 1999).

Selected references: Ayarzagiiena (1992); Duellman

(1997); Gorzula and Sefiaris (1998); ICZN (1999);

Duellman and Sefiaris (2003); Sefiaris and Ayarzagtiena

(2005); Kok and Castroviejo-Fisher (2008); Kok and

Kalamandeen (2008); Myers and Donnelly (2008);

Barrio-Amords and Duellman (2009); Castroviejo-

Fisher et al. (2009); Guayasamin et al. (2009); Sefiaris

et al. (2014).

Vitreorana helenae (Ayarzaguena 1992)

Holotype: MHNLS 9431.

Type locality: “Quebrada Jaspe, San Ignacio de Yuruani,

Edo. Bolivar. Venezuela.”

Distribution: Region 5. Venezuela and Guyana. In

Venezuela the species 1s known from several localities in

the Gran Sabana Region.

Remarks: Sefiaris (1997) reported Cochranella

oyampiensis from Salto Karuay, a locality in the Gran

Sabana Region, Bolivar State. Kok and Castroviejo-

Fisher (2008) demonstrated that those specimens

correspond to Vitreorana helenae. Purported differences

between both taxa listed by Sefiaris and Ayarzagtiena

(2005) were interpreted as intraspecific variation.

Selected references: Ayarzagiiena (1992); Duellman

(1993); Sefiaris (1997); Sefiaris and Ayarzagtiena

(2005); Kok and Castroviejo-Fisher (2008); Kok and

Kalamandeen (2008); Guayasamin et al. (2009); Sefiaris

et al. (2014).

Subfamily Hyalinobatrachinae

Guayasamin, Castroviejo-Fisher, Trueb,

Ayarzagiiena, Rada and Vila, 2009

Genus Celsiella Guayasamin, Castroviejo-Fisher, Trueb,

Ayarzagutena, Rada and Vila, 2009

Type species: Centrolenella revocata Rivero, 1985, by

original designation.

Celsiella revocata (Rivero 1985)*

Holotype: UPR-M 5295.

Type locality: Colonia Tovar, 1,800 m, D.F., Venezuela.

Distribution: Region 2. Southern versant of the

Amphib. Reptile Conserv.

Cordillera Litoral and the western sector of the Cordillera

de la Costa, 1,200—1,800 m asl.

Selected references: Rivero (1985); Ruiz-Carranza and

Lynch (1991, 1998); Myers and Donnelly (1997); Sefiaris

and Ayarzagtiena (2005); Guayasamin et al. (2009).

Celsiella vozmedianoi

(Ayarzagtiena and Sefiaris 1997)*

Holotype: MHNLS 13355.

Type locality: “Cerro El Humo, Peninsula de Paria,

Estado Sucre, Venezuela. (10°42’N-62°37’W), 750 m

snm.”

Distribution: Region 2. Endemic on the Peninsula de

Paria, Sucre State.

Selected references: Ayarzagiiena and Sefiaris (1997);

Sefiaris and Ayarzagtiena (2005); Guayasamin et al.

(2009).

Genus Hyalinobatrachium

Ruiz-Carranza and Lynch, 1991

Type species: Hylella fleischmanni Boettger, 1893, by

original designation.

HAyalinobatrachium cappellei

(van Lidth de Jeude 1904)

Holotype: RMNH 4463.

Type locality: River Saramacca and neighboring areas,

Suriname.

Distribution: Region 5. Brazil, Venezuela, Guyana,

Suriname, and French Guiana. Widespread in the

Venezuelan Guayana (Amazonas and Bolivar States).

Remarks: Many references to this species are found

under the names: Hyalinobatrachium crurifasciatum

Myers and Donnelly, 1997, H. eccentricum Myers and

Donnelly, 2001, H. ignioculus Noonan and Bonett,

2003 and H. taylori. Barrio-Amoros and Castroviejo-

Fisher (2008b) commented on variation, vocalization

and several morphological traits (under H. ignioculus).

Castroviejo-Fisher et al. (2011) placed the three previous

names into the synonymy of H. cappel/lei and identified

many published records of H. taylori as H. cappellei.

Selected references: Goin (1964); Sefiaris and

Ayarzagtiena (1994, 2005); Myers and Donnelly

(1997, 2001); Duellman and Sefiaris (2003); Barrio-

Amoros and Castroviejo-Fisher (2008b); Kok and

Kalamandeen (2008); Barrio-Amords and Duellman

(2009); Guayasamin et al. (2009); Castroviejo-Fisher et

al. (2011); Sefiaris et al. (2014).

HAyalinobatrachium duranti (Rivero 1985)*

Holotype: UPR-M 5811.

Type locality: La Mucuy, 2,172 m, Estado Mérida,

Venezuela (also type locality of synonyms H.

July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

31A. Centrolene venezuelensis. Female. Paramo Tétari, Sierra de 31B. Centrolene venezuelensis. Male. La Motus, Mérida. Photo:

Perija, Zulia. Photo: Fernando Rojas-Runjaic. César Barrio-Amoros.

9 "

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32A. Espadarana andina. Male. San Juan de Colon, Tachira. 32B. Espadarana andina. Male. La Azulita, Mérida. Photo:

Photo: César Barrio-Amoros. César Barrio-Amoros.

' :

a. sy ca

“ai ' = act :

33. Vitreorana antisthenesi. Male. Palmichal, Beyuma, Carabobo. 34. Vitreorana castroviejoi. Male. Las Melenas, Irapa, Sucre.

Photo: César Barrio-Amoros. Photo: César Barrio-Amoros.

loreocarinatum (Rivero 1985) and H. ostracodermoides Ayalinobatrachium fragile (Rivero 1985)*

(Rivero 1985).

Distribution: Region 1. Andes of Mérida State, 1,830-— Type: UPR-M 5938.

2,400 m asl. Type locality: “Mundo Nuevo, entre Manrique y La

Remarks: Sefiaris (1999) considered Hyalinobatrachium Sierra, 396 m, Estado Cojedes, Venezuela.”

loreocarinatum, H. _ pleurostriatus, and H. _ Distribution: Region 2. Western and central sectors of

ostracodermoides synonymous with H. duranti. the Cordillera de la Costa, 100—1,200 m asl.

Selected references: Rivero (1985); Sefiaris (1999); Remarks: Hyalinobatrachium fleischmanni has been

Sefiaris and Ayarzagtiena (2005). listed as occurring in Venezuela until recently, but is no

Amphib. Reptile Conserv. 24 July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

longer considered part of the Venezuelan herpetofauna.

Most of the older records were misidentifications

between H. fragile and H. guairarepanense.

Selected references: Rivero (1985); Sefiaris and

Ayarzagtiena (2005); Barrio-Amoros (2006c).

Hyalinobatrachium guairarepanense Sefiaris, 1999*

Holotype: MHNLS 13731.

Type locality: “Quebrada Chacaito, Parque Nacional

El Avila, (10°30’36”N, 66°51°44”W), 980 m, Distrito

Federal, Venezuela.”

Distribution: Region 2. Known only from the central

sector of Cordillera Litoral, in Cordillera de la Costa,

720-1,200 m asl.

Remarks: Hyalinobatrachium fleischmanni was listed as

occurring in northern Venezuela in error (Rivero 1961).

Most records were misidentifications of H. fragile and

H. guairarepanense. The image presented here (Image

41) is of an individual from near the type locality (but

at around 1,200 m; J. Vieira, pers. comm.), where it had

not been seen since its original description. Considered

Endangered under Venezuelan Red Book criteria (Sefiaris

et al. 2015a). Here the ending of the original species

name guairarepanensis is emended as guairarepanense,

as Hyalinobatrachium is a neuter genus (Ruiz-Carranza

and Lynch 1991), following the Code’s article 30.1.2.

(ICZN 1999).

Selected references: Ginés (1959); Rivero (1961,

1964a,b); Goin (1964); Tello (1968); Cannatella and

Lamar (1986); Manzanilla et al. (1995); Myers and

Donnelly (1997); ICZN (1999); Sefiaris (1999); Barrio-

Amoros (2013); Sefiaris et al. (2015).

HAyalinobatrachium iaspidiense (Ayarzagiiena 1992)

Holotype: EBD 28803.

Type locality: “Quebrada Jaspe, San Ignacio de Yuruani,

Edo. Bolivar. Venezuela.”

Distribution: Region 5. Considered until recently

endemic to the Venezuelan Gran Sabana. For several

years, reports of this species have come from French

Guiana, Surinam, Brazil, Ecuador, and Peru (Guayasamin

and North 2009; Yanez-Mufioz et al. 2009), showing one

of the widest distributions among centrolenid frogs. In

Venezuela, however, only known from the southeastern

sector (east of the Parima-Maigualida mountain chain).

Selected references: Ayarzagiiena (1992); Sefiaris and

Ayarzagtiena (2004, 2005); Guayasamin and North

(2009); Yanez-Mufioz et al. (2009); Castroviejo-Fisher

et al. (2011); Sefiaris et al. (2014).

HAyalinobatrachium mesai

Barrio-Amoros and Brewer-Carias, 2008 *

Holotype: EBRG 4644.

Type locality: Southern slope of Sarisarifiama-tepul,

Amphib. Reptile Conserv.

Bolivar State, Venezuela (4°25’N, 64°7’W), elevation

420 m.

Distribution: Region 5. Only known from the type

locality.

Remarks: Guayasamin and North (2009) believe that due

to the similarity with Hyalinobatrachium iaspidiense, H.

mesai is likely a synonym. However, CBA field notes

clearly note green bones as the main distinguishing

character (Barrio-Amoros and Brewer-Carias 2008).

Selected references: Barrio-Amoros and Brewer-Carias

(2008); Guayasamin and North (2009); Castroviejo-

Fisher et al. (2011).

Hyalinobatrachium mondolfi

Ayarzaguena and Sefiaris, 2001

Holotype: MHNLS 12710.

Type locality: “Primer raudal del Cafio Acoima,

afluente del Rio Grande (8°22’N, 61°32’W), 15 m snm,

estribaciones de la serrania de Imataca, Estado Delta

Amacuro.”

Distribution: Regions 4, 5, 6. Widespread in lowlands

of Amazonia in southern Venezuela and the Guianas, and

from Colombia and Brazil into Bolivia. In Venezuela,

only known from Delta Amacuro State.

Selected references: Sefiaris and Ayarzagtiena (2001,

2004, 2005); Castroviejo-Fisher et al. (2011).

Hyalinobatrachium orientale (Rivero 1968)

Holotype: MCZ 72497.

Type locality: “Cerro Turumiquire, 1,200 m, Estados

Sucre-Monagas, Venezuela.”

Distribution: Region 2. Tobago and _ northeastern

Venezuela. In Venezuela, restricted to the eastern part of

Cordillera de la Costa (Serrania de Paria and Macizo de

Turimiquire), in Monagas and Sucre States.

Remarks: Hyalinobatrachium orientale was believed to

be a species complex including H. orocostale (Cannatella

and Lamar 1986). Castroviejo-Fisher et al. (2008)

demonstrated that H. orocostale may be differentiated, as

H. orientale remains only known from NE Venezuela and

Tobago (Hardy 1984b; Jowers et al. 2014). The genetic

distance between Venezuelan and Tobagoan populations

is low enough that Jowers et al. (2014) did not elevate

subspecies tobagoensis to species status. However, a

biogeographic rationale could support its elevation, but

would require further investigation. CBA and G. Rivas

collected H. orientale in Macuro (Peninsula de Paria) at

near sea level (elevation 5 m; specimens at CVULA),

extending the elevational range from 5—1,200 m.

Selected references: Rivero (1968b); Hardy (1984b);

Cannatella and Lamar (1986); Ruiz-Carranza and

Lynch (1991); Sefiaris and Ayarzagtiena (1993, 2005);

Manzanilla et al. (1995); Ayarzagtiena and Sefiaris

(1996); Duellman (1997); Gorzula and Sefiaris (1998);

Castroviejo-Fisher et al. (2008); Jowers et al. (2014).

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Catalogue of the amphibians of Venezuela

35A. Vitreorana gorzulae. Male. Las Lajas, Sierra de Lema, 35B. Vitreorana gorzulae. Female. Las Lajas, Sierra de Lema,

Bolivar. Photo: Charles Brewer-Carias. Bolivar. Photo: César Barrio-Amoros.

: t fade :

36. Vitreorana helenae. Male. Chivaton, Gran Sabana, Bolivar. 37. Celsiella vozmedianoi. Male. Las Melenas, Irapa, Sucre.

Photo: César Barrio-Amoros. Photo: César Barrio-Amoros.

38A. Hyalinobatrachium cappellei. Male. Chivaton, Gran 38B. Hyalinobatrachium cappellei. Male. Quebrada de Jaspe,

Sabana, Bolivar. Photo: César Barrio-Amoros. Bolivar. Photo: César Barrio-Amoros.

Amphib. Reptile Conserv. July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

Sie

r |

Ee , ae _ -

38C. Hyalinobatrachium cappellei. Detail of the iris. La Laja,

Sierra de Lema, Bolivar. Photo: Charles Brewer-Carias.

40A. Hyalinobatrachium fragile. Haciendo la Elvira, Guatopo,

Guarico. Photo: Walter Schargel.

ee eos =

41. Hyalinobatrachium guairarepanense. Quebrada Chacaito,

Caracas. Photo: Jose Vieira.

HAyalinobatrachium orocostale (Rivero 1968)*

Holotype: MCZ 47501

Type locality: “Cerro Platillon (Hacienda Picachitos),

Cordillera del Interior, 1,200 m, Estado Guarico,

Venezuela.”

Distribution: Region 2. Apparently restricted to humid

mountainous forests of Serrania del Interior, a small

Amphib. Reptile Conserv.

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39. Hyalinobatrachium duranti. Male. Chorotal, Mérida. Photo:

César Barrio-Amoros.

40B. Hyalinobatrachium fragile. Cerro El Tigre, Sierra de Aroa,

Yaracuy. Photo: Fernando Rojas-Runjaic.

42. Hyalinobatrachium iaspidiense. Salto Rio Lindo, Sierra de

Imataca, Delta Amacuro. Photo: César Barrio-Amoros.

mountain chain parallel to the Serrania del Litoral, both

within the central sector of Cordillera de la Costa in

Venezuela.

Remarks: Erected from synonymy with

Hyalinobatrachium orientale by Castroviejo-Fisher et al.

(2008).

Selected references: Rivero (1968); Sefiaris and

Ayarzagtiena (2005); Castroviejo-Fisher et al. (2008).

July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

HAyalinobatrachium pallidum (Rivero 1985)*

Holotype: UPR-M 4554.

Type locality: “Guacharaquita, entre La Grita y Paramo

de La Negra, 1,768 m, Edo Tachira, Venezuela.”

Distribution: Region 1. Cordillera de Mérida in its

western, southern, and eastern versants (Mérida, Tachira,

and Barinas States), and Sierra de Perija (Zulia State).

Remarks: This species has a wider distribution than

previously known. Formerly only known from its type

locality until Sefiaris and Ayarzagtiena (2005) redescribed

the species based on new topotypic specimens. Later,

Guayasamin et al. (2009) assigned a population

from the eastern versant of the Andes, reported

as Hyalinobatrachium ibama_ by _ Barrio-Amoros

(2006d) to H. cf. pallidum. Based on molecular work

(Guayasamin et al. 2009 and unpub.), it is doubtful that

H. ibama is the species occupying the eastern versant

of the Cordillera de Mérida (Barrio-Amoros 2006d),

but is more likely H. pallidum (Image 47A). Actually,

this implies that H. ibama from Colombia could be a

junior synonym of H. pallidum. Another population

from Quebrada La Rana, near La Macana, Mérida, is

morphologically slightly different from the H. pallidum

population at San Isidro, Barinas (Image 47B). More

research is needed to resolve the status of H. pallidum

in the Venezuelan Andes. More recently, Rojas-Runjaic

et al. (2012) reported four additional localities, along

the eastern versant of the Sierra de Perija, at elevations

1,132-1,832 m asl.

Selected references: Rivero (1985); Sefiaris and

Ayarzagiena (2005); Barrio-Amorés (2006d);

Guayasamin et al. (2009); Rojas-Runjaic et al. (2012).

HAyalinobatrachium tatayoi

Castroviejo-Fisher, Ayarzaguiena and Vila, 2007

Holotype: MHNLS 17174.

Type locality: “Stream near Tokuko (09°50’30.6’N,

72°49’ 13.6°W; 301 masl.), Estado Zulia, Venezuela.”

Distribution: Regions 1, 6. Known from four localities

in lowlands and uplands of the eastern versant of Sierra

de Perija, Zulia State (Rojas-Runjaic et al. 2012). CBA

and Erik Arrieta also collected this species at Rio Frio

(CVULA 8201-09) at 40 m, in a small stream flowing

from Cordillera de Mérida through the lowlands facing

Lake Maracaibo, Merida State. Elevation range 40-512

m asl.

Remarks: Castroviejo-Fisher et al. (2009) showed that

Hyalinobatrachium tatayoi is embedded in a clade of

specimens of H. fleischmanni and these two species could

be conspecific. If H. fleischmanni is a species complex it

requires further taxonomic and nomenclatural revision.

Selected references: Castroviejo-Fisher et al. (2007,

2009); Rojas-Runjaic et al. (2012).

Amphib. Reptile Conserv.

Ayalinobatrachium taylori (Goin 1968)

Holotype: BM 1939.1.1.65.

Type locality: “750 ft. (228 m) along the New River,

Guyana”

Distribution: Region 5. Venezuela, Guyana, Suriname,

and French Guyana. Widespread in the Venezuelan

Guayanas. Elevational distribution in Venezuela from

450 m (La Laja, Sierra de Lema) to 1,850 on Auyan-tepul

(Sefiaris and Ayarzagtiena 1994). Lower elevations are

known in Guyana, French Guiana, and Suriname (Kok

and Castroviejo-Fisher 2008). Amazonas state reports

(Sefiaris and Ayarzagtiena 2005) are doubtful and must

be verified.

Remarks: Hyalinobatrachium taylori has a long history

of misidentifications. The type series contained three

different species, and many published records refer to H.

cappellei (Castroviejo-Fisher et al. 2011).

Selected references: Goin (1968); Ruiz-Carranza

and Lynch (1991); Ayarzagtiena (1992); Sefiaris and

Ayarzagtiena (1993, 2005); Gorzula and Sefiaris

(1998); Lescure and Marty (2000); Noonan and Bonet

(2003); Kok and Castroviejo-Fisher (2008); Kok and

Kalamandeen (2008); Barrio-Amords and Duellman

(2009); Castroviejo-Fisher et al. (2011); Sefiaris et al.

(2014).

Family Ceratophryidae Tschudi, 1838

Remarks: Recovered Leptodactylidae by _ Frost

(2006). Faivovich et al. (2014) treated its phylogenetic

relationships.

Selected references: Frost (2006); Faivovich et al.

(2014).

Genus Ceratophrys Wied-Neuwied, 1824

Type species: Ceratophrys varius Wied-Neuwied, 1824

(=Bufo auritus Raddi, 1823), by subsequent designation

of Fitzinger (1843).

Remarks: Ceratophrys cornuta (Linnaeus 1758)

was considered to occur in Venezuela (Rivero 1961;

Lynch 1982). This was accepted by many subsequent

authors until Barrio-Amords (2004) deleted it from

the Venezuelan checklist for lack of any voucher or

photographs of the species in the country.

Ceratophrys calcarata Boulenger, 1890

Holotype: BMNH 1947.2.17.28 (formerly 89.12.16.168).

Type locality: “Colombia.”

Distribution: Region 3, 6. Caribbean lowlands of

Colombia and Venezuela. In Venezuela, distribution 1s in

the northwestern portion of the country (Maracaibo Lake

Basin, Falcon and Lara States), with a disjunct population

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

43. Hyalinobatrachium mesai. Male. Southern slope of — 44. Hyalinobatrachium mondolfii. Male. Salto Rio Lindo, Sierra

Sarisarifiama tepui, Bolivar. Photo: Mark Moffett. de Imataca, Delta Amacuro. Photo: Fernando Rojas-Runjaic.

45. Hyalinobatrachium orientale. Male. Macuro, Peninsula de 46. Hyalinobatrachium orocostale. Cerro Platillon, Guarico.

Paria, Sucre. Photo: César Barrio-Amoros. Photo: Santiago Castroviejo-Fisher.

47A. Hyalinobatrachium aff. pallidum. Male. San Isidro, 47B. Hyalinobatrachium pallidum. Male. Quebrada La Rana,

Barinas. Photo: César Barrio-Amoros. south of Santa Cruz de Mora, Mérida. Photo: César Barrio-

Amoros.

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Catalogue of the amphibians of Venezuela

48A. Hyalinobatrachium tatayoi. Male. Rio Frio, Mérida. Photo:

César Barrio-Amoros.

48B. Hyalinobatrachium tatayoi. Female. Ipika, Tokuko river

basin, Sierra de Perija, Zulia. Photo: Fernando Rojas-Runjaic.

49. Hyalinobatrachium taylori. Female. Quebrada de Jaspe,

Gran Sabana, Bolivar. Photo: César Barrio-Amoros.

50A. Gaaipnas calcarata. pene Nuevo, Falcon. Photo:

César Barrio-Amoros.

50B. Cua Ramat Falcon. Photo: Edward Camargo.

in the surroundings of Puerto Ayacucho, southeast of the

Orinoco River in Amazonas State.

Remarks: Faivovich et al. (2014) recovered the

phylogenetic relationships of the family Ceratophryidae,

placing Ceratophrys calcarata as a sister species of C.

cornuta.

Selected references: Boulenger (1890); Lutz (1927);

Ginés (1959); Rivero (1961, 1964b); Lynch (1982);

La Marca (1986, 1995a); Mijares-Urrutia and Arends

Amphib. Reptile Conserv.

50C. Cenaoniae ne Bete: Ayacucho, Anarene

Photo: Zelimir Cernelic.

(1993); Frost et al. (2006); Faivovich et al. (2014).

Family Ceuthomantidae

Heinicke, Duellman, Trueb, Means,

MacCulloch and Hedges, 2009

Remarks: Includes Pristimantinae sensu Padial et al.

(2014), including Ceuthomantinae Heinicke et al. (2009),

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Barrio-Amoros et al.

and follows Heinicke et al (2018) intended for a robust

framework for a standarized and revised Terraranae

taxonomy.

Genus Ceuthomantis Heinicke, Duellman, Trueb,

Means, MacCulloch and Hedges, 2009

Type species: Ceuthomantis smaragdinus Heinicke,

Duellman, Trueb, Means, MacCulloch, and Hedges,

2009, by original designation.

Ceuthomantis aracamuni

(Barrio-Amoros and Molina 2006)*

Holotype: MHNLS 17042.

Type locality: Summit of Cerro Aracamuni (01°28’36”N,

65°50’7”"W), elevation 1,493 m, Amazonas State,

Venezuela.

Distribution: Region 5. Endemic to Cerro Aracamuni,

a granitic mountain in Amazonas State, southern

Venezuela.

Remarks: Described originally in genus

Eleutherodactylus (Barrio-Amords and Molina

2006), subsequently transferred to Pristimantis based

on Heinicke et al. (2007), and finally transferred to

Ceuthomantis by Heinicke et al. (2009).

Selected references: Barrio-Amoros and Molina (2006);

Heinicke et al. (2009); Barrio-Amoros (2010c).

Ceuthomantis cavernibardus

(Myers and Donnelly 1997)

Holotype: AMNH 131537.

Type locality: “North base of Pico Tamacuari, 1,160-

1,200 m elevation. Sierra Tapirapecd, Amazonas,

Venezuela (1°13’N, 64°42’W).”

Distribution: Region 5. Restricted to the Sierra

Tapirapeco, in the extreme south of Amazonas State in

Venezuela along the northern border of Brazil.

Remarks: Described originally as a member of genus

Eleutherodactylus (Myers and Donnelly 1997), and

considered a member of Pristimantis by Barrio-Amoros

and Brewer-Carias (2008), before being transferred to

Ceuthomantis by Heinicke et al. (2009).

Selected references: Myers and Donnelly (1997);

Caramaschi and Niemeyer (2005); Barrio-Amoros and

Brewer-Carias (2008); Heinicke et al. (2009); Barrio-

Amoros (2010c).

Ceuthomantis duellmani Barrio-Amoros, 2010*

Holotype: EBRG 4676.

Type locality: Edge of Sima Mayor, Sarisarifiama-tepui

(1°28°36"N, 65°50’7°W), elevation 1,350 m, Bolivar

State, Venezuela.

Distribution: Region 5. Endemic to Cerro Sarisarifiama-

tepui in Bolivar State; southeastern Venezuela.

Amphib. Reptile Conserv.

Selected references: Barrio-Amoros and Brewer-Carias

(2008); Barrio-Amoros (2010c).

Family Craugastoridae

Hedges, Duellman and Heinicke, 2008

Remarks: This very diverse group of Terrarana is

still under revision (e.g., Padial et al. 2014; Heinicke

et al. 2015, 2018). Pyron and Wiens (2011) found

Craugastoridae embedded in Strabomantidae. Padial et al.

(2014) validated a monophyletic Craugastoridae including

subfamily Ceuthomantinae Heinicke et al., 2009. Heinicke

et al. (2018) showed Strabomantidae monophyletic with

respect to Craugastoridae, a view shared here. Heinicke

et al. (2015) did not allow unequivocal placement of

Tachiramantis into a subfamily, however, Heinicke et al.

(2018) placed it into Craugastoridae.

Genus Tachiramantis

Heinicke, Barrio-Amoros and Hedges, 2015

Type species: Eleutherodactylus prolixodiscus Lynch,

1978, by original designation.

Remarks: Recently erected by Heinicke et al. (2015)

to accommodate a clade of terraranan frogs deeply

divergent from Pristimantis. More species currently in

Pristimantis will probably be shown to be members of

Tachiramantis, such as P. melanoproctus, P. mondolfii

and P. tayrona (due to its similarity to P. prolixodiscus).

Tachiramantis lentiginosus (Rivero 1984)*

Holotype: UPR-M 6060.

Type locality: “Guacharaquita, 1,768 m, entre La Grita y

Paramo de La Negra, Edo. Tachira, Venezuela.”

Distribution: Region 1. Known from scattered localities

in cloud forests of Tachira and Merida States.

Selected references: Rivero (1982c).

Tachiramantis prolixodiscus (Lynch 1978)

Holotype: KU 132726.

Type locality: “30 km ENE Bucaramanga, road to

Cucuta, Departamento Santander, Colombia, 2,485 m.”

Distribution: Region 1. Cordillera Oriental in Colombia

and Venezuela. In Venezuela, present in the Cordillera de

Mérida and the Sierra de Perija. Barrio-Amoros (2010a)

reported it from Calderas, Barinas State, and Barrio-

Amoréos et al. (20101) from Perija; Camargo et al. (2014)

provided first report from Lara State.

Remarks: According to Lynch (2003), Eleutherodactylus

chlorosoma Rivero, 1984 is a junior synonym. Heinicke

et al. (2015) assigned prolixodiscus to Tachiramantis as

its type species.

Selected references: Lynch (1978, 2004); Rivero (1982);

Barrio-Amoros (2010a); Barrio-Amoros et al. (2010);

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Catalogue of the amphibians of Venezuela

Javier Mesa.

53. Tachiramantis prolixodiscus. Los Alcaravanes, Calderas;

Barinas. Photo: César Barrio-Amoros.

55. Allobates caribe. Southern slopes Cerro El Humo, Peninsula

de Paria, Sucre. Photo: César Barrio-Amoros.

Camargo et al. (2014); Heinicke et al. (2015).

Family Dendrobatidae Cope, 1865

Remarks: Herein, family Dendrobatidae is treated to

include subfamilies Aromobatinae and Dendrobatinae,

which does not follow Grant’s et al. (2006, 2017)

classification as two separate families.

Amphib. Reptile Conserv. 32

52. Zachiramantis lentiginosus. Female. Guaraque, Mérida.

Photo: César Barrio-Amoros.

* és ; i-o4 a + +

César Barrio-

54. Allobates algorei. Rio Frio, Tachira. Photo:

Amoros.

f. fh ig a

a A" a.

56. Allobates femoralis. Yasuni, Ecuador. Photo: César Barrio-

Amoros.

Subfamily Aromobatinae Grant, Frost, Caldwell,

Gagliardo, Haddad, Kok, Means, Noonan, Schargel

and Wheeler, 2006

Remarks: Grant et al. (2006) erected family

Aromobatidae for dendrobatoid frogs without alkaloids.

Santos et al. (2009) returned to a comprehensive

Dendrobatidae with Aromobatinae as a subfamily, which

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Barrio-Amoros et al.

is followed here. Pyron and Wiens (2011) opined that

there was not a need for subfamilies in Dendrobatidae.

Grant’s et al. (2017) claim against Santos (2009) and

Barrio-Amoros et al. (2010b) opinions.

Genus Prostherapis Cope, 1868

Type species: Prostherapis inguinalis Cope, 1868, by

original designation.

Prostherapis dunni (Rivero 1961)*

Holotype: FMNH 35987.

Type locality: “above Caracas,

Venezuela.”

Distribution: Region 2. Only found inthe Caracas Region.

Remarks: Little 1s known about this frog. Grant et al.

(2006) did not include this species in their molecular

analysis, and its morphological characteristics prevent

clear assignment to an established genus. Until molecular

data become available, it should remain in the genus to

which it was originally assigned, which is not currently

used for any other dendrobatid species. Its unknown

phylogenetic relationships (La Marca 2004; Grant et al.

2006) require more data to determine its proper place

among dendrobatids. It has likely suffered a dramatic

decline in populations due to habitat destruction in the

Caracas area (La Marca 2004). Recent surveys by FRR

and collaborators in June, July, and December 2015

failed to detect any individuals. It is currently considered

Critically Endangered in the Venezuelan Red Book

(Rojas-Runjaic and Sefiaris 2015a) but could be already

extinct.

Selected references: Rivero (1961, 1964a, 1988); Tello

(1968); Myers et al. (1991); La Marca (2004); Grant et

al. (2006); Barrio-Amorés (2013); Rojas-Runjaic and

Sefiaris (2015a).

Distrito Federal,

Genus Allobates Zimmermann and Zimmermann, 1988

Type species: Prostherapis femoralis Boulenger, 1884,

by original designation.

Remarks: Barrio-Amoros et al. (2011) and Barrio-

Amoros and Santos (2012) noted the difficulty of

distinguishing some A//obates and Aromobates species

using the morphological traits proposed by Grant et al.

(2006).

Allobates algorei Barrio-Amoros and Santos, 2009*

Holotype: EBRG 5560.

Type locality: Rio Negro, Municipio Cordoba, Estado

Tachira, Venezuela, “07°34.723°N, 72°10.739°W,”

elevation 482 m.”

Distribution: Region 1. Cordillera Oriental de Colombia

(Acevedo et al. 2019) and extreme southwestern

piedmont of the Venezuelan Andes, on both northern and

Amphib. Reptile Conserv.

southern sides of the Tachira depression.

Selected references: Barrio-Amoros and Garcia Porta

(2003); Barrio-Amoros and Santos (2009).

Allobates bromelicola (Test 1956)*

Holotype: UMMZ 113027.

Type locality: “1,375 m on Pico Periquito, Rancho

Grande, Estado Aragua, Venezuela.”

Distribution: Region 2. Surroundings of the type

locality, 1,310—1,375 m asl.

Remarks: Nothing is known of this species beyond

the original description and some natural history notes

by Dixon and Rivero Blanco (1985). It appears to be

quite rare, difficult to observe and collect, or it may have

suffered a severe decline.

Selected references: Test (1956); Ginés (1959); Rivero

(1961, 1964a, 1988); Dixon and Rivero Blanco (1985);

Myers et al. (1991); Manzanilla et al. (1995); La Marca

and Mijares-Urrutia (1997); Barrio and Fuentes (1999a),;

Barrio-Amoros (2006c).

Allobates caribe

(Barrio-Amoros, Rivas and Kaiser 2006)*

Holotype: MHNLS 17462.

Type locality: “Southern slope of Cerro El Humo,

Peninsula de Paria, Estado Sucre, Venezuela

(10°41’094”N, 62°37°147"W), elevation 1,050 m.”

Distribution: Region 2. Only known from type locality.

Remarks: Described as Colostethus caribe, it was

transferred to Allobates by Lotters et al. (2007: 55) a shift

agreeable to the current authors, but later (mistakenly)

transferred to Anomaloglossus (Lotters et al. 2007: 60)

without explanation. Frost (2018) used A//obates for

this species by implication. Several recent attempts to

secure more specimens in the same general area failed

(G. Rivas, pers. comm. ).

Selected references: Barrio-Amoroés et al. (2006a);

Lotters et al. (2007).

Allobates femoralis (Boulenger 1884)

Syntypes: BMNH — 1947.2.14.21-22; Lectotype

designated by Silverstone 1976: BMNH 1947.2.14.21

(sensu Frost 2017).

Type locality: “Yurimaguas, Huallaga

(Departamento Loreto), Northern Peru.”

Distribution: Widely distributed in the Amazonian

Region (Colombia, Peru, Ecuador, Bolivia, Brazil,

Venezuela, Guyana, Suriname, and French Guiana). In

Venezuela, only known from two localities in the extreme

east of Bolivar State.

Remarks: Allobates femoralis was reported for

Venezuela by Duellman (1997) based on one specimen

(KU 167335), which was confused with Ameerega picta

(see explanation in Barrio-Amords 2004 and Barrio-

River,

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Catalogue of the amphibians of Venezuela

Amoros and Santos 2009). Nevertheless, its presence in

the country was shown by Barrio-Amoros and Santos

(2009) using two different recorded calls; voucher

specimens from Venezuela are still lacking.

Selected references: Duellman (1997); Barrio-Amoros

and Santos (2009).

Allobates humilis (Rivero 1980)*

Holotype: UPR-M 3526.

Type locality: “Bocond (Laguneta artificial del

Ministerio de Agricultura), Edo. Trujillo, Venezuela,

1,470 m.”

Distribution: Region 1. Eastern slopes of Venezuelan

Andes, in Trujillo and Barinas States, 1,100—1,470 m asl.

Remarks: Misidentified specimens of A/lobates humilis

reported by Barrio-Amorés and Garcia Porta (2003)

from Tachira State actually represent A. algorei (Barrio-

Amoréos and Santos 2009).

Selected references: La Rivero (1980, 1988); Myers

et al. (1991); Marca et al. (2002); Barrio-Amoroés and

Garcia Porta (2003); Barrio-Amoros (201 0a).

Allobates mandelorum (Schmidt 1932)*

Holotype: FMNH 17788.

Type locality: “Camp at altitude of 8,000 feet (2,630 m)

on Mount Turumiquire, (Estados Sucre and Monagas),

Venezuela.”

Distribution: Region 2. Endemic to Macizo de

Turimiquire, a mountain in the Cordillera de la Costa

Oriental, between Monagas and Sucre states. Known

only from type locality and from Elvecia in Sucre State

(La Marca 1993).

Remarks: Despite the taxonomic revision of La Marca

(1993), where it was not closely related to Mannophryne

or the Colosthetus alboguttatus group (currently

Aromobates), its status remains unsolved. It was

allocated to A/lobates by Grant et al. (2006) without any

substantiation. The relationships within Aromobatinae

remain unknown and it has not been collected since 1932.

Selected references: Schmidt (1932); Ginés (1959);

Rivero (1961, 1982a, 1988); Hardy (1984a); Myers et al.

(1991); La Marca (1993); Grant et al. (2006).

Allobates pittieri

(La Marca, Manzanilla and Miyares-Urrutia 2004)*

Holotype: ULABG 5564.

Type locality: “Venezuela: Estado Aragua: Municipio

Ocumare de la Costa de Oro (antes Municipio Mario

Bricefio Iragorri): quebrada afluente del Rio La Trilla,

170 m snm. entre la Estacion Biologica de Rancho

Grande y Ocumare de la Costa, vertiente Norte del Parque

Nacional Henri Pittier (10°22’52”N, 67°44’67”°W)”

Distribution: Region 2. Distributed throughout

the western Cordillera de la Costa (including

Amphib. Reptile Conserv.

Aragua, Carabobo, and Falcon States) towards the

northeasternmost sector of the Andes (Lara State), 150—

1,700 m asl.

Selected references: La Marca et al. (2004).

Allobates sanmartini

(Rivero, Langone and Prigioni 1986)*

Holotype: MHNM 540.

Type locality: “Las Mayjadas, rio Orinoco, Estado

Bolivar, Venezuela.”

Distribution: Region 5. Known only from the type

locality.

Remarks: Known only from the type series (two

females). Despite several attempts (by CBA, J.C. Santos,

and FRR) no additional individuals have been located.

The type locality, Las Majadas, could be either where

the expedition gathered to send the specimens, or the

last campsite where collected material was processed

(P. Langone, pers. comm.). In the authors’ opinion,

this area 1s unsuitable for dendrobatid frogs, as it is dry

forest similar to that in the Llanos which is devoid of

dendrobatoids. Most likely, the specimens came from

the Campamento Cecilia Magdalena, upper Caura River,

where most of the other amphibian specimens were

collected (Rivero et al. 1986). A recently metamorphed

(CVULA 7860) was collected by CBA in Santa Maria

de Erebato, Upper Caura River area, in the same main

area of Campamento Cecilia Magdalena, which might

be an individual of this species. However, its small

size, and absence of molecular data, prevent a positive

identification.

Selected references: Rivero et al. (1986); Rivero (1988);

Myers et al. (1991); La Marca (1997); Grant et al. (2006).

Allobates undulatus (Myers and Donnelly 2001)*

Holotype: EBRG 3021.

Type locality: “Forest stream on Cerro Yutajé, 1,750 m

(5°46’N, 66°8’W), Amazonas, Venezuela.”

Distribution: Region 5. Endemic to Yutajé, a tepui in the

north of Amazonas State.

Selected references: Myers and Donnelly (2001); Grant

et al. (2006).

Genus Anomaloglossus

Grant, Frost, Caldwell, Gagliardo, Haddad, Kok,

Means, Noonan, Schargel and Wheeler, 2006

Type species: Colostethus beebei Noble, 1923, by

original designation.

Remarks: Genus Anomaloglossus, recognized by

the presence of a median lingual process (MLP), was

described by Grant et al. (1997). Currently 28 species

are known from the Guiana Shield and northern Amazon

(Frost 2018). The seven trans-Andean species with MLP

now belong to Ectopoglossus Grant, Rada, Anganoy-

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

ae wee a=

ae re

ees ae z, en

57. Allobates humilis. Calderas, Barinas. Photo: César Barrio-

Amoros.

59. Anomaloglossus breweri. Cueva El Fantasma, Aprada-tepui,

Bolivar. Photo: Charles Brewer-Carias.

- fd

-'olt

61. Anomaloglossus parkerae. La Escalera region, 860 m asl,

Bolivar. Photo: William E. Duellman.

Criollo, Batista, Dias, Jeckel, Machado and Rueda-

Almonacid, 2017.

Anomaloglossus ayarzaguenai (La Marca 1997)*

Holotype: MHNLS 12949.

Type locality: “Sector central de Cerro Jaua, Estado

Bolivar (4°49’55”N, 64°25’54”W).”

Amphib. Reptile Conserv. 35

58. Allobates pittieri. Nirgua massif, Yaracuy. Photo: Sebastian

Lotzkat.

moffetti. Sarisarifiama-tepui, Bolivar. Photo:

a ?

Bett if : a -7

ie a

nay

62. Anomaloglossus praderioi. Maringma-tepui, Cuyuni-

Mazaruni District, Guyana. Photo: Philippe Kok.

Distribution: Region 5. Known only from type locality,

a tepui in Bolivar State.

Selected references: La Marca (1997a).

Anomaloglossus breweri (Barrio-Amoros 2006)*

Holotype: MHNLS 17044.

Type locality: “Entry of Cueva del Fantasma,

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Catalogue of the amphibians of Venezuela

63. Anomaloglossus roraima. Wei Assipu-tepui, Cuyuni-

Mazaruni District, Guyana. Photo: Philippe Kok.

64A. Anomaloglossus rufulus. Churi-tepui, Chimanta Massif,

Bolivar. Dorso-lateral view. Photo: César Barrio-Amoros.

64B. Anomaloglossus rufulus. Churi-tepui, Chimanta Massif,

Bolivar. Ventral view. Photo: César Barrio-Amoros.

66. Anomaloglossus triunfo. Southern slopes of Cerro Santa

Rosa, Bolivar. Photo: César Barrio-Amoros.

northwestern slope of Aprada tepui, 05°27’N, 62°27’ W,

660 m above sea level, Estado Bolivar, Venezuela.”

Distribution: Region 5. Apparently endemic to Aprada-

tepul.

Selected references:

Sefiaris et al. (2014).

Barrio-Amoros (2006a, 2013);

Amphib. Reptile Conserv.

65. Anomaloglossus eae El Peiton, Siher sigpes

Auyan-tepui, Bolivar. Photo: César Barrio-Amoros.

67. Anomaloglossus verbeeksnyderorum. Male. Tobogan de la

Selva, Amazonas. Photo: Zelimir Cernelic.

Anomaloglossus guanayensis (La Marca 1997)*

Holotype: MHNLS 10708.

Type locality: “Alto rio Parguaza, Serrania de Guanay

(5°55’N y 66°23’W), Estado Amazonas.” Type locality

is actually in Bolivar State.

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Barrio-Amoros et al.

Distribution: Region 5. Endemic to Guanay, a tepui in

Bolivar State.

Selected references: La Marca (1997a); Gorzula and

Sefiaris (1998).

Anomaloglossus moffetti

Barrio-Amoros and Brewer-Carias, 2008*

Holotype: EBRG 4645.

Type locality: Southern slope of Sarisarifiama-tepul,

Camp IV, Estado Bolivar, Venezuela (4°29’N, 64°8’W),

elevation 1,108 m.

Distribution: Region 5. Known only from Sarisarifiama,

a tepui in Bolivar State.

Remarks: It should be compared to Anomaloglossus

ayarzaguenai, as both are from neighboring tepuis, and

could represent the same taxon.

Selected references: Barrio-Amoros and Brewer-Carias

(2008).

Anomaloglossus murisipanensis (La Marca 1997)*

Holotype: MHNLS 11385.

Type locality: “Murisipan-tepui (05°53’N y 62°04’W),

Estado Bolivar, Venezuela, 2,350 m snm.”

Distribution: Region 5. Known only from Murisipan, a

tepui in Bolivar State.

Remarks: Listed as Vulnerable (VU) in the current

Venezuelan Red Book (Royas-Runjaic and Sefiaris

2015b).

Selected references: La Marca (1997a); Gorzula and

Sefiaris (1998); Sefiaris et al. (2014); Rojas-Runjaic and

Sefiaris (2015b).

Anomaloglossus parimae (La Marca 1997)*

Holotype: ULABG 4221.

Type locality: “Pista Constitucién (02°13’49”N,

63°20°00’W’) en las cercanias del Cerro Delgado

Chalbaud, (Amazonas State, Venezuela), 670 m snm.”

Distribution: Region 5. Known only from type locality.

Remarks: It should be compared to Anomaloglossus

tamacuarensis, aS both are from the same mountain

range, and could represent the same taxon.

Selected references: La Marca (1997a).

Anomaloglossus parkerae

(Meinhardt and Parmelee 1996)*

Holotype: KU 167332.

Type locality: “km 112, El Dorado-Santa Elena de

Uairén road (06°01'N, 61°24’ W; 860 m), Estado Bolivar,

Venezuela.”

Distribution: Region 5. La Escalera Region, Sierra

de Lema; probably extended through the Gran Sabana

Region to El Pauji (Bolivar State).

Remarks: Key _ species for

understanding — the

Amphib. Reptile Conserv.

phylogenetic relationships of Anomaloglossus, as 1t was

the first species described from Bolivar State, where the

majority of Anomaloglossus species were discovered

afterwards. It inhabits the northern versant of the Sierra

de Lema, especially in the sector known as La Escalera.

Many expeditions to that area from 2006-2011 failed

to find the species, even in pristine zones. Two putative

populations, one from Kavanayen (EBRG 5729-32,

5775-78) and another from El Pauji (CBA, unpub. data)

could demonstrate its survivorship. However, molecular

data for parkerae (from the type locality) have been

never been generated, which prevents determination of

its affinities.

Selected references: Meinhardt and Parmelee (1996);

Duellman (1997); La Marca (1997a); Gorzula and

Sefiaris (1998); Sefiaris et al. (2014); Fouquet et al.

(2015); Barrio-Amoros (2016).

Anomaloglossus praderioi (La Marca 1997)

Holotype: ULABG 4196.

Type locality: “Monte Roraima, tercera quebrada a

partir de la base (5°10’N y 60°47’W). Parque Nacional

Canaima, sector Oriental (Gran Sabana), Estado Bolivar,

Venezuela. 1,950 m snm.”

Distribution: Region 5. Eastern Gran Sabana in Bolivar

State, and adjacent Guyana, 1,310—1,850 m asl. Known

in Venezuela from Roraima and Sierra de Lema (Kok

2010).

Remarks: The original description (La Marca 1996e)

was based on two adult males. Kok (2010) redescribed

the species with fresh material.

Selected references: La Marca (1997a); Grant et al.

(2006); Kok (2010); Cole et al. (2013); Sefiaris et al.

(2014).

Anomaloglossus roraima (La Marca 1997)

Holotype: ULABG 4197.

Type locality: “Paso de la Muerte,” 60—70 m antes de

la cumbre del tepuy, en el camino “La Rampa,” que

conduce desde la base a la cima del Monte Roraima,

Estado Bolivar, Venezuela. 2,700 m snm.”

Distribution: Region 5. Roraima in Venezuela and Wei

Assipu and Maringma tepuis in Guyana, 1,860—2,700 m

asl.

Remarks: Kok et al. (2013) redescribed the species

with fresh material. Originally described from a single,

subadult female.

Selected references: La Marca (1997a); Grant et al.

(2006); Cole et al. (2013); Kok et al. (2013); Sefiaris et

al. (2014).

Anomaloglossus rufulus (Gorzula 1990)*

Holotype: MHNLS 10361.

Type locality: “Porcion central de Murey Tepui

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Catalogue of the amphibians of Venezuela

oa ; 7.

af . }

= —— ¥ t

68. Anomaloglossus wothuja. Female. Tobogan del Cuao, Cerro

Sipapo, Amazonas. Photo: César Barrio-Amordos.

69B. Aromobates cannatellai. Female. Parque Cascada La

Escalera, Mesa de Pérez, Tachira. Photo: César Barrio-Amoros.

71A. Aromobates ericksonae. Olinda, La Azulita, Mérida. Photo:

César Barrio-Amoros.

en el Macizo de Chimanta (CHIMANTA XVIII)

05°22’N-62°05’W. 2,600 m, Estado Bolivar, Venezuela.”

Distribution: Region 5. Endemic to Chimanta, a tepui

in Bolivar State.

Remarks: Described originally as Dendrobates rufulus,

its the generic allocation changed to Epipedobates (Walsh

1994; Myers 1997) and Allobates (Jungfer and Bohme

2004) until Barrio-Amoros and Santos (2011) allocated it

Amphib. Reptile Conserv.

69A. Aromobates cannatellai. Male. Parque Cascada La

Escalera, Mesa de Pérez, Tachira. Photo: Liz del Valle.

Bk. os i . 4 i ea 2

70. Aromobates duranti. Paramo La Culata, Mérida. Photo:

Jaime Péfaur.

ha!

71B. Aromobates ericksonae. Olinda, La Azulita, Mérida. Photo:

César Barrio-Amoros.

to its current genus based on the presence of an MLP and

(unpub. ) genetic data.

Selected references: Gorzula (1988, 1992); Walsh

(1994); Myers (1997); Barrio and Fuentes (1999a, 2012);

Barrio-Amoros (2001c); Barrio-Amorés and Santos

(2011); Sefiaris et al. (2014).

Anomaloglossus shrevei (Rivero 1961)*

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Barrio-Amoros et al.

Holotype: MCZ 28567.

Type locality: Monte Marahuaca (1,524—1,829 m),

Estado Amazonas, Venezuela.

Distribution: Region 5. Cerros Duida and Marahuaka to

the Upper Orinoco, Amazonas State.

Remarks: Little 1s known about this species beyond

its original description. An additional specimen was

reported by Rivero (1964b), and a male was described by

La Marca (1997).

Selected references: Rivero (1961, 1964b, 1967a, 1988);

La Marca (1997a).

Anomaloglossus tamacuarensis

(Myers and Donnelly 1997)

Holotype: AMNH 131347.

Type locality: “North base of Pico Tamacuari, 1,160—

1,200 m elevation. Sierra Tapirapecd, Amazonas,

Venezuela (1°13'N, 64°42’W).”

Distribution: Region 5. Known at type locality in

Venezuela, and an additional locality on the Brazilian

side of Tapirapeco (Caramaschi and Niemeyer 2005a).

Selected references: Myers and Donnelly (1997);

Caramaschi and Niemeyer (2005a).

Anomaloglossus tepuyensis (La Marca 1997)*

Holotype: ULABG 2557.

Type locality: “Entre Danto y Pifion (= Pefidn), a casi

una hora caminando desde Danto, en el trayecto desde

Kamarata hasta las laderas del Auyan-tepui, Estado

Bolivar, Venezuela, 1,650 m snm.”

Distribution: Region 5. Endemic from the slopes of

Auyan-tepui in Bolivar State. Probably extended to

neighboring tepuis.

Remarks: Myers and Donnelly (2008) comprehensively

redescribed this species. The Anomaloglossus

“tepuyensis’ cited from Guyana (Grant et al. 2006;

MacCulloch and Lathrop 2009), is actually a similar

species, A. megacephalus Kok, MacCulloch, Lathrop,

Billaert and Bossuyt, 2010. This taxon could be a

synonym of A. parkerae (P. Kok, pers. comm.).

Selected references: Grant et al. (1997, 2006); La

Marca (1997a); La Marca et al. (2002); Barrio-Amoros

and Brewer-Carias (2008); Myers and Donnelly (2008);

MacCulloch and Lathrop (2009); Sefiaris et al. (2014).

Anomaloglossus triunfo

(Barrio-Amoros, Fuentes and Rivas 2004)*

Holotype: EBRG 4756.

Type locality: “Summit of Cerro Santa Rosa, Serrania

del Supamo, 685 m above sea level. 6°40°39”N,

62°24’26” W, Estado Bolivar.”

Distribution: Region 5. Only known from two localities,

350-680 m asl, at the northwestern slopes of Sierra de

Lema, Bolivar State.

Amphib. Reptile Conserv.

Remarks: This name could be a synonym of A. parkerae

(P. Kok, pers. comm. ).

Selected references: Barrio-Amoros et al. (2004,

2011b); Myers and Donnelly (2008); Kok et al. (2013).

Anomaloglossus verbeeksnyderorum

Barrio-Amoros, Santos and Jovanovic, 2010*

Holotype: MHNLS 19649.

Type locality: “Tobogan de la Selva, Municipio Atures,

Estado Amazonas, Venezuela, 5°23’N, 67°34’W, 56 m

asl., 5.4109°N, 67.6197°W.”

Distribution: Region 5. Tobogan de la Selva and

nearby granitic domes around Puerto Ayacucho area in

NW Amazonas State; also at Serrania de los Pijiguaos,

Bolivar State (Camargo et al. 2014).

Remarks: Kok et al. (2012) showed that Anomaloglossus

verbeeksnyderorum and A. wothuja are not genetically

differentiated. There are subtle morphological differences

among these species (noted in Barrio-Amoros et al.

2011), but more detailed work is necessary to assess its

proper identity.

Selected references: Barrio-Amoros et al. (2010);

Camargo et al. (2014).

Anomaloglossus wothuja

(Barrio-Amoros, Fuentes and Rivas 2004)*

Holotype: MBUCV 6689.

Type locality: “Base of Cerro Sipapo, Tobogan del Cuao,

150 m above sea level, 5°05’09”’N, 67°27’07” W, Estado

Amazonas, Venezuela.”

Distribution: Region 5. Apparently endemic from

the uplands of the Cuao Massif; probably occurring

more extensively throughout similar granitic areas in

Amazonas State.

Remarks: Frost (2016) erroneously used acronym

EBRG (instead of MBUCV) for the catalog number

of the holotype of Anomaloglossus wothuja. See other

remarks for this species under A. verbeeksnyderorum.

Selected references: Barrio-Amoros et al. (2004); Frost

(2016).

Genus Aromobates Myers, Paolillo and Daly, 1991

Type species: Aromobates nocturnus Myers, Paolillo

and Daly, 1991 by original designation.

Remarks: Aromobates was known from a single species,

A. nocturnus until Grant et al. (2006) moved species in

the Colostethus alboguttatus group (sensu Rivero 1988)

and Nephelobates (sensu La Marca 1994) to Aromobates,

increasing the number to the current 18 species. The

current authors recover A. inflexus from synonymy (see

below in its own account). More species await description

from the Venezuelan Andes and beyond. Barrio-Amoros

and Santos (2012) provided a phylogeny and a list with

current threats for all species of the genus (except A.

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Catalogue of the amphibians of Venezuela

inflexus).

Selected References: Myers et al. (1991); La Marca

(1994); Grant et al. (2006, 2017); Barrio-Amoros et

al. (2011); Barrio-Amoros and Santos (2012); Barrio-

Amoréos (2016).

Aromobates alboguttatus (Boulenger 1903)*

Holotype: BM 1947.2.13.88.

Type locality: “Mérida, Estado Mérida, Venezuela, a

1,600 m.”

Distribution: Region 1. Andes of Estado Mérida.

Remarks: The species appears to have vanished in

the wild (Barrio-Amoroés and Santos 2012). Rivero

(1982a) included Colostehus inflexus Rivero, 1978 in

the synonymy of C. alboguttatus. La Marca (1997)

recovered the species as Nephelobates inflexus, without

explanation. Barrio-Amoros and Santos (2012) consider

this species different from A. alboguttatus and valid, but

awaiting proper attention and a redescription.

Selected references: Boulenger (1903); Ginés (1959);

Rivero (1961, 1963b, 1978, 1988); Péfaur (1985); Myers

et al. (1991); La Marca (1992, 1994); Pifiero and La

Marca (1996); Mijares and La Marca (1997); Barrio

and Fuentes (1999a); Barrio-Amoros and Santos (2012);

Barrio-Amoros (2013).

Aromobates cannatellai: Barrio-Amoros and Santos, 2012*

Holotype: CVULA 8327.

Type locality: “Parque Cascada de la Escalera, at the

entrance of Mesa de Pérez, Municipio Uribante, Estado

Tachira, Venezuela (8.0031N, 71.7316W), elevation

1,140 m.”

Distribution: Region 1. Only known from its type locality.

Remarks: Some populations of Aromobates saltuensis

reported from Colombia should probably be compared

to A. cannatellai (Barrio-Amorés and Santos 2012).

Acevedo et al. (2018) report 4. cannatellai from Norte

de Santander, Colombia.

Selected references: Barrio-Amoros and Santos (2012).

Aromobates capurinensis (Péfaur 1993)*

Holotype: CVULA IV.1063.

Type locality: “Paramo El Molino, via Canagua, Sierra

Nevada, 2,420m, Distrito Arzobispo Chacon, Municipio

Libertad, Estado Mérida, Venezuela.”

Distribution: Region 1. Known only from type locality.

Has been not observed since the year of its collection.

Selected references: Péfaur (1993); Barrio-Amoros and

Santos (2012).

Aromobates duranti (Péfaur 1985)*

Holotype: CVULA IV-1608.

Type locality: “Paramo de La Culata, Distrito Libertador,

Amphib. Reptile Conserv.

Estado Mérida, Venezuela, 2,880 m.”

Distribution: Region 1. Surroundings of type locality, in

streams of paramo and subpdramo 2,600-3,000 m asl.

Selected references: Péfaur (1985); Rivero (1988);

Myers et al. (1991); Mijares and La Marca (1997);

Barrio-Amoros and Santos (2012).

Aromobates ericksonae

Barrio-Amoros and Santos, 2012*

Holotype: CVULA 8309.

Type locality: “Los Ranchos, Santa Cruz de Mora,

Estado Mérida, Venezuela (8.3989N, 71.6801W),

elevation 1,193 m.”

Distribution: Known from five localities on the western

versant of Cordillera de Mérida, at both sides of the

Chama River Valley, 676—1,193 m asl.

Selected references: Barrio-Amoros and Santos (2012).

Aromobates haydeeae (Rivero 1978)*

Holotype: UPRM 4706.

Type locality: “El Vivero, entre Paramo El Zumbador

y Mesa del Aura, 2,570 m, Estado Tachira, Venezuela.”

Distribution: Region 1. Cloud forests of the Andes of

Tachira state.

Remarks: No recent data available; this species could be

affected by declines (Barrio-Amoros and Santos 2012).

Selected references: Rivero (1978, 1988); Myers et al.

(1991); La Marca (1994); Mijares and La Marca (1997);

Barrio-Amoros and Santos (2012).

Aromobates inflexus (Rivero 1980)*

Holotype: UPRM 4696.

Type locality: “El Almogral, entre Boca de Monte y el

cruce La Grita-Bailadores, Carr. De Pregonero, Estado

Tachira, Venezuela, 3,075 m.”

Distribution: Region 1. Surroundings of type locality.

Remarks: Here recovered from synonymy with

Aromobates alboguttatus, based on the considerations by

Barrio-Amoros and Santos (2012). See comment under

A. alboguttatus. The relationships between it and A.

orostoma must be evaluated.

Selected references: Rivero (1978, 1980, 1988); Myers

et al. (1991); La Marca (1997); Barrio-Amoroés and

Santos (2012).

Aromobates leopardalis (Rivero 1978)*

Holotype: UPRM 5157.

Type locality: “Mucubaji, 3,300 m, Edo. Mérida,

Venezuela.”

Distribution: Region 1. Paramo and subpdramo habitat

in Sierra Nevada, Merida State, 2,400—3,300 m asl.

Remarks: Not seen in recent years, probably very

endangered or extinct. Barrio-Amoros and Santos (2012)

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Barrio-Amoros et al.

ee ae

72. Aromobates haydeeae. Paramo Zumbador, Tachira. Photo: 73. Aromobates mayorgai. Surroundings of Chorotal, Mérida.

Amelia Diaz de Pascual. Photo: Fernando Rojas-Runjaic.

— | L . tae Oe

74A. Aromobates meridensis. Altos de San Luis, La Azulita, 74B. Aromobates cf. meridensis. Way from Mérida to El Morro,

Merida. Photo: César Barrio-Amoros.

J ae a a. i. : - 5 ‘ a "

75. Aromobates ornatissimus. Aguas de Obispos, Trujillo. Photo: 76. Aromobates saltuensis. Coloncito, Tachira. Photo: César

César Barrio-Amoros. Barrio-Amoros.

Amphib. Reptile Conserv. 41 July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

state that 1995 was the last year it was seen.

Selected references: Rivero (1978, 1988); La Marca

(1991b); Miyjares-Urrutia (1991); Myers et al. (1991);

Barrio-Amoros (200Ic, 2013); Barrio-Amords and

Santos (2012).

Aromobates mayorgai (Rivero 1980)*

Holotype: UPRM 5160.

Type locality: “El Chorotal (El Sirenal), Carretera

Merida a La Azulita, 1,800 m, Edo. Mérida, Venezuela.”

Distribution: Region 1. Western Sierra de la Culata,

1,700—2,400 m asl.

Remarks: La Marca and Otero (2012) redescribed the

species.

Selected references: Rivero (1980, 1988); La Marca and

Mijares (1988); Myers et al. (1991); La Marca (1994);

Mijares and La Marca (1997); Barrio-Amoros (2001c);

Barrio-Amoros and Santos (2012); La Marca and Otero

(2012).

Aromobates meridensis (Dole and Durant 1972)*

Holotype: MBUCV 6168.

Type locality: “Chorotal, 15 km. al SE de La Azulita,

1,880 m, Estado Mérida, Venezuela.”

Distribution: Region 1. Southwestern Sierra de la

Culata, Mérida, 1,800 to 3,300 m asl. Recently reported

along the way from Merida to El Morro (Image 74B).

Remarks: This species has received some attention

(Barrio-Amoros et al. 2010; La Marca and Otero 2012).

The first authors reported a surviving population of

around 56 to 129 adults in 2006, and redescribed the

species from topotypic material. That population had

a high presence of fungal pathogen Batrachochytrium

dendrobatidis (Lampo et al. 2008; Barrio-Amoros

and Lampo 2009). La Marca and Otero (2012) also

redescribed the species based on the holotype and

clarified the confusion regarding the catalog data of the

type series and commented on its conservation status.

Selected references: Dole and Durant (1972); Rivero

(1988); La Marca (1991a “1994”): Myers et al. (1991);

Mijares and La Marca (1997); Barrio-Amoros (2001c);

Lampo et al. (2008); Barrio-Amoroés and Lampo (2009);

Barrio-Amoros et al. (2010g); Barrio-Amoros and Santos

(2012); La Marca and Otero (2012).

Aromobates molinarii (La Marca 1985)*

Holotype: CVULA 2820.

Type locality: “Las Playitas, 2,270 m, near Bailadores

(8°15’N, 71°50’ W), Estado Mérida, Venezuela.”

Distribution: Region 1. Surroundings of type locality

and road to Estanques, Merida State.

Selected references: La Marca (1985, 1991la “1994”);

Rivero (1988); Myers et al. (1991); Mijares and La Marca

(1997); Barrio and Fuentes (1999); Barrio-Amoros and

Amphib. Reptile Conserv.

Santos (2012).

Aromobates nocturnus Myers, Paolillo and Daly, 1991*

Holotype: AMNH 130005.

Type locality: “Cloud forest at 2,250 m elevation, about

2 km. Airline ESE Agua de Obispos, Estado Trujillo,

Venezuela (9°42’N, 70°05’ W).”

Distribution: Region 1. Known only from type locality.

Remarks: Apparently highly endangered, not found in

type locality since its original discovery. Several parties

have failed to find the species (La Marca 2005; Barrio-

Amoréos et al. 2011). Considered as Critically Endangered

(CR) by the current Venezuelan Red Book (La Marca

2015a) but could be extinct.

Selected references: Myers et al. (1991); Barrio and

Fuentes (1999a); Barrio-Amoros (2001c, 2013, 2016);

Grant et al. (2006); Barrio-Amoros and Santos (2012);

La Marca (2015a).

Aromobates ornatissimus

Barrio-Amoros, Rivero and Santos, 2011*

Holotype: EBRG 5292

Type locality: Las Palmas, Municipio Carache, Estado

Trujillo, Venezuela, 09°41°47”N, 70°08’24” W (9.6964

N-70.1400 W)); elevation 2,350 m

Distribution: Region 1. Apparently endemic from type

locality and surroundings.

Remarks: In the phylogenies recovered by Grant et al.

(2006; as Nephelobates sp. 1321 and ULABG 4445) and

Barrio-Amoros and Santos (2012) this species appears as

the sister taxon of all other Aromobates.

Selected references: Barrio-Amoros et al. (2011a);

Barrio-Amoros and Santos (2012).

Aromobates orostoma (Rivero 1978)*

Holotype: UPRM 4509.

Type locality: “Boca del Monte, Camino del Pregonero,

2,615 m, Estado Tachira, Venezuela.”

Distribution: Region 1. Known from the type locality.

Remarks: The species needs taxonomic attention.

Selected references: Rivero (1978, 1988); Péfaur

(1985); La Marca (1991a “1994”); Mijares and La Marca

(1997); Barrio-Amoros and Santos (2012).

Aromobates saltuensis (Rivero 1978)*

Holotype: UPRM 5147.

Type locality: “de la Fria a Michelena, Edo. Tachira,

Venezuela, 830 m.”

Distribution: Region 1. The species is known from

the southwestern extreme of the Cordillera de Mérida

and northeastern Cordillera Oriental de Colombia in

its Venezuelan side. Mentioned from Colombia by

Grant et al. (2006) and Anganoy-Criollo (2012) from

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Barrio-Amoros et al.

as yep e serranus. El Morro, Vien Photo: Jaime

Péfaur.

79. Aromobates zippeli. Surroundings of Mucuchies, Mérida.

Photo: Fernando Rojas-Runjaic.

81. Mannophryne aff. caquetio. Sierra de San Luis, Falcon.

Photo: César Barrio-Amoros.

Departamentos of Norte de Santander, Boyaca and Cesar,

respectively.

Remarks: Redescribed by Barrio-Amorés and Santos

(2012). Data presented by Barrio-Amoroés and Santos

(2012) show Colombian populations are more closely

related with Aromobates cannatellai than with A.

Amphib. Reptile Conserv.

5 wy

a. ne

78. Womb: tokuko. Female ane MHNLS 18490. 0. Ipika,

rio Tokuko, Sierra de Perija, Zulia. Photo: Edwin Infante.

stvd sami

80. Waa: sp. l. Calderas, Barinas. Photo: César Barrio-

Amoros.

82A. Mannophryne collaris. Male. Estanquillo, Chama river

valley, Mérida. Photo: César Barrio-Amoros.

saltuensis, or could represent new species. See Barrio-

Amoros and Santos (2012) for detailed account of the

Colombian records of A. cf. saltuensis.

Selected references: Rivero (1978, 1988); Péfaur

(1985); Myers et al. (1991); Grant et al. (2006, 2017);

Barrio-Amoros and Santos (2012).

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Catalogue of the amphibians of Venezuela

Aromobates serranus (Péfaur 1985)*

Holotype: CVULA: IV-2847.

Type locality: “El Morro, Distrito Libertador, Estado

Merida, Venezuela, 2,300 m.”

Distribution: Region 1. Surroundings of type locality.

Remarks: Not seen or reported in a long time (Barrio-

Amoros and Santos 2012). Very likely extinct.

Selected references: Péfaur (1985); Rivero (1988);

Myers et al. (1991); Mijares and La Marca (1997):

Barrio-Amoros and Santos (2012).

Aromobates tokuko

Rojas-Runjaic, Infante and Barrio-Amoros, 2011*

Holotype: MHNLS 18479.

Type locality: “Surroundings of Ipika, Yukpa indigenous

community, Rio Tokuko Basin, Municipio Machiques

de Perija, Sierra de Perijé, Estado Zulia, Venezuela

(09°52’N, 72°51’ W; elevation 595 m).”

Distribution: Region |. Known from four nearby localities

in the central-eastern sector of the Sierra de Periya.

Remarks: Only member of this genus present in Periya.

More species are expected in such unexplored Sierra.

Considered as Critically Endangered by the Venezuelan

Red Book (Rojas-Runjaic and Sefiaris 2015c).

Selected references: Rojas-Runjaic et al. (2011); Barrio-

Amoros and Santos (2012); Rojas-Runjaic and Sefiaris

(2015c).

Aromobates walterarpi La Marca and Otero, 2012*

Holotype: ULABG 2087.

Type locality: “Stream at about 500 m away from ‘Plaza

Bolivar’ of Pifiango, close to the cemetery, on the road

from Pifiango to Pico El Aguila 2,325 m (9°01°59.8”N,

70°53’02.5” W), Estado Mérida, Venezuela.”

Distribution: Region 1. Known only from type locality.

Selected references: La Marca and Otero (2012).

Aromobates zippeli Barrio-Amoros and Santos, 2011*

Holotype: CVULA 8329.

Type locality: “Surroundings of Mucuchies, Estado

Merida, Venezuela, 8.75N, 70.8833W, elevation 2,970

m.”

Distribution: Region 1.

surroundings of type locality.

Selected references: Barrio-Amoros and Santos (2012).

Apparently restricted to

Genus Mannophryne La Marca, 1992

Type species: Colostethus yustizi La Marca, 1989 by

original designation.

Mannophryne caquetio Mijares and Arends, 1999*

Amphib. Reptile Conserv.

Holotype: EBRG 3570.

Type locality: “Toma de agua de Maparari, Municipio

Federacion, Sierra de Churuguara, Estado Falcon,

Venezuela (aprox. 10° 47’N, 69° 25’W), 800 m.”

Distribution: Region 2. Apparently endemic from Sierra

de Churuguara, Estado Falcon.

Selected references: Mijares and Arends (1999b);

Barrio-Amoros et al. (2010b); Mijares-Urrutia and La

Marca (2015a).

Mannophryne collaris (Boulenger 1912)*

Lectotype: BM 1947.2.14.42.

Type locality: “Mérida, 5,200 feet, and Rio Albireggas

(Albarregas), 11,300 feet, Venezuela.”

Distribution: Region 1. Once believed widespread along

the Venezuelan Andes. Today restricted to the Chama

River Valley and its tributaries (rio Mocoties), between

195—1,900 m by Barrio-Amoros et al. (2010b).

Remarks: This species is vicariant of Mannophryne

urticans (Barrio-Amoros et al. 2010b).

Selected references: Boulenger (1912); Ginés (1959);

Rohl (1959); Rivero (1961, 1963b, 1988); Dole and

Durant (1974b); Durant and Dole (1975); Péfaur and

Diaz De Pascual (1982, 1987); Péfaur (1987); La Marca

(1991c “1994,” 1992, 1994a, 1995a,b); Myers et al.

(1991); Barrio (1996a); Barrio and Fuentes (1999a);

Barrio-Amoros et al. (2010b); Barrio-Amoros (2013).

Mannophryne cordilleriana La Marca, 1994*

Holotype: ULABG 763.

Type locality: “Presa Hidraulica Jose Antonio Paez,

1,600 m, near La Mitusus on road Santo Domingo-

Barinas, Estado Mérida, Venezuela.”

Distribution: Region 1. Once believed to only inhabit

the type locality and its surroundings, recent evidence

(Barrio-Amoros et al. 2010b) shows that it 1s widespread

along the Eastern versant of the Cordillera de Mérida in

states Mérida and Barinas (Barrio-Amoros 2010), 200 to

at least 1,950 m asl, and still quite abundant.

Remarks: This is the vicariant species of Mannophryne

orellana from Tachira state (J.C. Santos and CBA,

unpub. data).

Selected references: La Marca (1994a, 1995b); Barrio-

Amoros (2010a); Barrio-Amoros et al. (2010b).

Mannophryne herminae (Boettger 1893)*

Syntypes: SMF. Presumably SMF 7286 is a Lectotype

by designation, fide Edwards, 1974.

Type locality: “Puerto Cabello in Venezuela.”

Distribution: Region 2. Central Cordillera de la Costa

(to be better determined).

Remarks: The name herminae has long been

associated with the abundant populations found in

the Cordillera de la Costa. Unpublished genetic data

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

fa, ; <b ‘e* a

82B. Mannophryne collaris. Female. Estanquillo, Chama river 83A. Mannophryne cordilleriana. Female. Acequias, Barinas.

valley, Mérida. Photo: César Barrio-Amoros. Photo: César Barrio-Amoros.

* . % =

83B. Mannophryne cordilleriana. Female. Calderas, Barinas. 84A. Mannophryne sp. | aff. herminae. Female. Rancho Grande,

Photo: César Barrio-Amoros. Henri Pittier National Park, Aragua. Photo: César Barrio-Amoros.

84B. Mannophryne sp. | aff. herminae. Female. Rancho Grande, 85A. Mannophryne lamarcai. Male. Cerro Socopo, Falcon. Photo:

Henri Pittier National Park, Aragua. Photo: César Barrio-Amoros. Arlene Cardozo.

Amphib. Reptile Conserv. July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

(by J.C. Santos) indicates some of these populations

belong to an undescribed species. Mannophryne

herminae sensu stricto apparently is found only in San

Esteban (Carabobo state) and surrounding areas, near

Puerto Cabello. The identity of other populations from

Cordillera de la Costa traditionally determined as M

herminae need confirmation. Rojas-Runjaic et al. (2018)

redefined M. herminae and restricts Mannophryne

herminae sensu Stricto to the northern slope of Cordillera

de la Costa between Carabobo and Aragua states. In the

Mamnnophryne collaris species group (Manzanilla et al.

2007; Grant et al. 2017).

Selected references: Boettger (1893); Stejneger (1901);

Lutz (1927); Schmidt (1932); Aleman (1952); Mertens

(1957b); Ginés (1959); Sexton (1960); Rivero (1961,

1964a, 1988); Test (1962); Gremone et al. (1986); La

Marca (1991c “1994,” 1992, 1994a, 1995b); Myers

et al. (1991); Manzanilla et al. (1995); Barrio-Amoroés

(2006c); Barrio-Amoros et al. (2010b); Rojas-Runjaic et

al. (2018).

Mannophryne lamarcai Mijares and Arends, 1999*

Holotype: EBRG 3281.

Type locality: “Cerro Sopocdé, 30 km SW de Guajiro,

Municipio Mauroa, Estado Falcon, Venezuela (10° 28’N,

70° 48’W), 1,250 m.”

Distribution: Region 2. Only known from type locality

and four additional localities at Serrania de Ziruma

(Moran et al. 2016).

Selected references: Barrio and Fuentes (1999a);

Mijares and Arends (1999a); Mijares-Urrutia and La

Marca (2015b), Moran et al. (2016).

Mannophryne larandina (Yustiz 1991)*

Holotype: UCLA 0087.

Type locality: “Hato Arriba, Distrito Moran, Sierra de

Barbacoas, 1,800 m snm.” Estado Lara, Venezuela.

Distribution: Region 1. Surroundings of type locality.

Remarks: Based on genetic and morphological evidence

this species is very similar of Mannophryne yustizi and

could be a synonym of it.

Selected references: Yustiz (1991); Mijares and Arends

(1999).

Mannophryne leonardoi Manzanilla, La Marca,

Jowers, Sanchez, and Garcia-Paris, 2007*

Holotype: EBRG 4899

Type locality: “Caserio El Toyano, Parroquia Pozuelos,

Municipio Sotillo, Estado Anzoategui, Venezuela,

10°7°30.7”N, 64°29°35.7” W, 875 m”.

Distribution: Region 2. Distributed along the Turimiquire

massif, among Anzoategui, Sucre and Monagas states.

Remarks: Considered as Endangered by the current

Venezuelan Red Book (Rojas-Runjaic and Sefiaris

Amphib. Reptile Conserv.

2015d).

Selected references: Yustiz (1991); Mijares and Arends

(1999); Manzanilla et al. (2007b); Barrio-Amoros et al.

(2010b); Rojas-Runjaic and Sefiaris (2015d).

Mannophryne molinai

Rojas-Runjaic, Matta-Pereira and La Marca, 2018*

Holotype: MHNLS 21355.

Type locality: “Quebrada La Rondona, Sierra de

Aroa, Sucre municipality, Yaracuy state, Venezuela

(10°19’20.8”N, 68°52’24.0”W; 1,180 m asl).”

Distribution: Endemic of the Sierra de Aroa and only

known from type locality at the southeastern slope of

this mountain chain. Other populations known from

northeastern and western foothills of Sierra de Aroa

should be evaluated to determine if they correspond to

this new species.

Remarks: Apparently pertaining to the species group of

Mamnnophryne collaris (sensu Manzanilla et al. 2007, and

Grant et al. 2017) and closely related to M. herminae,

though easily distinguished by the advertisement call

(Rojas-Runjaic et al. 2018). Its phylogenetic position

and relationships into the genus remain unevaluated.

Conservation status has not been evaluated by the IUCN,

but based on restricted distribution and loss of habitat

Rojas-Runyaic et al. (2018) proposed to classify it as VU

(vulnerable).

Selected references: Grant et al. (2017), Rojas-Runjaic

et al. (2018).

Mannophryne neblina (Test 1956)*

Holotype: UMMZ 113001.

Type locality: “Portachuelo Pass, Rancho Grande,

Estado Aragua, Venezuela.”

Distribution: Region 2. Known only from type locality

and surroundings, 900—1,100 m asl.

Remarks: Despite numerous searches in the type

locality and surroundings during recent decades, no other

specimens have been found since its description. At least

a local extinction is presumed. Lotzkat (2007) apparently

found M. neblina at Macizo de Nirgua (Yaracuy State).

However, vouchers are not mentioned and the specimens

were identified by J. Manzanilla by checking photos.

Considered as Critically Endangered (CR) by the current

Venezuelan Red Book (La Marca 201 5b).

Selected references: Ginés (1959); Rivero (1961, 1964a,

1988); La Marca (1991c “1994,” 1994a, 1995b, 2015b);

Myers et al. (1991); Manzanilla et al. (1995); Rodriguez

and Rojas-Suarez (2008); Barrio-Amoros et al. (2010b).

Mannophryne oblitterata (Rivero 1984)*

Holotype: UPR-M 3492.

Type locality: “Carretera de Sta. Teresa a Higuerote, 10

km hacia abajo del cruce Santa Teresa-Altagracia, 150 m,

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Barrio-Amoros et al.

Falcon.

85B. Mannophryne lamarcai. Female. Cerro Socopo,

Photo: Arlene Cardozo.

‘ar ts : ee a

87. Mannophryne cf. neblina. Cerro Zapatero, Nirgua Massif,

ie? Pepa = 4

es. |

aia Tas

88B. Mannophryne oblitterata. Female. Guatopo National Park,

Guarico. Photo: Javier Mesa.

Edo. Miranda, Venezuela.”

Distribution: Region 2. Endemic to Guatopo, in the

Interior Coastal Range.

Remarks: The description of Co/ostethus guatopoensis

Dixon and Rivero-Blanco, 1985 (a current synonym of

M. oblitterata fide Rivero, 1988), and the redescription

of La Marca (1994a) are more comprehensive than

Rivero’s description.

Selected references: Rivero (1984, 1988); Dixon and

Amphib. Reptile Conserv.

86. Mannophryne leonardoi. Cueva del Guacharo, Monagas.

Photo: César Barrio-Amoros.

w ing Ty

baie ai: Se ee ef

88A. Mannophryne oblitterata. Male. Guatopo National Park,

Guarico. Photo: César Barrio-Amoros.

: ; Stee x —

+ ~ a Bie ad ver

oe,

89. Mannophryne orellana. Female. Pregonero-La Trampa road,

Tachira. Photo: César Barrio-Amoros.

Rivero-Blanco (1985); La Marca (1991c “1994,” 1992,

1994a, 1995b); Myers et al. (1991); Barrio-Amoréos et al.

(2010b).

Mannophryne orellana

Barrio-Amoros, Molina and Santos, 2010*

Holotype: CVULA 7165.

Type locality: “Road from Pregonero to La Trampa,

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Catalogue of the amphibians of Venezuela

Estado Tachira,

elevation 1,192 m.”

Distribution: Region |. Southwestern edge of Cordillera

de Merida in Uribante Valley, and northeastern side of

Cordillera Oriental de Colombia in its Venezuelan side.

Probably present in Colombia.

Remarks: Vicariant of Mannophryne_ cordilleriana

(Barrio-Amoros et al. 2010b).

Selected references: Barrio-Amoros et al. (2010b).

Venezuela (O8°01’N, 71°43’W),

Mannophryne riveroi

(Donoso-Barros 1964) *

Holotype: MZUC 8566 (Coleccion de Donoso Barros

V-307; see Barrio-Amoros and Ortiz 2015).

Type locality: “Cerro Azul, Macuro,” Estado Sucre,

Venezuela.

Distribution: Region 2. Endemic of Peninsula de Paria,

Sucre state.

Remarks: Barrio-Amor6s et al. (2010b) reported natural

history, variation, and malformations. Barrio-Amoros and

Ortiz (2015) assign the proper number to the holotype in

the collection of the Universidad de Concepcion in Chile.

Considered as Endangered by the current Venezuelan

Red Book (Rojas-Runjaic and Sefiaris 2015e).

Selected references: Donoso-Barros (1964); Rivero

(1967c, 1968a, 1988); La Marca (1991c “1994,”

1994a, 1995b); Myers et al. (1991); Barrio-Amoros et

al. (2010b,c); Barrio-Amoros and Ortiz (2015); Rojas-

Runjaic and Sefiaris (2015e).

Mannophryne speeri

La Marca, 2009*

Holotype: ULABG 5393.

Type locality: “Approximately 1 km NNE from

Laguneta, 960 m asl., 65.9 km on the road from the

crossroads Guanare-Suruguapo heading to Villanueva,

Sierra de Portuguesa, Municipio Moran, Estado Lara,

Venezuela.”

Distribution: Region 1. Currently known only from type

locality and nearby.

Selected references: La Marca (2009); Barrio-Amoros

et al. (2010b).

Mannophryne trujillensis

Vargas and La Marca, 2007*

Holotype: ULABG 1160.

Type locality: “Paseo Los Ilustres, Quebrada Los

Cedros, 840 m elevation, 9°21°46.0’N, 70°26'41.8” W,

Trujillo, Estado Trujillo, Venezuela.”

Distribution: Region 1. Currently known only from type

locality and nearby.

Selected references: Vargas and La Marca (2007);

Barrio-Amoros et al. (2010b).

Amphib. Reptile Conserv.

Mannophryne urticans

Barrio-Amoros, Molina and Santos, 2010*

Holotype: CVULA 7224.

Type locality: Rio Frio, northwestern slope of the

Cordillera de Mérida, Estado Mérida, Venezuela

(08°51°N, 71°17’W), 676 m asl.

Distribution: Region 1. Known only from type locality,

possibly more widespread along similar habitats in the

western piedmont of the Cordillera de Mérida.

Remarks: Vicariant of Mannophryne collaris (Barrio-

Amoros et al. 2010b).

Selected references: Barrio-Amoros et al. (201 0b).

Mannophryne yustizi (La Marca 1989)*

Holotype: CVULA: IV-2842.

Type locality: “14 km. SSE Sanare, on stream along road

Sanare-Parque Nacional Yacambu, 1,475 m, (ca. 9°43’N,

69°39’°W), Distrito Jiménez, Estado Lara, Venezuela.”

Distribution: Region |. Extended throughout the eastern

versant of Cordillera de Mérida in Lara state.

Selected references: La Marca (1989, 1992, 1994a,

1995b); Myers et al. (1991); Yustiz (1996); Barrio-

Amoros et al. (2010b).

Mannophryne venezuelensis

Manzanilla, Jowers, La Marca and Garcia-Paris, 2007*

Holotype: EBRG 4924.

Type locality: “From approximately 4.0 km east of San

Juan de Las Galdonas, Municipio Arismendi, Estado

Sucre, Venezuela, 10°43’N, 62°48’ W, altitude 180 m.”

Distribution: Region 2. Distributed along the Peninsula

de Paria, 0—1,000 m asl.

Remarks: Many populations of Mannophryne from

Coastal Range were previously assigned to Mannophryne

trinitatis (including M. venezuelensis) until Barrio-

Amoros et al. (2006b) based on previously published

evidence (Rivero 1961; La Marca 1994; Kaiser et al.

2003) restricted its distribution to Trinidad. First species

of the genus defined based on an integrative approach

with morphological, genetic, and bioacoustic evidence

(Manzanilla et al. 2007b).

Selected references: Barrio-Amoros et al. (2006b;

2010b); Manzanilla et al. (2007a, 2009).

Mannophryne vulcano

Barrio-Amoros, Molina and Santos, 2010*

Holotype: CVULA 7170

Type locality: Northern slope of Cerro El Volcan, Baruta,

Estado Miranda, Venezuela, collected by Charles Brewer

on 30 June, 2007 (10°25’N, 66°51’ W), 1,064 m asl.

Distribution: Region 2. Valley of Caracas.

Remarks: Previous mentions of Mannophryne trinitatis

and M. herminae from area around Caracas should be

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Barrio-Amoros et al.

90A. Mannophryne riveroi. Male. Southern slopes Cerro Humo, 90B. Mannophryne riveroi. Female. Southern slopes Cerro Humo,

Peninsula de Paria, Sucre. Photo: César Barrio-Amoros. Peninsula de Paria, Sucre. Photo: Hinrich Kaiser.

91A. Mannophryne speeri. Male. Chavasquén-Guarico road, 91B. Mannophryne speeri. Female. Chavasquén-Guarico

Portuguesa. Photo: César Barrio-Amoros. Portuguesa. Photo: César Barrio-Amoros.

92. Mannophryne aff. trujillensis. Monte Carmelo, Trujillo. Photo: 93A. Mannophryne urticans. Male. Rio Frio, Mérida. Photo:

Juan Pablo Diasparra. César Barrio-Amoros.

Amphib. Reptile Conserv. July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

Nee =~ .

. *

. 2

. ~!

93B. Mannophryne urticans. Female. Rio Frio, Mérida. Photo:

César Barrio-Amoros.

95. Mannophryne vulcano. Male. Cota Mil, Caracas. Photo: César

Barrio-Amoros.

4 i} ‘aa

97. Mannophryne sp. 1. aff. herminae. Male. Casa Maria, Bejuma,

Carabobo. Photo: César Barrio-Amoros.

referred to M. vulcano (Barrio-Amoros et al. 201 0b).

Selected references: Barrio-Amoros et al. (2006b;

2010b).

Subfamily Dendrobatinae Cope, 1865

Genus Ameerega Bauer, 1986

Type species: Hyla trivittata Spix, 1824 by original

designation.

Amphib. Reptile Conserv.

ie. ® J = + 5

awe os. aae

—— 7°

ice, Oa

et eta Ce at “S oe ee ,

ge eS ee ee eee |

Ee eae So a ee es ae

94. Mannophryne venezuelensis. Female. Road to Las Melenas,

Peninsula de Paria, Sucre. Photo: César Barrio-Amoros.

ara

a =

96. Mannophryne ustizi Female. Yacambu National Park, Lara

Photo: César Barrio-Amoros.

Guatopo National Park, Guarico. Photo:

“ 4

98. Mannophryne sp. 2.

César Barrio-Amoros.

Ameerega picta (Tschudi 1838)

Holotype: MNHNP 4910

Type locality: Santa Cruz, Bolivia.

Distribution: Region 5. Widely distributed in Amazon

Region lowlands, through Colombia, Peru, Bolivia,

Paraguay, Brazil, and Venezuela. In Venezuela, only

known from Northeastern Bolivar and adjacent Delta

Amacuro States.

Remarks: Barrio-Amoros (2004) treated the Guayanan

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

populations of this taxon as Epipedobates guayanensis,

following Schulte (1999) without arguing about the

change. Ameerega guayanensis (Heatwole, Solano and

Heatwole 1965; type locality: forest between Rancho

Alegre and base of Altiplanicie (de Nuria), on trail to

Quebrada Cabeza de Burro, 5 km east of Las Chicharras,

47 km north of Tumeremo; Altiplanicie de Nuria,

100-250 m, Estado Bolivar, Venezuela) would be the

proper name of Guianan populations if demonstrated

to be genetically different from A. picta from the type

locality. The southwestern Amazonian population is

separated from the Guiana population by at least 3,000

km in a straight line (Lotters et al. 2007). However, J.C.

Santos (unpub. data) preliminary results do not indicate

a clear genetic separation and thus, until more data are

published, we consider guayanensis under picta. As we

do not use subspecies, the name must be regarded in its

synonymy. See also comment by Avila Pires et al. (2010).

Selected references: Heatwole et al. (1965); Silverstone

(1976); Gremone et al. (1986); Walsh (1994); Duellman

(1997); Gorzula and Sefiaris (1998); Barrio and Fuentes

(1999a); Schulte (1999); Sefiaris and Ayarzagtiena (2004

2002”); Lotters et al. (2007); Barrio-Amoros (2016).

Ameerega trivittata (Spix 1824)

Syntypes: ZSM 43/0; RMNH 1836 (lectotipo).

Type locality: “Iuxta flumen Teffé,” Brazil.

Distribution: Region 5. Widely distributed in Amazon

Region lowlands, through Colombia, Peru, Bolivia,

Brazil, Venezuela, Guyana, and Suriname. In Venezuela,

known only from northeastern Bolivar State and adjacent

Delta Amacuro.

Selected references: Zimmermann and Zimmermann

(1988); Gorzula (1991); Walsh (1994); Gorzula and

Sefiaris (1998); Barrio and Fuentes (1999a); Sefiaris and

Ayarzagtiena (2004 “2002”); Grant et al. (2006); Lotters

et al. (2007); Barrio-Amoros (2016).

Genus Dendrobates Wagler, 1830

Type species: Rana tinctoria Cuvier, 1797 by subsequent

designation by Dumeril and Bibron, 1841.

Dendrobates leucomelas Steindachner, 1864

Holotype: NHMW 19188.

Type locality: “Colombia.”

Distribution: Regions 4, 5. Widespread south of the

Orinoco (see map in Barrio and Fuentes (1999), Lotters

et al. (2007), and Molina-Rodriguez and Kahn (2016)).

Much more common in Venezuela south of the Orinoco

(in Amazonas, Bolivar, and Delta Amacuro States), also

present but scattered in southeastern Colombia, northern

Brazil, and western Guyana.

Remarks: Different morphs recognized, but so far, not

of taxonomic importance. The “sapito minero” has a

Amphib. Reptile Conserv.

key importance in some ethnic mythologies like the

Ye’ kwana (Barrio-Amoros and Brewer-Carias 2008).

Selected references: Steindachner (1864); Ginés

(1959); Rivero (1961, 1964b,d, 1967a); Heatwole et al.

(1965); Silverstone (1975); Paolillo (1977); Hoogmoed

and Gorzula (1979); Gremone et al. (1986); Rivero et

al. (1986); Walsh (1994); Rodriguez and Rojas-Suarez

(1995); Duellman (1997); Fuentes and Rodriguez-Acosta

(1997); Barrio and Fuentes (1998, 1999a); Gorzula and

Sefiaris (1998); Sefiaris and Ayarzagtiena (2004 “2002”);

Grant et al. (2006); Lotters et al. (2007); Barrio-Amoroés

and Brewer-Carias (2008); Barrio-Amoros et al. (2011b),;

Sefiaris et al. (2014); Molina-Rodriguez and Kahn (2016).

Genus Minyobates Myers, 1987

Type species: Dendrobates steyermarki Rivero, 1971 by

original designation.

Minyobates steyermarki (Rivero 1971)*

Holotype: UPR-M 3399.

Type locality: “Cerro Yapacana, 1,200 m Territorio

Federal Amazonas, Venezuela.”

Distribution: Region 5. Endemic to Yapacana, a tepul in

Amazonas State.

Remarks: See considerations about conservation in

Barrio-Amorés and Fuentes (1999a), Rodriguez and

Rojas-Suarez (2009), Barrio-Amoros and Torres (2010),

and La Marca (2016). Accounts on morphology and natural

history by Lotters et al. (2007) and La Marca (2016).

Selected references: Rivero (197 1c); Silverstone (1975);

Gremone et al. (1986); Myers (1987); Walsh (1994);

Rodriguez and Rojas-Suarez (1995); Gorzula and Sefiaris

(1998); Barrio and Fuentes (1999a); Vences et al. (2000,

2003); Grant et al. (2006); Lotters et al. (2007); Barrio-

Amoros and Torres (2010); Barrio-Amoros (2016); La

Marca (2016).

Family Eleutherodactylidae Lutz, 1954

Subfamily Eleutherodactylinae Lutz, 1954

Genus Adelophryne Hoogmoed and Lescure, 1984

Type species: Adelophryne adiastola Hoogmoed and

Lescure, 1984, by original designation.

Adelophryne gutturosa Hoogmoed and Lescure, 1984

Type: BM 1983.1139.

Type locality: “Between camp IV and V, northern slopes

of Mount Roraima, Guyana (60°46’W, 5°17’N) 3,000

feet (914 m).”

Distribution: Region 5. Known from a few localities in

Sierra de Lema and Gran Sabana, eastern Venezuela.

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Catalogue of the amphibians of Venezuela

99. Pee picta. Sita ree wee Amacuro. Photo: oon 100. ees trivittata. Siete Imataca, Delta are Photo:

Barrio-Amoros. César Barrio-Amoros.

101A. DERG BAIS Teicomslas Beds me la iteer! Sierra de 101B. Dendrobates iencanislas Serrania del Cuao, Manneonee

Lema, Bolivar. Photo: César Barrio-Amoros. Photo: Oswaldo Fuentes Ramos.

102. Minyobates steyermarki. Yapacana summit, Amazonas. 103. Eleutherodactylus Aiton ‘Ciudad Bolivar, Bolivar.

Photo: Wolfgang Schmidt. Photo: César Barrio-Amoros.

Selected references: Ayarzagtiena and Diego-Aransay Genus Eleutherodactylus Dumeril and Bibron, 1841.

(1985); Duellman (1997); MacCulloch et al. (2008);

Barrio-Amoros and Duellman (2009); Sefiaris et al. |§ Type species: Hy/odes martinicensis Tschudi, 1838, by

(2014). monotypy.

Remarks: After taxonomic rearrangement of Terraranans

Amphib. Reptile Conserv. 52 July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

based on molecular phylogeny by Heinicke et al. (2007)

Eleutherodactylus was restricted to the Caribbean clade

of direct developing frogs. The only E/eutherodactylus in

Venezuela is an introduced but well established species.

Eleutherodactylus johnstonei Barbour, 1914

Syntypes: MCZ 2759 (two specimens).

Type locality: St. George Parish, Grenada.

Distribution: Region 2. Introduced populations in

different cities with variable success. Likely originally

from Sta. Lucia or the Antigua and Barbuda bank (Frost

2016). Currently established in most of Lesser Antilles,

several countries in Central America and northern South

America (Panama, Colombia, Venezuela, Guyana,

Suriname, and French Guiana). In Venezuela, established

populations reported in several cities and towns (e.g.,

Caracas, Cumana, Irapa, Maracay, Valencia). See a

resumé in Kaiser et al. (2002). Despite being introduced,

not reported to affect any autochthonous species, and

only known from anthropogenic sites like gardens and

parks.

Selected references: Hardy and Harris (1979); Rada

(1981b); Gorzula (1989); La Marca (1992); Gorzula and

Sefiaris (1998); Kaiser et al. (2002); Rojas-Runyaic et al.

(2007); Barrio-Amoros (2013).

Family Hemiphractidae Peters, 1862

Remarks: Long considered a subfamily of Hylidae.

Darst and Cannatella (2004) suggested this group is not

monophyletic nor related to Hylidae. For an account

of subsequent changes in phylogenetic relationships

and taxonomy see Frost (2018). See Duellman (2015)

for a thorough review of genera and species included.

Recently Castroviejo et al. (2015) inferred a new family

phylogeny based on total evidence, recovering it (all six

genera) as monophyletic, and sister of Athesphatanura.

Several species of Stefania await description. In

Peninsula de Paria, Sucre State, a Gastrotheca, possibly

new to science, has been heard calling in the canopy

during rainy days and nights.

Subfamily Cryptobatrachinae Frost, Grant,

Faivovich, Bain, Haas, Haddad, de Sa, Channing,

Wilkinson, Donnellan, Raxworthy, Campbell,

Blotto, Moler, Drewes, Nussbaum, Lynch, Green and

Wheeler, 2006

Genus Cryptobatrachus Ruthven, 1916

Type species: Cryptobatrachus boulengeri Ruthven,

1916, by original designation.

Remarks: Cryptobatrachus nicefori Cochran and Goin,

1970 from Colombia was excluded from the genus by

Infante-Rivero et al. (2009), despite Lynch’s (2008)

Amphib. Reptile Conserv.

contrary opinion, and placed into Hyla incertae sedis.

In their recent phylogenetic analysis Castroviejo et al.

(2015) inferred Cryptobatrachus as sister of Flectonotus

and included these two genera in Cryptobatrachinae.

Cryptobatrachus remotus \nfante-Rivero, Rojas-

Runjaic and Barrio-Amoros, 2009

Holotype: MHNLS 17661.

Type locality: “Fundo ‘El Progreso,’ cuenca alta del

rio Socuy, sierra de Perijaé, Municipio Jests Enrique

Losada, Estado Zulia, Venezuela (10°43’13,30°N,

72°29’ 16,60”O; + 845 m).”

Distribution: Region 1. Eastern versant of Sierra de

Perija, Zulia State, Venezuela; very probably in adjacent

Colombia.

Remarks: While the description of this species was

in press, another work dealing with Cryptobatrachus

(Lynch 2008) provided the description of C. pedroruizi, a

species from the Colombian side of Perija relatively near

the known distribution area of C. remotus. Comparing

both sets of type material (both morphologically and

genetically) is imperative to discern if they represent one

or two species. Cryptobatrachus remotus 1s considered

Endangered (EN) under criteria of the Venezuelan Red

Book (Rojas-Runjaic 2015).

Selected references: Infante-Rivero et al. (2008), Lynch

(2008); Duellman (2015); Rojas-Runjaic et al. (2015).

Genus Flectonotus Miranda-Ribeiro, 1920

Type species: Nototrema pygmaeum, Boettger, 1893, by

original designation.

Remarks: In their recent phylogenetic analysis

Castroviejo et al. (2015) inferred Flectonotus as

sister of Cryptobatrachus including both genera in

Cryptobatrachinae.

Flectonotus fitzgeraldi (Parker 1933)

Holotype: BM 1947.2.22.41.

Type locality: “Mt. Tucuche, Trinidad.”

Distribution: Region 2. Northern range of Trinidad

(Trinidad and Tobago), and Venezuela, where restricted

to the Peninsula de Paria.

Selected references: Duellman and Maness (1980);

Duellman and Gray (1983); Duellman et al. (1988, 2011).

Flectonotus pygmaeus (Boettger 1893)

Lectotype: SMF 2679.

Type locality: “Puerto Cabello, Estado Carabobo,

Venezuela.”

Distribution: Regions 1, 2. Venezuela and Colombia.

In Venezuela, on both sides of the Andes (Cordillera de

Merida) below 1,500 m asl, and Cordillera de la Costa,

above 800 m asl.

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Catalogue of the amphibians of Venezuela

104. Adelophryne gutturosa. La Laja, Sierra de Lema, Bolivar.

Photo: César Barrio-Amoros.

‘ F he

105B. Cryptobatrachus remotus. Female with eggs. Cerro Las

Antenas, Sierra de Perija, Zulia. Photo: Fernando Rojas-Runjaic.

106A. Flectonotus fitzgeraldi. Las Melenas, Peninsula de Paria,

Sucre. Photo: César Barrio-Amoros.

Remarks: Populations from Cordillera de la Costa and

Andes are indistinguishable genetically (D. Blackburn

and W.E. Duellman, pers. comm.).

Selected references: Boettger (1893); Lutz (1927);

Rivero (1961, 1964a, 1971a); Mertens (1967); Duellman

Amphib. Reptile Conserv. 54

105A. Cryptobatrachus remotus. Male. Cerro Las Antenas, Sierra

de Perija, Zulia. Photo: Fernando Rojas-Runjaic.

105C. Cryptobatrachus remotus. Female with recently hatched

froglets. Cerro Las Antenas, Sierra de Perija, Zulia. Photo:

Fernando Rojas-Runjaic.

Pa ds *

eg ue

sae aE ae ASPET

106B. Flectonotus fitzgeraldi. Macuro, Peninsula de Paria, Sucre.

Photo: Diego A. Flores.

er tar

and Maness (1980); Duellman and Gray (1983); Péfaur

and Diaz De Pascual (1987); Duellman et al. (1988,

2011); Mijares-Urrutia and Arends (1993); Manzanilla

et al. (1995); Yustiz (1996); Rivas and Barrio-Amoros

(2005); Barrio-Amoros (2006c, 2010a).

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Subfamily Hemiphractinae Peters, 1862

Genus Gastrotheca Fitzinger, 1843

Type species: Hyla marsupiata Dumeéril and Bibron,

1841, by original designation.

Remarks: In the recent phylogeny of hemiphractids

by Castroviejo et al. (2015) none of the taxonomies

historically proposed for Gastrotheca were found

monophyletic (including some subgenera proposed by

Duellman 2015). Consequently, they proposed a new

infrageneric taxonomy based on four species groups

(fissipes, longipes, marsupiata, and microdiscus).

Gastrotheca helenae Dunn, 1944

Holotype: MLS 268.

Type locality: Paramo de Tama, Departamento Norte de

Santander, Colombia.

Distribution: Region 1. Endemic of paramo de Tama,

between Colombia and Venezuela.

Remarks: Acevedo et al. (2011) presented natural history

data on this rare species. In subgenus Amphignathodon

by Duellman (2015), or in the Gastrotheca longipes

species group sensu Castroviejo-Fisher et al. (2015).

Selected references: Duellman (1980, 1989b, 2015);

Duellman and Ruiz-Carranza (1986); Duellman et al.

(1988); Acevedo et al. (2011); Castroviejo-Fisher et al.

(2015).

Gastrotheca nicefori Gaige, 1933

Holotype: UMMZ 73242.

Type locality: Pensilvania,

Colombia.

Distribution: Region 1. Panama, Colombia, and

Venezuela. In Venezuela, distributed throughout the

Cordillera de Mérida. Very likely present in Sierra de

Pera.

Remarks: Duellman (1989b) synonymized Gastrotheca

yacambuensis Yustiz, 1976 with G. nicefori, without

explanation. La Marca (1992) recognized the synonymy,

but Barrio-Amoros (1998, 2004, 2009) was reluctant to

do so as it was done without providing any comparison.

Duellman (2015) presented arguments for the synonymy

(see below), but molecular evidence (Castroviejo et

al. 2015) indicated G. yacambuensis could be a valid

species, different from G. nicefori. Further evidence is

necessary to assess this issue adequately. The species is

included in the subgenus Duellmania (Duellman 2015)

and in the Gastrotheca marsupiata species group sensu

Castroviejo-Fisher et al. (2015).

Selected references: Yustiz (1976a,b, 1996); Duellman

(1980, 1989b, 2015); Duellman et al. (1988); La Marca

(1991b “1994,” 1992): Castroviejo-Fisher et al. (2015).

Departamento Caldas,

Gastrotheca ovifera (Lichtenstein and Weinland 1854)*

Amphib. Reptile Conserv.

Holotype: ZMB 3073.

Type locality: Puerto Cabello, Carabobo, Venezuela.

Distribution: Region 2. Cordillera de la Costa in states

Aragua, Cojedes, Miranda, Distrito Federal, Carabobo,

and Yaracuy, 890—2,060 m asl.

Remarks: Reported from the Delta of the Orinoco River

(MHNLS 2942) by Valera-Leal et al. (2011) without

further comment; but current authors believe this is an

error as no Gastrotheca specimen has ever been reported

from that area. Included in subgenus Opisthodelphys

(Duellman 2015) or in Gastrotheca marsupiata species

group sensu Castroviejo-Fisher et al. (2015). Despite

a 646 man-hour effort, Valera-Leal et al. (2011) failed

to locate any surviving population in the Henri Pittier

National Park in 2006-2007. Barrio-Amoros (1998)

reported it very abundant at Galipan in March 1997,

while CBA has heard the species in Cerro Volcan,

Caracas, Miranda, as recently as 2009.

Selected references: Lutz (1927); Mertens (1957a,b,

1967); Ginés (1959); Rivero (1961, 1964a,d); Tello

(1968); Duellman (1980, 2015); Duellman et al. (1988),

La Marca (1995a); Manzanilla et al. (1995); Barrio

(1999b); Barrio-Amoros (2001, 2006c, 2013); Schmidt

et al. (2002); Manzanilla and Sanchez (2003); Rivas

and Barrio-Amoroés (2005); Valera-Leal et al. (2011);

Castroviejo-Fisher et al. (2015).

Gastrotheca walkeri Duellman, 1980*

Holotype: UMMZ 117177.

Type locality: “From between the Estacion Bioldgica

Rancho Grande and Paso Portachuelo, Estado de Aragua,

Venezuela,” 1,100 m.

Distribution: Region 2. Cordillera de la Costa in states

Aragua, Cojedes, D.F., Carabobo, and Yaracuy.

Remarks: Duellman (2015) stated that no call is

associated to this species, but the current authors find

it commonly heard from high bromeliads in Coastal

cloud forest (e.g., in Rancho Grande, Estado Aragua;

see Barrio-Amoros 2006c). In Gastrotheca longipes

species group sensu Castroviejo-Fisher et al. (2015) or in

subgenus Cryptotheca by Duellman (2015).

Selected references: Duellman (1980, 2015); Duellman

et al. (1988); Manzanilla et al. (1995); Barrio (1999c),;

Rivas and Barrio-Amorés (2005); Barrio-Amoros

(2006c).

Gastrotheca williamsoni Gaige, 1922*

Holotype: UMMZ 55559.

Type locality: San Esteban, Carabobo, Venezuela.

Distribution: Region 2. Known only from type locality.

Remarks: Not collected since original description. Either

extremely rare, or extinct, or Gastrotheca walkeri could

be a junior synonym. A review of this species is needed to

clarify its taxonomy and conservation status. Duellman

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Catalogue of the amphibians of Venezuela

107A. Flectonotus pygmaeus.

César Barrio-Amoros.

107C. Flectonotus pygmaeus. Female. Rancho Grande, Henri

Pittier National Park, Aragua. Photo: César Barrio-Amoros.

BU Sonar ‘eh?

109. Gastrotheca nicefori Altos de San Luis, La Azulita, Mérida.

Photo: César Barrio-Amoros.

(2015) distinguished G. williamsoni from G. walkeri

based on four morphological characters (absence of

supraciliary processes — vs present in wal/keri—, fingers

v2 webbed — vs 4 webbed in walkeri—, first finger

longer than second — shorter in wal/keri—, and granular

tympanic annulus — smooth in walkeri. The only known

locality of G. williamsoni lies within the distribution

of G. walkeri, and no additional specimen obtained for

almost a century. As both species share a unique character

Amphib. Reptile Conserv.

107B. Flectonotus pygmaeus. Male. Casa Maria, Bejuma,

Carabobo. Photo: César Barrio-Amoros.

a * ce oc

“let ieee

Pn eke.

, Sake eS, a ee,

108. Gastrotheca helenae. Paramo Tama, Norte de Santander,

Colombia. Photo: Aldemar A. Acevedo.

110A. Gastrotheca ovifera. Altos de Pipe, Miranda. Photo:

Aldemar A. Acevedo.

among marsupial frogs (paired retroperitoneal brood

pouches), have overlapping geographic distributions,

and only differ in morphological traits prone to

preservation artifacts, makes us doubt the validity of

G. walkeri, assessing their taxonomic status requires

further investigation. Included in subgenus Cryptotheca

by Duellman (2015) or in Gastrotheca longipes species

group sensu Castroviejo-Fisher et al. (2015).

Selected references: Gaige (1922); Lutz (1927); Ginés

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

(1959); Rohl (1959); Rivero (1961, 1964a); Duellman

(1980, 2015); Duellman et al. (1988); Barrio-Amoros

(2013).

Gastrotheca yacambuensis Yustiz, 1976*

Holotype: MBUCO 6015.

Type locality: “Quebrada El Cedral, Parque Nacional

Yacambu, ladera Sur de Sierra Portuguesa, Estado Lara,

Venezuela, 1,700 m.”

Distribution: Region 1. Parque Nacional Yacambu,

Lara State, and probably all Cordillera de Mérida in the

Venezuelan Andes.

Remarks: Duellman (1989b) synonymized Gastrotheca

yacambuensis with G. nicefori without providing

any argument. Barrio-Amorés (1998, 2004, 2013)

and Barrio-Amoros et al. (2009) did not follow the

synonymy, waiting for additional evidence. Duellman

(2015) compared overall external similarities among

names he synonymized (Duellman 1989b) to support

his previous statement. Castroviejo et al. (2015)

showed that topotypical specimens of G. nicefori are

significantly different genetically from specimens from

the Venezuelan Andes; so, G. yacambuensis could be the

name for populations previously referred to G. nicefori

in the Venezuelan Andes. The current authors consider

G. yacambuensis as a valid species, at least for central

Andean populations, while awaiting a comprehensive

review of the Andean populations of Gastrotheca

nicefori.

Selected references: Yustiz (1976a,b, 1996); Duellman

(1989b); Castroviejo et al. (2015).

Genus Stefania Rivero, 1968

Type species: Hy/a evansi Boulenger, 1904, by original

designation.

Remarks: Infrageneric groups in Stefania (S. evansi and

S. goini species group) are no longer recognized, since

Kok et al. (2012) and Castroviejo-Fisher et al. (2015)

inferred these two groups as non-monophyletic.

Stefania breweri Barrio-Amoros and Fuentes, 2003*

Holotype: MBUCV 6574.

Type locality: “Summit of Cerro Autana (Wahari

Kuaway), near the north ridge (4°52’N, 67°27’W),

1,250 m elevation, Municipio Atures, Estado Amazonas,

Venezuela.”

Distribution: Region 5. Endemic to Autana, a tepui in

Estado Amazonas.

Remarks: Not seen during two expeditions to the top of

Autana after the collection of the only known specimen

in 1971. Autana has an extremely small surface of 1.9

km?, so this is one of the most geographically restricted

frog species in the world. However, it is only DD by

IUCN (Stuart 2006) and Rodriguez and Rojas-Suarez

Amphib. Reptile Conserv.

(2008) keep it under IUCN category VU/D2; it is not

even mentioned in the last edition of the Venezuelan Red

Book (Rodriguez et al. 2015). We argue that the species

(known from a single specimen, not seen during two

expeditions with herpetologists in 1972 and 2001, and

restricted to an area of 1.9 km’) should be classified CR

B2a.

Selected references: Barrio-Amoros and Fuentes

(2003); Stuart (2006); Rodriguez and Rojas-Suarez

(2008); Barrio-Amords and Torres (2010); Barrio-

Amoréos (2013).

Stefania ginesi Rivero, 1968*

Holotype: FMNH 74041.

Type locality: “Chimanta-tepui, Venezuela, rock

outcrops near E. branch of headwater Rio Tirica, 7,300

fy (22225 min)

Distribution: Region 5. Endemic to Chimanta massif

in Estado Bolivar. Restricted to Chimanta-tepui (type

locality) by Kok et al. (2016).

Selected references: Rivero (1966, 1967c, 1970);

Duellman and Hoogmoed (1984); Gorzula (1992);

Sefiaris et al. (1996, 2014); Gorzula and Sefiaris (1998);

Barrio-Amoros and Fuentes (2003, 2012); McDiarmid

and Donnelly (2005); Salerno and Pauly (2012); Kok et

al. (2016, 2017).

Stefania goini Rivero, 1968*

Holotype: AMNH 23193.

Type locality: “Vegas Falls, Mte. Duida, Amazonas,

Venezuela, 4,600 ft.”

Distribution: Region 5. Endemic to Duida and

Huachamakari, tepuis in Estado Amazonas.

Selected references: Rivero (1966, 1967c, 1970);

Duellman and Hoogmoed (1984); Sefiaris et al. (1996);

Barrio-Amorés and Fuentes (2003); McDiarmid and

Donnelly (2005).

Stefania marahuaquensis (Rivero 1961)*

Holotype: MCZ 285566.

Type locality: Cafio Caju, Cerro Marahuaca, Estado

Amazonas, Venezuela, 120 m.

Distribution: Region 5. Endemic to Duida and

Marahuaka, tepuis in Amazonas State.

Selected references: Rivero (1961, 1964b, 1970);

Duellman and Hoogmoed (1984); Sefiaris et al. (1996);

Barrio-Amorés and Fuentes (2003); McDiarmid and

Donnelly (2005).

Stefania oculosa

Sefiaris, Ayarzaguena and Gorzula, 1997*

Holotype: MHNLS 12961.

Type locality: “Tepuy Jaua, Estado Bolivar, Venezuela

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Catalogue of the amphibians of Venezuela

110B. Gastrotheca ovifera. Altos de Pipe, Miranda. Photo: J.

Celsa Sefiaris.

111. Gastrotheca walkeri. Rancho Grande, Henri Pittier National

Park, Aragua. Photo: Juan Vicente Rueda-Almonacid.

112. Stefania breweri. Holotype MBUCV 6574. Top of Cerro

Autana, Amazonas. Photo: César Barrio-Amoros.

113B. Stefania ginesi. Female carrying froglets. Churi-tepui,

Chimanta massif, Bolivar. Photo: Charles Brewer-Carias.

(04°49°55”N, 64°25’59”W). 1,600 m snm.”

Distribution: Region 5. Endemic to Jaua, a tepui in

Bolivar State.

Selected references: Sefiaris et al. (1996); Barrio-

Amoros and Fuentes (2003).

Stefania percristata

Sefiaris, Ayarzaguena and Gorzula, 1997*

Holotype: MHNLS 12952.

Type locality: “Tepuy Jaua, Estado Bolivar, Venezuela

(04°49’55”N, 64°25°59”"W), 1,600 m snm.”

Amphib. Reptile Conserv.

oe a

and Brewer-

Carias (2008). Sima Mayor Sarisarifiama-tepui, Bolivar. Photo:

Javier Mesa.

Distribution: Region 5. Endemic to Jaua, a tepui in

Bolivar State.

Selected references: Sefiaris et al. (1996); Barrio-

Amoros and Fuentes (2003).

Stefania riae Duellman and Hoogmoed, 1984*

Holotype: KU 174688.

Type locality: “Cerro Sarisarifiama, 1,400 m, Estado

Bolivar, Venezuela.”

Distribution: Region 5. Endemic to Sarisarifiama, a

tepui in Bolivar State.

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

Remarks: Barrio-Amoros and Brewer-Carias (2008)

present variation of the species not previously noted.

Selected references: Duellman and Hoogmoed (1984);

Sefiaris et al. (1997); Barrio-Amoros and Fuentes (2003);

McDiarmid and Donnelly (2005); Barrio-Amoros and

Brewer-Carias (2008); Kok et al. (2017).

Stefania riveroi

Sefiaris, Ayarzaguena and Gorzula, 1997*

Holotype: MHLNS 10413.

Type locality: Yuruani-tepui, Estado Bolivar, Venezuela

(05°19’N, 60°51’ W), 2,300 m snm.

Distribution: Region 5. Endemic to Yuruani, a tepui in

Bolivar State.

Selected references: Sefiaris et al. (1996, 2014); Gorzula

and Sefiaris (1998); Barrio-Amoros and Fuentes (2003);

Sinsch and Juraske (2006); Kok et al. (2017).

Stefania satelles

Sefiaris, Ayarzaguena and Gorzula, 1997*

Holotype: MHNLS 10433.

Type locality: “Aprada-tepui I, Estado Bolivar,

Venezuela (05°24’N, 62°27’W), 2,500 m snm.”

Distribution: Region 5. Described as endemic to several

small tepuis in Estado Bolivar: Aprada-tepui, Angasima-

tepui (Adanta), and Upuigma-tepui (El Castillo) sensu

Sefiaris et al. (1996). Kok et al. (2016) restricted it to

Aprada tepui, its type locality, while the rest of Stefania

on other tepuis formerly identified as sate/les are

confirmed as candidate species (see Appendix 1).

Selected references: Gorzula (1992); Sefiaris et al.

(1996, 2014); Gorzula and Sefiaris (1998); Barrio-

Amoros and Fuentes (2003); Salerno and Pauly (2012);

Kok et al. (2016, 2017).

Stefania scalae Rivero, 1970

Holotype: MCZ 64373.

Type locality: “La Escalera, road between El Dorado

and Sta. Elena de Uairén, around 1,200 m, Edo. Bolivar,

Venezuela.”

Distribution: Region 5. Widespread in uplands of

eastern Estado Bolivar.

Remarks: Stefania evansi was considered part of

the Venezuelan batrachofauna after S. scalae was

synonymized with S. evansi by Duellman and Hoogmoed

(1984). MacCulloch and Lathrop (2002) found further

evidence to distinguish between S. evansi and S. scalae.

Barrio-Amoros and Fuentes (2003) pointed out, after the

resurrection of S. scalae by Sefiaris et al. (1997), that

there is no record of S. evansi from Venezuela, although

its presence Is likely.

Selected References: Rivero (1970); Gorzula et al.

(1983); Duellman and Hoogmoed (1984); Sefiaris et al.

(1996, 2014); Duellman (1997); Gorzula and Sefiaris

Amphib. Reptile Conserv.

(1999); MacCulloch and Lathrop (2002); Barrio-Amoroés

and Fuentes (2003); McDiarmid and Donnelly (2005);

Sinsch and Juraske (2006); Barrio-Amoros and Duellman

(2009); Barrio-Amoros et al. (2011b).

Stefania schuberti

Sefiaris, Ayarzaguena and Gorzula, 1997*

Holotype: MHNLS 12917.

Type locality: Sector este, cima del Auyan-tepui, Estado

Bolivar, Venezuela (05°53’36”N, 62°29712”W), 1,750 m

asl.

Distribution: Region 5. Endemic to the summit of

Auyan-tepui, Bolivar State.

Selected references: Sefiaris et al. (1996, 2014); Barrio-

Amoros and Fuentes (2003, 2012); Myers and Donnelly

(2008); Kok et al. (2017).

Stefania tamacuarina Myers and Donnelly, 1997

Holotype: AMNH 131428.

Type locality: “Camp on ridge north Pico Tamacuari,

1,270 m elevation, Sierra Tapirapecd, Amazonas,

Venezuela (1°13’N, 64°42’W).”

Distribution: Region 5. Only known from type locality

in Venezuela and one nearby locality on the Brazilian

side of Tapirapeco (Caramaschi and Niemeyer 2005b).

Selected references: Myers and Donnelly (1997);

Barrio-Amoros and Fuentes (2003); Caramaschi and

Niemeyer (2005b); McDiarmid and Donnelly (2005).

Family Hylidae Rafinesque, 1815

Remarks: We follow Duellman’s et al. (2016) recent

revision, where Phyllomedusinae was raised to family

level (Phyllomedusidae, see below). Five subfamilies

are considered under the family rank in Venezuela:

Cophomantinae, Dendropsophinae, Lophyohylinae,

Pseudinae, and Scinaxinae. Dubois (2017) argues that

Boana Gray, 1825 has priority over Hypsiboas.

Subfamily Cophomantinae Hoffmann, 1878

Remarks: After Duellman et al. (2016), this subfamily

contains six genera: Aplastodiscus, Bokermannohyla,

Colomascirtus (described by them to accommodate

species previously assigned to Hyloscirtus larinopygion

and H. armatus species groups), Hyloscirtus, Hypsiboas

(now Boana), and Myersiohyla. However, based on

molecular evidence, Rojas-Runjaic et al. (2018),

synonymized Colomascirtus with Hyloscirtus.

Genus Boana Gray, 1825

Type species: Rana boans Linnaeus, 1758, by monotypy.

Coined as subgenus of Hyla according to Dubois (2017).

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Catalogue of the amphibians of Venezuela

114B. Stefania riae. Pattern B in Barrio-Amoros and Brewer-Carias

(2008). Sima Mayor Sarisarifiama-tepui, Bolivar. Photo: Javier Mesa.

115A. Stefania aera Sammit of ST Bolivar. Photo:

ees Kok.

11SB. Stefania riveroi. Summit of SE tepui, ‘Bolivar. Photo:

Philippe Kok.

116. Stefania satelles: Aprada-tepui, Bolivar. Photo: Brad Wilson.

Remarks: Faivovich et al. (2005) assigned Centrolenella

pulidoi Rivero, 1968 to the Boana benitezi group. We

agree that description of holotype corresponds to a

juvenile B. benitezi. Faivovich et al. (2005) examined the

holotype (MCZ 72499) but was not sure if it was an adult

female or a juvenile. According to our combined field

experience, only juveniles of the B. benitezi group (B.

benitezi, B. lemai, and B. tepuianus at least) have reddish

Amphib. Reptile Conserv.

115C. Stefania riveroi. Female with eggs. Summit of Yuruani-

tepui, Bolivar. Photo: Steve Gorzula.

117A. Stefania scalae. Plain morph. Lowlands of Cuyuni river,

Bolivar. Photo: César Barrio-Amoros.

eyes, as Rivero (1968) reported in pulidoi. In Cerro Duida

the only known species of the Boana benitezi group 1s B.

benitezi. We here consider Centrolenella pulidoi Rivero,

1968 (recognized currently as Boana pulidoi by Frost

2018) as a junior synonym of Hyla benitezi (Rivero

1961), as Boana benitezi. Recently Dubois (2017)

defended the use of Boana for the clade preoccupied by

Hypsiboas, arguing nomenclatural precedence.

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Barrio-Amoros et al.

117B. Stefania scalae. Stripped morph. Lowlands of Cuyuni river,

Bolivar. Photo: César Barrio-Amoros.

° Kye

sgn

Bolivar. Photo: Elio Sanoja.

118. Stefania schuberti. Auyan-tepui Summit, Bolivar. Photo:

César Barrio-Amoros.

Boana alemani (Rivero 1964)*

Holotype: MHNLS 238.

Type locality: Cagua, Estado Aragua, Venezuela.

Distribution: Regions 1, 2. Cagua (Aragua State)

Kunana, Sierra de Perija (Zulia State), and Hato Los

Arrecifes (Guarico State); Cojedes and Falcon States.

Remarks: Currently in Boana punctata species group

(Faivovich et al. 2005). After original description no

Amphib. Reptile Conserv.

117C. Stefania scalae. Spotted morph. Lowlands of Cuyuni river,

Bolivar. Photo: César Barrio-Amoros.

117E. Stefania scalae. Female carrying recently hatched froglets.

Sierra de Lema, Bolivar. Photo: Alberto Blanco.

other specimens have been assigned with accuracy to

this taxon. Taxonomic status unclear since the species

description was based on two old and poorly preserved

specimens from disparate bioregions (holotype from

Cagua in Cordillera de La Costa, and paratype from

Kunana in Sierra de Perija) and probably not conspecific.

Rivero (1964) said this species is narrowly related to Hyla

granosa (= Boana cinerascens) and recognized that the

two species only can be differentiated by the dorsal spots

and the inclination of the loreal region. La Marca (1992)

said B. alemani and B. punctata are probably synonyms,

but B. punctata is not known from extreme northern

Venezuela. Juveniles of B. xerophylla and H. pugnax are

also green and somewhat similar to B. punctata. Based

on a preliminary examination of the type series (by

FRR) we also suspect B. alemani could be conspecific

with B. punctata, and the paratype (MHNLS 187) could

be a juvenile B. xerophylla. Specimens assigned to B.

alemani MHNLS 150 (also from Kunana) and MHNLS

407 (from Caracas) probably also correspond to juvenile

B. xerophylla. We provisionally consider this name

valid, but its taxonomic status should be confirmed, as

well as the identity of additional records from Cojedes

and Falcon (Manzanilla et al. 2000), Guarico (Camargo

et al. 2014), and the tadpole description (Miyjares-Urrutia

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Catalogue of the amphibians of Venezuela

119A. Hyloscirtus jahni. Day color. Monte Zerpa, Mérida. Photo:

César Barrio-Amoros.

120. Hyloscirtus japreria. Night color. Cerro Las Antenas, Zulia.

Photo: Fernando Rojas-Runjaic.

122. Ayloscirtus platydactylus. Cedral, San Luis, La Azulita,

Mérida. Photo: César Barrio-Amoros.

1992).

Selected references: Rivero (1964d, 1967c); Mijares-

Urrutia (1992); Manzanilla et al. (1995); Barrio-Amoros

(2013); Camargo et al. (2014).

Boana benitezi (Rivero 1961)

Amphib. Reptile Conserv.

119B. Hyloscirtus jahni. Day color. Monte Zerpa, Mérida. Photo:

César Barrio-Amoros.

121. Ayloscirtus lascinius. Male. La Macana, south of Santa Cruz

de Mora, Mérida. Photo: César Barrio-Amoros.

123A. Boana boans. La Laja, Sierra de Lema, Bolivar. Photo:

César Barrio-Amoros.

Type: MCZ 28564.

Type locality: Cafio Wanadi, Cerro Marahuaca, Estado

Amazonas, Venezuela.

Distribution: Region 5. Western Guayana uplands and

highlands, and northernmost Roraima State in Brazil.

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

Remarks: In Boana benitezi species group (Faivovich et

al. 2005). Boana tepuiana is a vicariant species from the

eastern Guayana uplands (Barrio-Amoros and Brewer-

Carias 2008).

Selected references: Rivero (1961, 1964b, 1967a,

1971b); Donnelly and Myers (1991); Heyer (1994b);

Gorzula and Sefiaris (1998); Barrio-Amoros and Brewer-

Carias (2008).

Boana boans (Linnaeus 1758)

Lectotype: ZIUU 27; designated by implication (Mertens

1940).

Type locality: “America.”

Distribution: Regions 1, 2, 3, 4, 5. Widely distributed

in northern South America, in the Darien and Choco

Regions, and Magdalena, Orinoco, and Amazon Basins

(Panama, Colombia, Ecuador, Venezuela, Brazil,

Guianas, and Trinidad). Common riparian element in

the lowlands of southern Venezuela (Amazonas, Bolivar,

and Delta Amacuro to as far north as Peninsula de Paria).

Also, in the eastern piedmont of the Venezuelan Andes

(Barinas and Tachira).

Remarks: The Andean piedmont population (Barrio-

Amoros 1999g, 2001a) is continuous from the eastern

versant of the Cordillera Oriental de Colombia and

thus from the Upper Amazon, fitting the Amazonian

distribution pattern for western Venezuela suggested by

Barrio-Amoros (1998). However, the Barinas and Tachira

populations are much smaller and with a different pattern

from those from the south of the Orinoco. Chacon-Ortiz

et al. (2005) probably confused by such difference,

reported B. rosenbergi, a Central American and Chocoan

species, from Tachira State, and the same specimens

mentioned by Chacon-Ortiz et al. (2005) were previously

reported correctly as B. boans (Barrio-Amoros 1999,

2001a). Specimens from throughout the whole species

distribution should be compared using molecular and

bioacoustic data, as they may be a species complex. In

the Boana semilineata species group (Faivovich et al.

2005). Some specimens are still confused in collections

with the very similar B. wavrini (Hoogmoed 1990b).

Selected references: Ginés (1959); Rivero (1961, 1964b,

1967a); Heatwole et al. (1965); Duellman (1971a, 1997);

Gremone et al. (1986); Hoogmoed (1990b); Donnelly

and Myers (1991); Magdefrau et al. (1991); Gorzula

and Sefiaris (1998); Barrio-Amorés (1999, 2001a);

Arrington and Arrington (2000); Lynch and Suarez-

Mayorga (2001); Chacon et al. (2005); Barrio-Amoros

and Brewer-Carias (2008); Barrio-Amoros et al. (2011b),;

Mendoza (2014); Sefiaris et al. (2014).

Boana calcarata (Troschel 1848)

Holotype: Not designated and probably lost.

Type locality: “British Guiana.”

Distribution: Venezuela, Brazil, Colombia, Ecuador,

Amphib. Reptile Conserv.

Bolivia, Guyana, and French Guiana. Widespread in

Southern Venezuela, in Amazonas, Bolivar, and Delta

Amacuro.

Remarks: Regions 4, 5. Caminer and Ron (2014)

reviewed the species and allies from the Upper Amazon,

especially Ecuador. They doubted the identity of

Colombian and Venezuelan samples despite the fact they

were well documented (Barrio-Amoros 1998; Sefiaris

and Barrio 2002). In the Boana albopunctata species

group (Faivovich et al. 2005).

Selected references: Duellman (1973); Donnelly and

Myers (1991); Sefiaris and Barrio (2002); Sefiaris and

Ayarzagtiena (2004 “2002”); Barrio-Amoros et al.

(2011b).

Boana cinerascens (Spix 1824)

Syntypes: ZSM 2498/0 (2 specimens), destroyed.

Type locality: “Ecga prope flumen Teffe” (= Ega, Tefé),

Amazonas, Brazil.

Distribution: Regions 4, 5. Widely distributed in the

Amazon Region (Colombia, Ecuador, Peru, Bolivia,

Brazil, Venezuela, Guyana, Suriname, and French

Guiana). Widespread in Southern Venezuela, in

Amazonas, Bolivar, and Delta Amacuro States.

Remarks: Inthe Boana punctata species group (Faivovich

et al. 2005). In Venezuela two species are confused under

this name, one from rainforest habitat (Image 125A)

and another from open areas (Image 125B). The nomen

cinerascens hides a complex of species, with several

taxa undescribed in the upper Amazon (D. Pareja, pers.

comm. ). Hoogmoed and Gruber (1983) suggested the use

of B. granosa (Boulenger 1882) instead of B. cinerascens

(Spix 1824) to stabilize the use of granosa. Barrio-

Amoros (2004) used the name cinerascens, overlooking

Hoogmoed and Gruber’s suggestion. Current situation

is complicated, with types of B. cinerascens from Tefé

apparently representing a different species than lectotype

of B. granosa from Canelos, Ecuador (P. Kok, pers.

comm. ).

Selected references: Spix (1824); Boulenger (1882);

Rivero (1961, 1964b,d, 1967c, 1971b); Duellman (1974a,

1997); Hoogmoed (1979a); Hoogmoed and Gruber

(1983); Gorzula and Sefiaris (1998); Frost (2004); Kok

and Kalamandeen (2008); Barrio-Amoros et al. (2011b);

Sefiaris et al. (2014).

Boana geographica (Spix 1824)

Holotype: Not designed. ZSM 35/0 is part of the type

series but is currently lost.

Type locality: Rio Tefé, Amazonas, Brazil.

Distribution: Region 4, 5. Widespread in tropical South

America, east of Andes, including Trinidad. Widely

distributed in southern and eastern Venezuela (Amazonas,

Bolivar, and Delta Amacuro States).

Remarks: Fouquet et al. (2016) show a wider variation of

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Catalogue of the amphibians of Venezuela

123B. Boana boans. Acequias river, Barinas. Photo: César Barrio-

Amoros.

125A. Boana cinerascens. Cuyuni river, Bolivar. Photo: César

Barrio-Amoros.

. rs i i — . =" = |

126. Boana geographica. Female. Imataca Forestal Reserve, Delta

Amacuro. Photo: César Barrio-Amoros.

both B. geographica and B. semilineata. They described

B. diabolica from the eastern Guiana Shield. A detailed

review should be done with Venezuelan populations. In the

Boana semilineata species group (Faivovich et al. 2005).

Selected references: Spix (1824); Gunther (1858);

Lutz (1927); Rivero (1961); Duellman (1971a, 1973,

1997); Hoogmoed and Gorzula (1979); Lynch (1979b),;

Gremone et al. (1986); Azevedo-Ramos (1995); Gorzula

and Sefiaris (1998); Barrio-Amoroés et al. (2011b);

Sefiaris et al. (2014); Fouquet et al. (2016).

Amphib. Reptile Conserv.

. ‘/., . eee

124. Boana calcarata. Cuyuni river, Bolivar. Photo: César Barrio-

Amoros.

125B. Boana cinerascens. Savannas surrounding Cerro Autana,

Amazonas. Photo: César Barrio-Amoros.

127. Boana_ hobbsi. Colombia. Photo:

Castroviejo-F isher.

Boana hobbsi (Cochran and Goin 1970)

Leticia,

Santiago

Holotype: MCZ 28052.

Type locality: “Cano Goacaya, a tributary of the Rio

Apaporis, in Amazonas, Colombia.”

Distribution: Region 4. Colombia and Venezuela. In

Venezuela known from a single record from the base of

Cerro Neblina in extreme southern Amazonas State.

Remarks: In the Boana punctata species group

(Faivovich et al. 2005).

Selected references: Cochran and Goin (1970);

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Duellman (1974a); Pyburn (1978); McDiarmid and

Paolillo (1988).

Boana jimenezi

(Sefiaris and Ayarzagtiena 2006)*

Holotype: MHNLS 16070.

Type locality: Quebrada Ataperé, headwater of the

Cucurital River (05°43’17.7”N, 62°34’21.1”’W), 970 m,

Canaima National Park, Bolivar State, Venezuela

Distribution: Region 5. Endemic to Venezuela, restricted

to the uplands of Gran Sabana and the summit of Auyan-

tepui. Expected in Guyana.

Remarks: Probably a member of Boana punctata species

group (Sefiaris and Ayarzagtiena 2006).

Selected references: Cochran and Goin (1970);

Duellman (1974a); Pyburn (1978); McDiarmid and

Paolillo (1988); Sefiaris et al. (2014).

Boana lanciformis (Cope 1871)

Holotype: Not designed. Type series unknown.

Type locality: “Pebas, Ecuador.”

Distribution: Regions 1, 2, 3, 4. Amazonia, eastern

slopes of the Andes, southern slopes of Cordillera de la

Costa, Peninsula de Paria.

Remarks: In the Boana albopunctata species group

(Faivovich et al. 2005). Hyla lanciformis guerreroi was

described by Rivero (1971) for the Venezuelan form.

Discussing with colleagues (L.F. Toledo, A. Chacon),

some differences are clear (including morphological and

osteological) and B. lanciformis could hide a species

complex. If so, the distinctiveness of the Venezuelan

form should be demonstrated by an integrative taxonomic

approach.

Selected references: Duellman (1971a, 1973); Rivero

(1971a); Péfaur and Diaz De Pascual (1987); Gorzula

and Sefiaris (1998); Barrio et al. (1999); Barrio-Amoros

(2010a).

Boana lemai* (Rivero 1971)

Holotype: UPR-M 3179.

Type locality: “Paso del Danto, La Escalera, entre

El Dorado y Santa Elena de Uairen, 1,300—1,400 m,

Serrania de Lema, Estado Bolivar, Venezuela.”

Distribution: Region 5. Eastern Guayana uplands

(Venezuela and Guyana). In Venezuela, known from

Sierra de Lema and some localities in the Gran Sabana.

Remarks: See variation in specimens from Guyana in

MacCulloch and Lathrop (2005). In the Boana benitezi

species group (Faivovich et al. 2005).

Selected references: Rivero (1971b); Duellman (1997);

MacCulloch and Lathrop (2005); Myers and Donnelly

(2008); Barrio-Amoros and Duellman (2009); Sefiaris et

al. (2009, 2014).

Amphib. Reptile Conserv.

Boana multifasciata (Gunther 1859)

Holotype: BM 1947.2.23.6

Type locality: “Para,” Brazil.

Distribution: Region 5. Brazil, Venezuela, Guyana,

Suriname, and French Guiana. In Venezuela, restricted

to the Eastern Guayana (Bolivar, Monagas, and Delta

Amacuro).

Remarks: In Boana albopunctata species group

(Faivovich et al. 2005).

Selected references: Rivero (1961, 1964a, 1967a);

Hoogmoed and Gorzula (1979); Duellman (1997);

Gorzula and Sefiaris (1998); Sefiaris and Ayarzagtiena

(2004); Sefiaris et al. (2014).

Boana ornatissima (Noble 1923)

Holotype: AMNH 13491.

Type locality: “Meamu, Mazaruni River, Guyana.”

Distribution: Region 5. Known in Venezuela from two

localities; one in the Gran Sabana (salto Karuay), and

another from middle Amazonas State. A picture was

shown to CBA of a B. ornatissima from Surumoni, near

La Esmeralda (Amazonas State), taken by the Austrian

Surumoni team from University of Vienna in 1998. This

locality fills the gap between the Gran Sabana locality

and that from Guainia, eastern Colombia (Lynch and

Vargas-Ramirez 2000).

Remarks: In Boana punctata species group (Faivovich

et al. 2005).

Selected references: Hoogmoed (1979a); Sefiaris

and Vernet (1997); Lynch and Vargas-Ramirez (2000);

Sefiaris et al. (2014).

Boana pugnax (Schmidt 1857)

Holotype: KM 1009.

Type locality: “Chiriqui, Flusse unneit Bocca (sic!) del

Toro.” Kohler (2011) is wrong stating the type locality as

“Neugranada.”

Distribution: Regions 1, 3, 6. Panama, Colombia, and

Venezuela. In Venezuela, widely distributed in open

lowlands of Maracaibo Lake Basin and Los Llanos

Region (Escalona et al. 2017).

Remarks: In Boana faber species group (Faivovich et

al. 2005). Many specimens in the Venezuelan collections

erroneously identified as Hyla or Hypsiboas crepitans

(FRR and CBA, personal observation: now Boana

xerophylla),; Escalona et al. (2017) distinguished

among B. pugnax and B. xerophylia in four Venezuelan

museums, expanding the previous known localities.

Selected references: La Marca (1996a); Chacon (2001);

Lynch and Suarez-Mayorga (2001); Tarano (2010);

Mendoza (2014); Infante-Rivero and Velozo (2015);

Escalona et al. (2017).

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~ _

128. Boana jimenezi. Quebrada Kawi, Gran Sabana, Bolivar.

Photo: César Barrio-Amoros.

129B. Boana lanciformis. Juvenile. Acequias river, Barinas.

Photo: César Barrio-Amoros.

130B. Boana lemai. Quebrada de Jaspe, Gran Sabana, Bolivar.

Photo: César Barrio-Amoros.

Boana punctata (Schneider 1799)

Syntypes: NHM 155 (two individuals)

Type locality: “Surinam.”

Distribution: Regions 3, 5. Widespread in South

America (Colombia, Venezuela, Ecuador, Peru, Bolivia,

Paraguay, Argentina, Brazil, Guyana, Suriname, French

Guiana, and Trinidad). Present in the open lowlands of

Venezuela, except in Maracaibo Lake Basin.

Remarks: In Boana punctata species group (Faivovich

et al. 2005).

Amphib. Reptile Conserv.

129A. Boana lanciformis. Male. Acequias river, Barinas. Photo:

César Barrio-Amoros.

130A. Boana lemai. Cuyuni river, Bolivar. Photo: César Barrio-

Amoros.

130C. Boana lemai. Juvenile. Quebrada Kawi, Gran Sabana,

Bolivar. Photo: César Barrio-Amoros.

Selected references: Duellman (1974a); Hoogmoed

(1979a); Gorzula and Sefiaris (1998); Barrio et al. (2000);

Sefiaris et al. (2014).

Boana roraima (Duellman and Hoogmoed 1992)

Holotype: BMNH 1979.560

Type locality: “North slope of Mt. Roraima (05°38'N,

60°44'W, elevation 1,480 m), Rupununi District,

Guyana.”

Distribution: Region 5. Guyana and Venezuela. In

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131A. Boana multifasciata. Santa Elena de Uairén, Bolivar. Photo:

César Barrio-Amoros.

132. Boana ornatissima. Karuay river, Gran Sabana, Bolivar.

Photo: Manuel Gonzalez.

133B. Boana pugnax. Ologa, southwestern Maracaibo Lake,

Zulia. Photo: César Barrio-Amoros.

Venezuela, restricted to the Eastern half of Bolivar State

(Barrio-Amoros et al. 2011).

Remarks: See variation from Guyana in MacCulloch and

Lathrop (2005). Barrio-Amoros et al. (2011) officially

reported the species from Venezuela, described the call,

and commented on distribution. Myers and Donnelly

(2008) described Hypsiboas angelicus from the summit

of Auyan-tepui. Barrio-Amoros et al. (2011) stated that

H. angelicus is a junior synonym of B. roraima. In Boana

benitezi species group (Faivovich et al. 2005).

Amphib. Reptile Conserv.

131B. Boana multifasciata. Amplexus. Chivaton, Gran Sabana,

Bolivar. Photo: César Barrio-Amoros.

24 r i ae a : 4

eo a)

133A. Boana pugnax. Male calling. San Vicente, Apure. Photo:

César Barrio-Amoros.

133C. Boana pugnax. Juvenile. Pagtiey river, Barinas. Photo:

César Barrio-Amoros.

Selected references: Duellman and Hoogmoed (1992);

MacCulloch and Lathrop (2005); Myers and Donnelly

(2008); Barrio-Amoros and Duellman (2009); Barrio-

Amoros et al. (2011c); Barrio-Amorés and Fuentes

(2012); Sefiaris et al. (2014).

Boana rhythmica (Sefiaris and Ayarzaguiiena 2002) *

Holotype: MHNLS 12957.

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Catalogue of the amphibians of Venezuela

Type locality: “Cerro Jaua, Parque nacional Jaua-

Sarisarifama, Bolivar State, Venezuela (4°49755”N,

64°25°54”W), 1,600 m.”

Distribution: Region 5. Endemic to Venezuela. Known

only from Cerro Jaua in Bolivar State.

Remarks: In Boana benitezi species group (Faivovich

et al. 2005).

Selected references: Sefiaris and Ayarzagiiena (2002);

Barrio-Amoros (2013).

Boana sibleszi (Rivero 1972)

Holotype: UPRM 3177.

Type locality: “Paso del Danto, La Escalera, entre

El Dorado y Santa Elena de Uairén, 1,300—1,400 m,

Serrania de Lema, Estado Bolivar, Venezuela.”

Distribution: Region 5. Venezuela and Guyana. In

Venezuela, distributed throughout the Eastern Guayana

uplands.

Remarks: In Boana punctata species group (Faivovich

et al. 2005). See variation from Guyana in MacCulloch

and Lathrop (2005).

Selected references: Rivero (1972); Hoogmoed (1979a);

Gorzula (1992); Duellman (1997); Gorzula and Sefiaris

(1998); MacCulloch and Lathrop (2005); Barrio-Amoros

and Duellman (2009); Sefiaris et al. (2014).

Boana tepuiana

(Barrio-Amoros and Brewer-Carias 2008)

Holotype: EBRG 4653.

Type locality: “Southern slope of Sarisarifiama-tepui,

Locality VI, Estado

Bolivar, Venezuela (4°25’N, 64°7’W), elevation 420 m.”

Distribution: Region 5. Eastern Guayana uplands.

Remarks: Must be in the Boana benitezi species group

(sensu Faivovich et al. 2005) by implication as it was

confused previously with B. benitezi, as it was already

observed by Frost (2019). See additional comments

under B. benitezi.

Selected references: Gorzula and Sefiaris (1998); Barrio-

Amoros and Brewer-Carias (2008); Sefiaris et al. (2014).

Boana wavrini (Parker 1936)

Type: IRSNB 1028.

Type locality: “Upper Orinoco, in the province of

Amazonas, Venezuela.”

Distribution: Region 4. Colombia, Venezuela and

Brazil. In Venezuela, restricted to the upper Orinoco, in

southern portion of Venezuelan Amazonas State.

Remarks: [n Boana semilineata species group (Faivovich

et al. 2005).

Selected references: Parker (1936); Ginés (1959);

Rivero (1961, 1967a,c); Hoogmoed (1990); La Marca

(1992); Gorzula and Sefiaris (1998).

Amphib. Reptile Conserv.

Boana xerophylla (Dumeril and Bibron 1841)

Lectotype: MNHN 652.

Type locality: Cayenne, French Guiana.

Distribution: Regions 1, 2, 3, 4, 5,6. Panama, Colombia,

Venezuela, Guyana, Suriname, French Guiana, and

northern Brazil. Widespread in Venezuela in open and

disturbed areas, 0O—2,300 m asl.

Remarks: In Boana faber species group (Faivovich

et al. 2005) by implication. Recently recovered from

synonymy of B. crepitans by Orrico et al. (2017). Several

authors gave evidence that B. crepitans was a complex

(Kluge 1979; Lynch and Suarez-Mayorga 2001; Gorzula

and Sefiaris 1999; Martins et al. 2009). The resurrection

of B. xerophylla as the northern South American

representative of the crepitans complex does not change

the panorama much. As reported previously by Barrio-

Amoros (2004), at least four different taxa were under

the name B. crepitans only in Venezuela. Now, all of

them pass automatically to be named B. xerophylla.

However, only the southeastern population (a blue morph

from rainforest lowlands in eastern Venezuela; Barrio-

Amoréos et al. 2011b) should be known as B. xerophylla

sensu stricto, aS it 1s conspecific with the French

Guiana population (Lescure and Marty 2000). All other

populations from Northern Venezuela, Colombia, and

Panama deserve further attention. So, in Venezuela three

more populations now being B. xerophylla represent

different taxa. One is from the Coastal Range and Andes

up to 2,300 m plus the upper Llanos; another is from the

western piedmont of the Cordillera de Mérida towards

the Maracaibo Lake Basin; another is a green morph

from the Gran Sabana (mentioned already as different by

Duellman 1997).

Selected references: Gunther (1858); Boettger (1892,

1893, 1896); Boulenger (1903); Lutz (1927); Aleman

(1952, 1953); Ginés (1959); Rohl (1959); Rivero (1961,

1963c, 1964a-d, 1967a, 1971a); Heatwole et al. (1965);

Tello (1968); Rivero and Esteves (1969); Donoso-

Barros and Ledn-Ochoa (1972); Staton and Dixon

(1977); Hoogmoed and Gorzula (1979); Rivero-Blanco

and Dixon (1979); Rada (1981); Péfaur and Diaz De

Pascual (1987); Ramo and Busto (1989, 1990); La Marca

(1991b “1994,” 1992, 1996a); Magdefrau et al. (1991);

Manzanilla et al. (1995); Péfaur and Pérez (1995); Barrio

(1996a); Yustiz (1996); Duellman (1997); Gorzula and

Sefiaris (1998); Lynch and Suarez-Mayorga (2002);

Barrio-Amoros (2006c); Barrio-Amoros and Brewer-

Carias (2008); Barrio-Amoros et al. (2011b); Mendoza

(2014); Sefiaris et al. (2014); Orrico et al. (2017).

Genus Ayloscirtus Peters, 1882

Type species: Hyloscirtus bogotensis Peters, 1882.

Remarks: Centrolenella estevesi Rivero, 1968 was

described from a creek close to Rio Albarregas north

of Merida city, at 2,400 m asl. This name was used as

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134. Boana punctata. Acequias river, Barinas. Photo: César

Barrio-Amoros.

135B. Boana roraima. Female. La Laja, Sierra de Lema, Bolivar.

Photo: César Barrio-Amoros.

whe =

Saban:

136A. Boana sibleszi. Spotted morph. Chivaton, Gran

Bolivar. Photo: César Barrio-Amoros.

136B. Boana sibleszi. Stripped morph. Chivaton, Gran Sabana,

Bolivar. Photo: César Barrio-Amoros.

Hyalinobatrachium by La Marca (1997), but more

recently Faivovich et al. (2005) revised the holotype

(MCZ 72498), reidentified the specimen as a juvenile

Hyloscirtus, and considered it as Hyloscirtus estevesi

(Frost 2018). Two species are reported from the same

area, both in the H. bogotensis group: H. jahni and H.

platydactylus.

From photographs of the holotype of Centrolenella

estevesi, the authors suspect it is a juvenile H. jahni,

due to snout shape projecting beyond the lip in ventral

Amphib. Reptile Conserv.

136C. Boana sibleszi. Plain morph, calling male. Paso el Danto,

La Escalera, Bolivar. Photo: César Barrio-Amoros.

view and absence of melanophores on fingers I and II.

Centrolenella estevesi Rivero, 1968 is thus considered a

junior synonym of Hyla jahni Rivero, 1961.

Ayloscirtus jahni (Rivero 1961)*

Holotype: UMMZ 46465.

Type locality: El Escorial, Mérida State, Venezuela.

Distribution: Endemic of cloud forest of Cordillera de

Merida (Andes), Merida State.

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137C. Boana tepuiana. Female. Pozo Esmeralda, El Pauji, Bolivar.

139A. Boana xerophylla. Night color. Parque Los Caobos,

Caracas. Photo: César Barrio-Amoros.

Remarks: Formerly in Hyloscirtus bogotensis species

group (Faivovich et al. 2005). Recently, Rojas-Runyaic

et al. (2018) recovered this species in his molecular

phylogeny as sister to all other Hyloscirtus. To promote

taxonomic stability and maintain taxonomy based on

monophyletic groups, they excluded H. jahni from the H.

bogotensis species group and created the monospecific

H. jahni species group, which is morphologically defined

by the presence of dermal spicules in adult males, and

by the high number of tooth rows in the larval oral disc.

Amphib. Reptile Conserv.

es a

137B. Boana tepuiana. Male paratype EBRG 4655. Southern

slopes of Sarisarifiama-tepui, Bolivar. Photo: César Barrio-Amoros.

138. Boana wavrini. Nifial, Casiquiare arm, Amazonas. Photo:

Fernando Rojas-Runjaic.

139B. Boana xerophylla. Day color. Imataca Forestal Reserve,

Delta Amacuro. Photo: César Barrio-Amoros.

Selected references: Rivero (1961, 1963b); La Marca

(1985a, 1994); Duellman (1989a).

Ayloscirtus japreria Rojas-Runjaic, Infante-Rivero,

Salerno et Meza-Joya, 2018

Holotype: MHNLS 19236.

Type locality: “Guacharaca Camp, Tetari Kopejoacha

creek, Rio Negro Basin, Sierra de Perijaé, Machiques de

Perija municipality, Zulia State, Venezuela (10°04’21.9N,

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139C. Boana xerophylla. Night color Chivaton, Gran Sabana,

Bolivar. Photo: César Barrio-Amoros.

139D. Boana xerophylla. Day color. El Playon, Caura river,

Bolivar. Photo: César Barrio-Amoros.

139E. Boana xerophylla. Day color. La Azulita, Mérida. Photo:

César Barrio-Amoros.

140. Boana sp. (cf. rufitela). MCZ 15369. Maracaibo, Zulia.

with permission of the Museum of Comparative Zoology, Harvard

University, Cambridge, Massachusetts, USA.

Amphib. Reptile Conserv.

139k Boat ET Night sie i ei Mérida, Photo:

César Barrio-Amoros.

SRE

way

141. Myersiohvla chamaeleo. Trice ie in life (AMNHA-131173).

Cerro de la Neblina, Amazonas. Photo: Roy McDiarmid.

72°51°16.7W; elevation 1,661 m asl.).”

Distribution: Known only from several localities on

the Venezuelan (Zulia State) and Colombian (Guajira

Department) slopes of the Sierra de Periya.

Remarks: In the Hy/oscirtus bogotensis species group

(Rojas-Runjaic et al. 2018).

Selected references: Rojas-Runjaic et al. (2018).

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Catalogue of the amphibians of Venezuela

Ayloscirtus lascinius (Rivero 1969)

Holotype: MCZ 65901.

Type locality: “Tabor, above de Delicias, paramo of

Tama, Tachira, Venezuela.”

Distribution: Region 1. Andes of Colombia (Norte de

Santander Department) and Venezuela (Tachira and

Merida States). Recently reported from Sierra de Perija

in Zulia State (Rojas-Runjaic et al. 2016).

Remarks: In Hy/loscirtus bogotensis species group

(Faivovich et al. 2005).

Selected references: Rivero (1969d); Duellman (1989a);

Rojas-Runjaic et al. (2016).

HAyloscirtus platydactylus (Boulenger 1905)

Holotype: BM 1947.2.13.14.

Type locality: “Merida, Andes of Venezuela.”

Distribution: Region 1. Andes of Colombia (Norte de

Santander Department) and Venezuela (Cordillera de

Merida and Sierra de Perija).

Remarks: In Hy/loscirtus bogotensis species group

(Faivovich et al. 2005).

Selected references: Boulenger (1905); Ginés (1959);

Rivero (1961, 1963b, 1969d); Duellman (1972a, 1979b,

1989a); La Marca (1985a, 1994b); Péfaur and Diaz De

Pascual (1987); Barrio-Amoros (2010a).

Genus Myersiohyla Faivovich, Haddad, Garcia, Frost,

Campbell and Wheleer, 2005

Type species: Myersiohyla neblinaria Faivovich,

McDiarmid and Myers, 2013. Faivovich et al. (2013)

stated the original type species of Myersohyla was Hyla

inparquesi, Ayarzagiena and Sefiaris, 1994 due to a

misidentified specimen of M. “inparquesi’ from Neblina,

which 1s now correctly identified as M. neblinaria.

Myersiohyla aromatica

(Ayarzagtiena and Sefiaris 1994)*

Holotype: MHNLS 12510

Type locality: “Cumbre del Tepuy Huachamacari,

Estado Amazonas, Venezuela (03°50’N, 65°45’O) 1,700

m snm.”

Distribution: Region 5. Endemic to Huachamacari, a

tepui in Amazonas State.

Selected references: Ayarzagiiena and Sefiaris (1993);

McDiarmid and Donnelly (2005); Faivovich et al. (2013).

Myersiohyla chamaeleo

Faivovich, McDiarmid and Myers, 2013*

Holotype: AMNH A-131173

Type locality: Venezuela: Departamento Amazonas:

Cerro de la Neblina: Camp I, 1,820—1,880 m.

Distribution: Region 5. Known from type locality and

Amphib. Reptile Conserv.

‘he

surroundings at Cerro de la Neblina in the southern

border of Amazonas State.

Selected references: Faivovich et al. (2013).

Myersiohyla inparquesi

(Ayarzagiiena and Sefiaris 1994)*

Holotype: MHNLS 12338.

Type locality: “Cumbre del Tepuy Marahuaca Sur,

Estado Amazonas, Venezuela (3°40’N, 65°27’O) 2,600

m snm.”

Distribution: Region 5. Endemic to Marahuaka, a tepui

in central Estado Amazonas.

Selected references: Ayarzagiiena and Sefiaris (1994);

McDiarmid and Donnelly (2005).

Myersiohyla loveridgei (Rivero 1961)*

Holotype: MCZ 28565.

Type locality: Pico Culebra, Cerro Duida, Amazonas

State, Venezuela.

Distribution: Known only from type locality.

Remarks: Region 5. Faivovich et al. (2013) noted the

similarity between Myersiohyla loveridgei and M.

aromatica, suggesting the possible synonymy of the

latter.

Selected references: Rivero (1961, 1963d, 1964b,

1971b); La Marca and Smith (1982b); Faivovich et al.

(2013).

Myersiohyla neblinaria

Faivovich, McDiarmid and Myers, 2013*

Holotype: USNM 562071

Type locality: “Venezuela: Departamento Amazonas:

Cerro de la Neblina: Camp VII, 1,730 m.”

Distribution: Region 5. Known from type locality and

surroundings at Cerro de la Neblina, in southern border

of Amazonas State. Probably also on the Brazilian side.

Selected references: Faivovich et al. (2013).

Subfamily Dendropsophinae Fitzinger, 1843

Genus Dendropsophus Fitzinger, 1843

Type species: Hyla frontalis Daudin, 1800 (= Rana

leucophyllata Beireis, 1783), by original designation.

Remarks: The genus was resurrected by Faivovich et al.

(2005) to comprise Hy/a species that have or are suspected

to have 30 chromosomes. Duellman et al. (2016) included

two genera in the subfamily Dendropsophinae, only one

being present in Venezuela: Dendropsophus.

Dendropsophus amicorum (Miyjares-Urrutia 1998)*

Holotype: USNM 216677.

Type locality: Cerro Socopd, 84 km al NO de Carora,

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_ Pa eS | ee rl =. =

142. Myersiohyla neblinaria. Holotype in life (USNM 562071).

Cerro de la Neblina, Amazonas. Photo: Roy McDiarmid.

144A. Dendropsophus luteoocellatus. Male calling. Calderas,

Barinas. Photo: César Barrio-Amoros.

145. Dendropsophus marmoratus. Male. Imataca, Delta Amacuro.

Photo: César Barrio-Amoros.

Estado Falcon, Venezuela (10°28’N, 70°48’O).

Distribution: Region 2. Known from type locality.

Remarks: Known only by holotype. No additional

specimens collected after it in 1968. Faivovich et al.

(2005) assigned this species to Dendropsophus but not to

any species group. Assessing this apparently rare species

is important, as it could be confused with a common

species like “Dendropsophus aff. minutus,” which would

have implications for conservation. Currently considered

Amphib. Reptile Conserv.

143. Dendropsophus battersbyi.

Caracas. Photo: Salvador Carranza.

Holotype BM _ 53.2.4.165.

144B. Dendropsophus luteoocellatus. Female. Calderas, Barinas.

Photo: César Barrio-Amoros.

ee Rca iat oa i]

Mec Pe

146A. Dendropsophus meridensis. Male calling, Los Suarez, way

from Mérida to la Azulita, Mérida. Photo: César Barrio-Amoros.

: te k

Critically Endangered by the Venezuelan Red Book

(Mijares-Urrutia et al. 2015).

Selected references: Mijares-Urrutia (1998); Faitvovich

et al. (2005); Mijares-Urrutia et al. (2015).

Dendropsophus battersbyi (Rivero 1961)*

Holotype: BM 53.2.4.165.

Type locality: Caracas, Venezuela.

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Catalogue of the amphibians of Venezuela

Distribution: Region 2. Only known from type locality.

Remarks: Not collected since original description. If this

is a valid species, it possibly disappeared from the Valley

of Caracas due to quick expansion of the city. Miyares-

Urrutia (1998) mentioned this species is apparently

related to Dendropsophus amicorum and D. minutus

(sensu lato). Faivovich et al. (2005) did not assign it to

any species group. A thorough examination of holotype by

CBA leades authors to believe Dendropsophus battersbyi

is a valid species and a member of the D. minutus species

group (sensu Gehara et al. 2014); alternatively, it could

be conspecific with populations of D. aff. minutus from

around Caracas that deserve further attention. A detailed

revision of the species/populations of Dendropsophus

minutus species group in the Venezuelan Andes and

the Cordillera de La Costa is required to clarify the

taxonomic status of this and close species.

Selected references: Rivero (1961, 1964a); Mijares-

Urrutia (1998); Faivovich et al. (2005); Barrio-Amoros

and Torres (2010); Barrio-Amoros (2013).

Dendropsophus luteoocellatus (Roux 1927)*

Lectotype: NHMB 3900.

Type locality: “El Mene, Provincia Falcon, Venezuela.”

Distribution: Regions 1, 2. Endemic to both sides of

Cordillera de Mérida and Cordillera de la Costa.

Remarks: In the Dendropsophus parviceps species

group sensu Faivovich et al. (2005).

Selected references: Roux (1927); Ginés (1959); Rivero

(1961, 1963c, 1969c, 1971a); Duellman and Crump

(1974); Yustiz (1996); Barrio-Amoros (2010a); Barrio-

Amoros and Molina (2010).

Dendropsophus marmoratus (Laurenti 1768)

Type: Unknown.

Type locality: “Surinam.”

Distribution: Regions 4, 5. Amazonian lowlands of

Colombia, Ecuador, Peru, Bolivia, Brazil, Venezuela,

Guyana, Suriname, and French Guiana. In Venezuela,

known from a few localities in Amazonas and Bolivar

States, south of the Orinoco River.

Remarks: In the Dendropsophus marmoratus species

group sensu Faivovich et al. (2005).

Selected references: Rivero (1961, 1964b); Bokermann

(1964).

Dendropsophus meridensis (Rivero 1961)*

Holotype: MCZ 2527.

Type locality: “Mérida, 1,630 m.”

Distribution: Region 1. Cordillera de Mérida

(Venezuelan Andes) in Mérida and Tachira States (see

current map by Armesto et al. 2015).

Remarks: So far, differences between Dendropsophus

meridensis and D. pelidnus are almost nonexistent or

Amphib. Reptile Conserv.

insignificant. Dendropsophus meridensis is known to

inhabit stagnant waters, but specimens of D. pelidnus

that Duellman (1989) obtained in Tachira were from a

stream. Guarnizo et al. (2012) found a maximum genetic

distance of 0.9% between central Cordillera de Mérida

specimens and others from near the type locality of

D. pelidnus, which is not significant. Aside from this

molecular evidence, calls and morphology should be

compared between these populations before proposing

a synonymy. Armesto et al. (2015) provide an updated

distribution map and model of potential distribution in

the Venezuelan Andes. If D. pelidnus is a junior synonym

of D. meridensis, then the latter would be present in

Colombia.

Selected references: Rivero (1961, 1963b, 1964d);

Duellman (1989a); Miares-Urrutia (1990b); La Marca

(1991b “1994,” 1992): Guarnizo et al. (2012); Armesto

et al. (2015).

Dendropsophus microcephalus (Cope 1886)

Syntypes: USNM 13473 (two specimens, lost).

Type locality: “Chiriqui,” Panama.

Distribution: Regions 3, 6. Widespread from

southeastern Mexico to northern South America

(Colombia, Venezuela, Guyana, Suriname, French

Guiana, Brazil, and Trinidad). In Venezuela, widely

distributed in open lowlands.

Remarks: Hy/la misera Werner, 1903 from Caracas,

Venezuela (holotype IRSNB 4549) (as Dendropsophus

miserus) Would be the name for Venezuelan populations

if, as Duellman (1970) suggested, the subspecies

currently recognized (as D. microcephalus miserus for

the Venezuelan and northern Colombian populations) is

proved to be a full species. Fouquette (1968), Duellman

(1970), and Savage (2002) already recognized the

differences among Central American and South American

populations. Molecular work is needed to assess its final

taxonomic identity. In the Dendropsophus microcephalus

Species group sensu Faivovich et al. (2005).

Selected references: Lutz (1927); Rivero (1961, 1963c,

1964d); Heatwole et al. (1965); Duellman and Fouquette

(1968); Fouquette (1968); Duellman (1970, 1974a,

1997); Staton and Dixon (1977); Hoogmoed and Gorzula

(1979); Rivero-Blanco and Dixon (1979); Rada (1981);

Yustiz (1996); Gorzula and Sefiaris (1998); Savage

(2002); Tarano (2010); Barrio-Amoros et al. (2011b).

Dendropsophus minusculus (Rivero 1971)

Holotype: UPR-M 3377.

Type locality: Nirgua, Yaracuy State, Venezuela.

Distribution: Regions 2, 3, 4, 5. Colombia, Venezuela,

Brazil, Guiana, Suriname, and French Guiana. Widespread

in Venezuela, from northern lowlands of Cordillera de la

Costa, throughout Los Llanos, to Orinoquian lowlands.

Remarks: In the Dendropsophus microcephalus species

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= =

146B. Dendropsophus meridensis. Los Suarez, way from Mérida

to la Azulita, Mérida. Photo: César Barrio-Amoros.

147B. Dendropsophus microcephalus. Arassari rafting camp,

Acequias, Barinas. Photo: César Barrio-Amoros.

148B. Dendropsophus minusculus. Plain morph. San Vicente,

Apure. Photo: César Barrio-Amoros.

group sensu Faivovich et al. (2005).

Selected references: Rivero (1971la); Hoogmoed and

Gorzula (1979); Rivero-Blanco and Dixon (1979); Ramo

and Busto (1989, 1990); Duellman (1997); Gorzula and

Sefiaris (1998); Tarano (2010); Barrio-Amoros et al.

(2011b); Sefiaris et al. (2014).

Dendropsophus minutus (Peters 1872)

Syntypes: ZMB 7456.

Type locality: Nova Friburgo, Rio de Janeiro, Brazil.

Distribution: Regions 1, 2, 4, 5. Widespread in

Amphib. Reptile Conserv.

147A. Dendropsophus microcephalus. Male calling. Chururt,

Tachira. Photo: César Barrio-Amoros.

148A. Dendropsophus minusculus. Male calling. El Palmar,

Bolivar. Photo: César Barrio-Amoros.

148C. Dendropsophus minusculus. Stripped morph. North of

Santa Elena de Uairén, Bolivar. Photo: César Barrio-Amoros.

Venezuela, with populations from the Coastal chain, the

Andes, the Amazon, and Guiana Shield.

Remarks: Gehara et al. (2014) shows this name hides

a complex of species, with at least four putative species

in Venezuela. The name Hyla goughi Boulenger, 1911

was mentioned to be available for populations clustered

together with Trinidadian populations. However,

CBA examined the type of Hyla goughi (BMNH

1947.2.13.12) and it is considered to be conspecific with

Hyla microcephala Cope, 1886. A cotype of H. goughi,

BMNH 1947.2.13.83, is a subadult Dendropsophus aff.

minutus. The status of other populations once attributed

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149A. Dendropsophus aff. minutus. Los Alcaravanes, Calderas,

Barinas. Photo: César Barrio-Amoros.

149C. Dendropsophus aff. minutus. Chivaton, Gran Sabana,

Bolivar. Photo: César Barrio-Amoros.

150. Dendropsophus parviceps. Yasuni, Ecuador. Photo: César

Barrio-Amoros.

to Dendropsophus minutus still need to be clarified.

Selected references: Rivero (1961, 1964b, 1971b);

Heatwole et al. (1965); Rivero and Esteves (1969):

Hoogmoed and Gorzula (1979); Donnelly and Myers

(1991); Magdefrau et al. (1991); Duellman (1997);

Gorzula and Sefiaris (1998); Barrio-Amords and

Duellman (2009); Barrio-Amorés (2010a); Barrio-

Amoréos et al. (2011b); Gehara et al. (2014); Sefiaris et

al. (2014).

Dendropsophus parviceps (Boulenger 1882)

Amphib. Reptile Conserv.

149B. Dendropsophus aff. minutus. Sierra de Lema, Bolivar.

Photo: César Barrio-Amoros.

149D. Dendropsophus aff. minutus. Mapurital, Caripe, Monagas.

Photo: Diego A. Flores.

Pt in

p eee . A

ge TS Lee ee

elidnus. Tama, Tachira Photo: Orlando

151. Dendropsophus P

Armesto.

Holotype: BM 1947.2.13.51

Type locality: “Sarayacu,” Provincia Pastaza, Ecuador.

Distribution: Region 4, 5. Widely distributed in Amazon

Basin: Colombia, Ecuador, Peru, Bolivia, Brazil,

and Venezuela. In Venezuela, only known from three

localities: lower Casiquiare (McDiarmid and Paolillo

1988), Surumoni in Amazonas State (MHNLS 20906-

20910), and Guaquinima-tepui in Bolivar State (Schltiter

and Magdefrau 1991).

Remarks: In Dendropsophus parviceps species group

(Faivovich et al. 2005).

Selected references: McDiarmid and Paolillo (1988);

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Magdefrau et al. (1991); Schltiter and Magdefrau (1991);

Amézquita and Hodl (2004).

Dendropsophus pelidnus (Duellman 1989)

Holotype: KU 181109.

Type locality: “Betania (07°30’N, 72°27’W, 2,220 m),

Estado Tachira, Venezuela.”

Distribution: Region |. Departments of Santander and

Boyaca in Colombia, and Tachira State in Venezuela,

2,200-—3,000 m asl.

Remarks: See comments under Dendropsophus

meridensis. If D. pelidnus is shown to be a junior

synonym of D. meridensis, then distribution of latter

would reach Colombia. The specific name ending is

amended to be consistent with the masculine genus,

from Greek adjective pelidnos (Duellman 1989a). In

Dendropsophus labialis species group (Duellman 1989a;

Faivovich et al. 2005).

Selected references: Duellman (1989a); Guarnizo et al.

(2012); Armesto et al. (2014).

Dendropsophus sarayacuensis (Shreve 1935)

Holotype: MCZ 19729.

Type locality: Sarayacu, Provincia Pastaza, Ecuador.

Distribution: Regions 4, 5. Amazon Basin, in Ecuador,

Peru, Bolivia, Brazil, and Venezuela. Only record for

Venezuela from base of Cerro La Neblina in the southern

extreme of Amazonas State (McDiarmid and Paolillo

1988).

Remarks: In Dendropsophus leucophyllatus species

group (Faivovich et al. 2005).

Selected references: Duellman (1974a); McDiarmid and

Paolillo (1988).

Dendropsophus yaracuyanus

(Miares-Urrutia and Rivero 2000)*

Holotype: EBRG 3311.

Type locality: “Los Bacos, Municipio Bolivar, Sierra de

Aroa, Estado Yaracuy, Venezuela.”

Distribution: Region 2. Endemic to Sierra de Aroa in

Yaracuy State.

Remarks: Not previously assigned to any species group

(Faivovich et al. 2005), herein assigned to D. parviceps

group, due to overall similarity with the other species in

this group.

Selected references: Mijares-Urrutia and Rivero (2000).

Subfamily Lophyohylinae Miranda-Ribeiro, 1926

Remarks: Duellman et al. (2016) included five

genera present in Venezuela in this subfamily: Apara-

sphenodon, Osteocephalus, Phytotriades, Tepuihyla, and

Trachycephalus.

Amphib. Reptile Conserv.

Genus Aparasphenodon Miranda-Ribeiro, 1920

Type species: Aparasphenodon brunoi Miranda Ribeiro,

1920, by monotypy.

Aparasphenodon venezolanus (Mertens 1950)

Holotype: SMF 22168.

Type locality: San Fernando de Atabapo, Amazonas

State, Venezuela.

Distribution: Region 4. Amazonian lowlands of

Venezuela and nearby Colombia.

Selected references: Mertens (1950, 1967); Ginés

(1959); Rivero (1961, 1964b); Paolillo and Cerda (1981);

Lynch and Vargas-Ramirez (2000).

Genus Osteocephalus Steindachner, 1862

Type species: Osteocephalus taurinus Steindachner,

1862, by subsequent designation of Kellogg (1932).

Osteocephalus helenae (Ruthven 1919)

Holotype: UMMZ 52681.

Type locality: Valley of the Demerara River, Dunoon,

Guyana.

Distribution: Region 5. Widely distributed in lowlands

of Amazon Region (Bolivia, Peru, Colombia, Brazil,

Colombia, Venezuela, Guyana, and French Guiana). In

Venezuela, known from several localities in Bolivar and

Delta Amacuro States (Barrio-Amoros 1998, Sefiaris and

Ayarzagtiena 2004).

Remarks: Osteocephalus helenae is the name that

must apply to southern Venezuela’s green, tuberculated,

medium-sized Osteocephalus. Specimens of this

species in this region have been known under two

names: Osteocephalus buckleyi (Boulenger 1882) with

distribution now restricted to Upper Amazon in Peru,

Colombia, and Ecuador (Jungfer et al. 2013); and

Osteocephalus cabrerai (Cochran and Goin 1970), first

reported from Venezuela by Gorzula and Sefiaris (1998).

All specimens examined by Jungfer et al. (2013) from the

Guiana Shield fall in the same clade, and only name O.

helenae 1s applicable. In Osteocephalus buckleyi species

group (Jungfer et al. 2013).

Selected references: Cochran and Goin (1970); Trueb

and Duellman (1971); Duellman and Mendelson (1995);

Gorzula and Sefiaris (1998); Sefiaris and Ayarzagtiena

(2004); Jungfer et al. (2013).

Osteocephalus leprieurii (Dumeril and Bibron 1841)

Holotype: MNHNP 4629.

Type locality: “Cayenne,” French Guiana.

Distribution: Regions 4, 5. Widely distributed in Guiana

Shield (Brazil, Venezuela, Guyana, Suriname, and French

Guiana). In Venezuela, known from several localities in

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152. Dendropsophus sarayacuensis. Sarayacu, Ecuador. Photo:

William E. Duellman.

153B. Dendropsophus yaracuyanus. Sierra de Aroa, Yaracuy.

Photo: Fernando Rojas-Runjaic.

155A. Osteocephalus helenae. Imataca, Delta Amacuro. Photo:

Fernando Rojas-Runjaic.

Amazonas and Bolivar States.

Remarks: In Osteocephalus leprieurii species group

(Jungfer et al. 2013). Osteocephalus ayarzaguenai

Gorzula and Sefiaris, 1997 is a junior synonym (Jungfer

and Hodl 2002).

Selected references: Rivero (1971la); Trueb and

Duellman (1971); Gorzula and Sefiaris (1996, 1998),

Duellman (1997); Barrio and Fuentes (2000a); Jungfer

and Hodl (2002); Sefiaris et al. (2009, 2014); Jungfer et

al. (2013).

Amphib. Reptile Conserv.

153A. Dendropsophus yaracuyanus. Sierra de Aroa, Yaracuy.

Photo: Walter Schargel.

154. Aparasphenodon venezolanus. Guainia, Colombia. Photo:

John D. Lynch.

155B. Osteocephalus helenae. Santa Maria de Erebato, Bolivar.

Photo: César Barrio-Amoros.

Osteocephalus taurinus Steindachner, 1862

Type: NHMW 16492.

Type locality: Barra do Rio Negro, Manaos, Amazonas,

Brazil.

Distribution: Region 5. Amazonian species, distributed

in southern, central, and upper Amazon in Brazil,

Peru, Ecuador, Bolivia, Colombia, and the Guianas. In

Venezuela all populations known as O. taurinus must be

revised. Jungfer et al. (2013) identified two candidate

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species under this name from Venezuela, one in NW

Amazonas State, and another from the extreme south

Amazonas State (Appendix 1). Jungfer et al. (2013)

restricts the presence of O. taurinus sensu stricto out

of Venezuela. However, populations of O. “taurinus”

from southeastern Venezuela would remain without

identification. For now, authors restrict distribution of

O. aff. taurinus to the lowlands and uplands of Bolivar

State.

Remarks: A_ well-recognized species complex. In

Venezuela at least three putative species are reported

under this name (Jungfer et al. 2013; Appendix 1).

Selected references: Ginés (1959); Rivero (1961,

1964b); Trueb and Duellman (1971); Donnelly and

Myers (1991); Duellman (1997); Gorzula and Sefiaris

(1998); Lynch and Vargas-Ramirez (2000); Barrio-

Amoros and Brewer-Carias (2008); Barrio-Amoros et al.

(2011b); Jungfer et al. (2013); Sefiaris et al. (2014).

Genus Phytotriades Jowers, Downie and Cohen, 2008

Type species: Amphodus auratus Boulenger, 1917, by

original designation.

Phytotriades auratus (Boulenger 1917)

Type: BMNH 1917.9.27.39-41

Type locality: “Mount Tucutche, a little above 3,000 feet

altitude,” Trinidad.

Distribution: Region 2. Only known from Trinidad

island and Cerro El Humo in Peninsula de Paria

(northwestern Venezuela), Sucre State. Probably in other

localities of Paria over 1,000 m where the giant bromeliad

Glomeropitcairnia erectiflora (its microhabitat) 1s

present.

Remarks: Considered as Critically Endangered by

current Venezuelan Red Book (De Freitas and Rivas

2015).

Selected references: Faivovich et al. (2005); Jowers et

al. (2009); De Freitas and Rivas (2015); Rivas and De

Freitas (2015).

Genus Tepuihyla

Ayarzagtena, Sefiaris and Gorzula, 1993

Type species: Hyla rodriguezi Rivero, 1968, by original

designation.

Remarks: The modern concept of 7epuihyla includes

several species ranging from the Upper Amazon

(Tepuihyla shushupe and T. tuberculosa) to the Guiana

Shield (Ron et al. 2016), some only occurring at the

tepui summits, and few others from uplands to lowlands.

Several new phylogenetic studies suggest that tepui

dwellers evolved quite recently from a common ancestor,

and not a long time ago as usually suspected for endemics

of high tepuis (Kok et al. 2012; Salerno et al. 2012).

Although Ayarzagtiena et al. (1993) defined the genus

Amphib. Reptile Conserv.

Tepuihyla as masculine, Myers and Stothers (2006)

clarified that Hyla is feminine so, 7epuihyla cannot be

assumed as masculine.

Tepuihyla aecii

(Ayarzagtiena, Sefiaris and Gorzula 1993)*

Holotype: MHNLS 12014.

Type locality: “Cumbre Sur del Monte Duida, Estado

Amazonas. Venezuela (3°19’N, 65°38’W). 2,150 m

snm.”

Distribution: Region 5. Endemic to Cerro Duida,

Amazonas State, Venezuela.

Selected references: Ayarzagtiena et al. (1993a,b).

Tepuihyla edelcae

(Ayarzagtiena, Sefiaris and Gorzula 1993)*

Holotype: MHNLS 10626.

Type locality: “Auyan-tepuy, 10.8 km al este del Salto

Angel. Estado Bolivar. Venezuela (5° 58’N, 62° 29° W).

1,970 m snm.”

Distribution: Region 5. Endemic to Auyan-tepul,

Bolivar State, Venezuela.

Remarks: Specimens referred as Tepuihyla edelcae from

the Chimanta massif correspond to 7’ obscura (Kok et

al. 2015).

Selected references: Ayarzagtiena et al. (1992a,b);

Gorzula (1992); Gorzula and Sefiaris (1998); McDiarmid

and Donnelly (2005); Barrio-Amoros and Fuentes

(2012); Kok et al. (2012, 2015); Salerno et al. (2012,

2014); Sefiaris et al. (2014).

Tepuihyla exophthalma (Smith and Noonan 2001)

Holotype: UG-CSBD HA-722.

Type locality: 30 km SE Imbaimadai, Mazaruni-Potaro

District, Guyana, ca. 585 m.

Distribution: Region 5. Venezuela and Guyana. In

Venezuela, only from high sector of Sierra de Lema.

Remarks: Described as Osteocephalus exophthalmus by

Smith and Noonan (2001) from Mazaruni-Potaro district

in Guyana. Later Kok and Kalamandeen (2008) reported

a second population from Kaieteur National Park also

in Guyana. Barrio-Amoros et al. (2010h) reported the

first Venezuelan specimens from the upper sector of La

Escalera in Sierra de Lema, Bolivar. Jungfer et al. (2013)

passed the species to 7epuihyla based on genetic data,

though this eliminated one of the most characteristic

morphological synapomorphies of Tepuihyla: the

absence of webbing between toes I and II. Osteocephalus

phasmatus MacCulloch and Lathrop, 2005 is a junior

synonym of 7. exophthalma according to Jungfer et al.

(2013).

Selected references: MacCulloch and Lathrop (2005);

Kok and Kalamandeen (2008); Barrio-Amoros et al.

(2010h); Jungfer et al. (2013); Sefiaris et al. (2014).

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156A. Osteocephalus leptreuirii. Cuyuni river, Bolivar. Photo:

César Barrio-Amoros.

157. Osteocephalus aff. taurinus. Male. La Laja, Sierra de Lema,

Bolivar. Photo: César Barrio-Amoros.

159, Tepuihyla edelcae. Auyan-tepui summit. Photo: César

Barrio-Amoros.

Tepuihyla luteolabris

(Ayarzagtena, Sefiaris and Gorzula 1993)*

Holotype: MHNLS 9376.

Type locality: “Tepuy Marahuaca Norte, Estado

Amazonas. Venezuela. (3° 45’N, 65° 30’W). 2,550 m.”

Distribution: Region 5. Endemic to Cerro Marahuaka, a

tepui in Amazonas State, Venezuela.

Remarks: Zepuihyla celsae was described from Cerro

Galicia in Falcon State by Miyares-Urrutia et al. (1999).

Jugfer et al. (2013) synonymized it with 7: /uteolabris,

based on communication by colleagues, but not a formal

Amphib. Reptile Conserv.

156B. Osteocephalus lepreuirii. “Osteocephalus ayarzaguenai”

pattern. Cuyuni river, Bolivar. Photo: César Barrio-Amoros.

158. Phytotriades auratus. Peninsula de Paria, Sucre. Photo:

Gilson Rivas.

160A. Tepuihyla exophthalma. Upper sector of La Escalera, Sierra

de Lema, Bolivar. Photo: César Barrio-Amoros.

comparison of type material. However, current authors

agree with that decision.

Selected references: Ayarzagtiena et al. (1993a,b);

Myyares et al. (1999); Jungfer et al. (2013).

Tepuihyla obscura

Kok, Ratz, Tagelaar, Aubret and Means, 2015*

Holotype: IRSNB 4192.

Type locality: “Summit of Chimanta-tepui (5°19’27”N,

62°12710’’W, 2,224 m asl.)”

Distribution: Region 5. Widespread in Chimanta massif

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(Bolivar State), on Abakapa, Akopan, Amuri, Apakara,

Chimanta, and Churi tepuis, 1,800—2,600 m asl.

Remarks: TZepuihyla edelcae was described from

Auyan-tepui and referred to date also to Chimanta massif

(Ayarzagtiena et al. 1992a), but these two populations

were recognized as a different species by Kok et al.

(2012) and Salerno et al. (2012, 2014).

Selected references: Salerno et al. (2014); Kok et al.

(2015).

Tepuihyla rodriguezi (Rivero 1968)

Holotype: MCZ 64740.

Type locality: Paso del Danto, Region de La Escalera,

entre El Dorado y Santa Elena de Uairén, 1,300—1,400 m,

Serrania de Lema, Estado Bolivar, Venezuela.

Distribution: Region 5. With synonymizing of Tepuihyla

galani, T: talbergae, and T. rimarum with T. rodriguezi

(Jungfer et al. 2013, Kok et al. 2015), the distribution of

the latter now extends from Paso el Danto in the northern

versant of Sierra de Lema (type locality) through the

higher sectors of the Gran Sabana, the foothills of

Guadacapiapu tepui (type locality of 7. galani), atop of

Ptari-tepui (type locality of 7. rimarum), and as low as

Kaieteur National Park in Guyana (type locality of 7.

talbergae), altitudinal range 366—2,440 m.

Remarks. Jepuihyla galani (Ayarzagtiena, Sefiaris et

Gorzula, 1993), T. talbergae Duellman and Yoshpa, 1996,

and 7! rimarum (Ayarzagtena, Sefiaris and Gorzula 1993)

recently synonymized with 7: rodriguezi by Jungfer et al.

(2013) and Kok et al. (2015).

Selected References: Rivero (1968); Ayarzagtiena et

al. (1992a,b); Duellman and Yoshpa (1996); Duellman

(1997); Gorzula and Sefiaris (1998); Kok and

Kalamandeen (2008); Jungfer et al. (2013); Sefiaris et al.

(2014); Kok et al. (2015).

Genus Trachycephalus Tschudi, 1838

Type species: 7rachycephalus nigromaculatus Tschudi,

1838, by monotypy.

Trachycephalus resinifictrix (Goeldi 1907)

Holotype: BM 1947.2.23.24.

Type locality: “Mission of San Antonio do Prata, at the

River Maracana,” Brazil.

Distribution: Regions 4, 5. Eastern part of Brazilian

Amazon Basin, Venezuela, Guyana, Suriname, and

French Guiana. In Venezuela known from the base

of Cerro La Neblina in Amazonian lowlands of south

Amazonas State, from Delta of the Orinoco in Delta

Amacuro State and from the upper Cuyuni area.

Selected references: McDiarmid and Paolillo (1988);

Sefiaris and Ayarzagtiena (2004 “2002”); Sefiaris et al.

(2009).

Amphib. Reptile Conserv.

Trachycephalus typhonius (Linnaeus 1758)

Holotype: UMMZ 134.

Type locality: “America.”

Distribution: Regions 1, 2, 3, 4, 5, 6. Widespread from

Mexico through Central America and South America

to Paraguay and Northern Argentina; also, in Trinidad

and Tobago. Widespread in lowland Venezuela, even in

disturbed areas.

Remarks: This taxon recently changed genus and

species. Most commonly known as_ Phrynohyas

venulosa, until Faivovich et al. (2005) transferred it to

the genus Trachycephalus, as T: venulosus. Then Lavilla

et al. (2010) elucidated the long controversial nomen

typhonius (used for a complex of bufonids as Bufo

typhonius, and for an Asian ranoid) to be applied as

Trachycephalus typhonius.

Selected references: Lutz (1927); Duellman (1956,

1971b, 1997); Rohl (1959); Rivero (1961, 1964a-d),;

Tello (1968); Rivero and Esteves (1969); Staton and

Dixon (1977); Hoogmoed and Gorzula (1979); Rivero-

Blanco and Dixon (1979); Ramo and Busto (1989, 1990);

Barrio (1996a); Yustiz (1996); Lavilla et al. (2010);

Tarano (2010); Barrio-Amoros et al. (2011).

Subfamily Pseudinae Fitzinger, 1843

Remarks: The long-recognized family Pseudidae was

passed to subfamily of Hylidae by Duellman (2001) and

supported by Haas (2003) based on larval characters.

Darst and Cannatella (2004) recovered Pseudis and

Scarthyla within hyaline frogs. Faivovich et al. (2005)

placed Pseudis in their tribe Dendropsophini. Garda

and Cannatella (2007) recognized the group with an

unranked name, Pseudae Fitzinger, 1843, for Pseudis

and Lysapsus. Aguiar et al. (2007) placed Lysapsus in

Pseudis, followed by Pyron and Wiens (2011), but not

by Wiens et al. (2010) who considered a monophyletic

Pseudis sensu stricto (not including Lysapsus). Duellman

et al. (2016) recovered Pseudinae as a subfamily of

Hylidae.

Genus Pseudis Wagler, 1830

Type species: Rana paradoxa Linnaeus, 1758, by

monotypy.

Pseudis paradoxa (Linnaeus 1758)

Syntypes: MHRM 114-148.

Type locality: Surinam.

Distribution: Regions 2, 3, 4, 5, 6. Widely distributed

from northern South America (Colombia, Venezuela,

Trinidad, Guyana, Suriname, French Guiana) throughout

Brazil to Peru and Bolivia. In Venezuela, in lowlands of

Maracaibo Lake Basin, Llanos, Guayana, and Delta of

the Orinoco.

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Catalogue of the amphibians of Venezuela

160B. Tepuihyla exophthalma. Upper sector of La Escalera, Sierra

de Lema, Bolivar. Photo: César Barrio-Amoros.

162A. Tepuihyla rodriguezi. Luepa, Gran Sabana, Bolivar. Photo:

César Barrio-Amoros.

163. Trachycephalus resinifictrix. Yasuni, Ecuador. Photo: Pau

Cardellach.

Remarks: Gallardo (1961) described different taxa

(subspecies) of P. paradoxa based mainly on chromatic

and biogeographic characters. None of them are regarded

currently as valid (Garda et al. 2010).

Selected references: Lutz (1927); Ginés (1959);

Gallardo (1961); Rivero (1961, 1964a); Heatwole et al.

(1965); Rivero and Esteves (1969); Staton and Dixon

(1977); Hoogmoed and Gorzula (1979); Gremone et al.

Amphib. Reptile Conserv.

vat Po, :

161. Tepuihyla obscura. Churi-tepui, Chimanta Massif, Bolivar.

Photo: César Barrio-Amoros.

»\A

162B. Tepuihyla rodriguezi. Guadacapiapu savannas (type locality of

syn. 7’ galani), Gran Sabana, Bolivar. Photo: César Barrio-Amoros.

> i .

—_

ae F,

“Sl ae

164A. Trachycephalus typhonius. Puerto Ayacucho, Amazonas.

Photo: César Barrio-Amoros.

(1986); Ramo and Busto (1989, 1990); Barrio (1996a);

Gorzula and Sefiaris (1998); Rojas-Runjaic et al. (2005);

Aguiar et al. (2007); Garda et al. (2010); Tarano (2010).

Genus Scarthyla Duellman and de Sa, 1988

Type species: Scarthyla ostinodactyla (= Hyla goinorum

Bokermann, 1962), by original designation.

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Barrio-Amoros et al.

Scarthyla vigilans (Solano 1971)

Type: MBUCV IV-6163

Type locality: “De la carretera entre Coloncito y El

Vigia (a los 20 km), Estado Zulia (sic!= Estado Tachira),

Venezuela.”

Distribution: Regions 1, 2, 3, 6. Colombia (Magdalena

Basin and Llanos), Venezuela and Trinidad (Smith et al.

2011). In Venezuela, apparent disjunct distribution in

lowlands of Maracaibo Lake Basin, high Llanos, Falcon

State (Barrio-Amoroés et al. 2006), northern Coastal

Range, and Delta of the Orinoco (Rojas-Runjaic et al.

2008). The type locality was situated in Zulia State by

Solano (1971), wrongly corrected to Mérida State by La

Marca (1992) and here corrected to Tachira State.

Remarks: Hy/a vigilans was regarded as incertae sedis by

Faivovich et al. (2005). Based on osteology and external

morphology, Barrio-Amoroés et al. (2006c) proposed

to assign H. vigilans to Scarthyla Duellman and de Sa,

1988 despite differences between the tadpoles of both

species. Lynch and Suarez-Mayorga (2011) assigned

vigilans to Pseudis, as P. vigilans, but that proposal was

never considered (Frost 2019). Despite the controversial

supraspecific position of this species, molecular evidence

indicates a close relationship between S. goinorum and S.

vigilans (B. Noonan and CBA, unpub. ).

Selected references: Solano (1971); Duellman and de

Sa (1988); La Marca (1992); Mijares and Hero (1997);

Barrio-Amoros (1998, 2004); Muares et al. (1998);

Suarez-Mayorga and Lynch (2001); Barrio-Amoros et

al. (2006c); Rojas-Runjaic et al. (2007, 2008); Lynch

and Suarez-Mayorga (2011); Smith et al. (2011); Frost

(2016).

Subfamily Scinaxinae

Duellman, Marion and Hedges, 2016

Remarks: Duellman et al. (2016) erected this

new subfamily for two of four genera (Scinax and

Sphaenorhynchus) present in Venezuela.

Genus Scinax Wagler, 1830

Type species: Hyla aurata Wied-Neuwied, 1821, by

subsequent designation of Stejneger (1907).

Scinax baumgardneri (Rivero 1961)*

Holotype: MCZ 28563.

Type locality: “Casa de Julian, entre Tapara y Cafio

Chana, 609 m, Estado Amazonas, Venezuela”.

Distribution: Region 4. Known from type locality and

two other localities (La Culebra and Puerto Ayacucho) in

Venezuelan Amazonia.

Remarks: In the Scinax ruber clade, but not assigned to

any species group (Faivovich et al. 2005). Scinax lindsayi

Pyburn, 1992 from Vaupes River between Brazil and

Amphib. Reptile Conserv.

Colombia seems similar to S. baumgardneri, and could be

a synonym. Fresh material of both and direct comparison

with types could corroborate this supposition.

Selected references: Rivero (1961, 1964b); Fouquette

and Delahoussaye (1977); Duellman and Wiens (1992);

Duellman (1997); Barrio-Amoros et al. (2004).

Scinax boesemani (Goin 1966)

Holotype: RMNH 12601.

Type locality: Zanderij, Suriname District, Suriname.

Distribution: Region 5. Venezuela, Guyana, Suriname,

French Guiana, and northern Brazil. In Venezuela,

several localities in Amazonas and Bolivar States.

Remarks: In Scinax ruber clade, but not assigned to any

Species group (Faivovich et al. 2005).

Selected references: Goin (1966); Rivero (1971b);

Duellman (1986, 1997); Duellman and Wiens (1992);

Gorzula and Sefiaris (1998); Barrio-Amoros et al. (2004,

2011b); Sefiaris et al. (2014).

Scinax danae (Duellman 1986)*

Holotype: KU 167073.

Type locality: “km 127 on El Dorado-Santa Elena de

Uairen Road, Estado de Bolivar, Venezuela, 1,250 m

(O5°57°N, 61°27°W).”

Distribution: Region 5. Apparently endemic to type

locality and surroundings in the high Sierra de Lema,

Bolivar State, Eastern Venezuela. Possibly also present

in Guyana.

Remarks: In Scinax ruber clade, but not assigned to any

Species group (Faivovich et al. 2005).

Selected references: Duellman (1986, 1997); Duellman

and Wiens (1992); Pyburn (1992); Gorzula and Sefiaris

(1998); Barrio-Amoros et al. (2004); Sefiaris et al. (2014).

Scinax exiguus (Duellman 1986)*

Holotype: KU 167094.

Type locality: “km 144 on the El Dorado-Santa Elena

de Uairén Road in the Gran Sabana, Estado Bolivar,

Venezuela, 1,210 m (05°53’N, 61°23’W).”

Distribution: Region 5. Endemic to Gran Sabana in

Bolivar State. Possibly also present in Guyana and

northern Brazil.

Remarks: In Scinax ruber clade, but not assigned to any

Species group (Faivovich et al. 2005).

Selected references: Duellman (1986, 1997); Duellman

and Wiens (1992); Gorzula and Sefiaris (1998); Barrio-

Amoros et al. (2004); Sefiaris et al. (2014).

Scinax fuscomarginatus (Lutz 1925)

Lectotype: AL-MN 845 designated by Cardoso and

Pombal (2010).

Type locality: “Sao Paulo et Bello Horizonte,” Brazil;

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Catalogue of the amphibians of Venezuela

164B. aeons enone Male. agai some valk 165A. Pseudis paradoxa. El Palmar, Bolivar. Photo: César

secretions. El Palmar, Bolivar. Photo: César Barrio-Amoros. Barrio-Amoros.

166A. Scarthyla vigilans. Male calling. Chururu, Tachira. Photo:

Amoré OS. César Barrio-Amoros.

166B. sSclatiyt vigilans. a Gonchi Zulia, PRO Céstar 167. Scinax baumgardneri. Capihuara, Casiquiare, Amazonas.

Barrio-Amoros. Photo: César Barrio-Amoréos.

restricted to Belo Horizonte, Minas Gerais, Brazil. distribution, formally synonymizing S. trilineatus with S.

Distribution: Region 5. Wide distribution area, from fuscomarginatus.

southern Brazil, Argentina and Paraguay to Venezuela, § Selected references: Literature refers to S. trilineatus in

Guyana, and Suriname. In Venezuela, several localities | Venezuela. Hoogmoed and Gorzula (1979); Duellman

in Bolivar State, from lowlands north of Sierrade Lema (1986); Duellman and Wiens (1992); Gorzula and

to uplands in the Gran Sabana. Sefiaris (1998); Martins (1998); Barrio-Amoros et al.

Remarks: In Scinax ruber clade, but not assigned to any = (2004); Cardoso and Pombal (2010); Brusquetti et al.

species group (Faivovich et al. 2005). Scinax trilineatus (2014); Sefiaris et al. (2014).

Hoogmoed and Gorzula, 1979 was synonymized with S.

fuscomarginatus without explanation by Martins (1998). Scinax garbei (Miranda-Ribeiro 1926)

Barrio-Amoros (2004, 2009) doubted the synonymy.

More recently, Brusquetti et al. (2014) comprehensively | Holotype: MZUSP 277.

reviewed Scinax fuscomarginatus throughout all its Type locality: Eirunepé, rio Jurua, Amazonas, Brazil.

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Barrio-Amoros et al.

ap oS Ee Re ae 6

169. Scinax danae. Uppers SSG of La Escalera, Sei de Lema,

Bolivar. Photo: César Barrio-Amoros.

170A. Scinax exiguus: Male. Kavanaven! Gran Sabana, ‘Bolivar. 170B. cee exiguus: peri North of Santa “Elena 7 we

Photo: César Barrio-Amoros. Bolivar. Photo: César Barrio-Amoros.

171A. Scinax fuscomarginatus. Male. North of Santa Elena de 171B. Scinax fuscomarginatus. Amplexus. North of Santa Elena

Uairén, Bolivar. Photo: César Barrio-Amorés. de Uairén, Bolivar. Photo: César Barrio-Amoros.

Distribution: Region 4. Colombia, Ecuador, Peru, Gaitan, Departamento Meta, Colombia.

Bolivia, Brazil, and Venezuela. In Venezuela, one locality Distribution: Region 4. Llanos of Colombia and

(Cafio Iguapo, Upper Orinoco) in southern Amazonas Amazonas State in Venezuela.

State. Remarks: In Scinax rostratus species group (Faivovich

Remarks: In Scinax rostratus species group (Faivovich — etal. 2005). Only locality known in Venezuela is Puerto

et al. 2005). Ayacucho (Amazonas State), based on the specimens

Selected references: Rivero (1967); Duellman (1970, § UPR-M 132-137 collected by Juan Rivero and referred by

1972b). him to Hyla boulengeri (Rivero 1961) but retrospectively

reidentified by Pyburn (1973) as S. kennedyi (La Marca

Scinax kennedyi (Pyburn 1973) 1992). Despite many herpetologists working for years

around Puerto Ayacucho, it has never been reported again

Holotype: UTA A-3697. in Venezuela.

Type locality: Around 110 miles ESE from Puerto Selected references: Pyburn (1973); Duellman and

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Catalogue of the amphibians of Venezuela

a / i» fe =

172. Scinax garbei. Male. Puyo, Ecuador. Photo: César Barrio-

Amoros.

174A. Scinax manriquei. Male. Olinda, La Azulita, Mérida. Photo:

César Barrio-Amoros.

175. Scinax nebulosus. Male. Cuyuni river, Bolivar. Photo: César

Barrio-Amoros.

Wiens (1992).

Scinax manriquei

Barrio-Amoros, Orellana and Chacon, 2004

Holotype: CVULA IV-1094.

Type locality: “Mesa Quintero, Guaraque, Estado

Merida, Venezuela, 1,700 m, 08°07’N, 71°36’ W.”

Distribution: Region 1. Andes of Venezuela

(States Mérida and Tachira) and adjacent Colombia

(departamento Norte de Santander).

Remarks: Scinax flavidus La Marca, 2004 is a junior

synonym of S. manriquei (Barrio-Amoros et al. 2010d).

In Scinax ruber clade, but not assigned to any species

group (Faivovich et al. 2005).

Selected references: Nicto-Castro (1999); Barrio-

Amoros (2004); Barrio-Amoros et al. (2004, 2010d), La

Amphib. Reptile Conserv.

SEES ‘a

173. Scinax kennedyi. Puerto Carrefio, Vichada, Colombia. Photo:

Juan David Jiménez.

174B. Scinax cf. manriquei. Doradas river, Tachira. Photo: César

Barrio-Amoros.

176A. Scinax rostratus. Casa Maria, Bejuma, Carabobo. Photo:

César Barrio-Amoros.

Marca (2004, 2007).

Scinax nebulosus (Spix 1824)

Neotype: MNRJ 4055 (holotype of Hyla egleri Lutz,

1968).

Type locality: “In sylvis prope flumen Teffé,” Amazonas,

Brazil. Neotype from Pirarucu pond of the Museu Goeldi

in Belem do Para, Para, Brazil (Hoogmoed and Gruber

1983).

Distribution: Region 5. Bolivia, Brazil, Venezuela,

Guyana, Suriname, and French Guiana. In Venezuela,

several localities in Bolivar State (Imataca, Cuyun,

Triunfo).

Remarks: In Scinax rostratus species group (Faivovich

et al. 2005). Some reports by Rivero (1961) as Hyla

boulengeri (especially those south of the Orinoco) can be

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Barrio-Amoros et al.

nebulosus, garbei, or kennedyi.

Selected references: Hoogmoed and Gruber (1983);

Duellman and Wiens (1992); Duellman (1997); Barrio-

Amoroés et al. (2004, 2011b); Barrio-Amoros and

Duellman (2009); Sefiaris et al. (2009).

Scinax rostratus (Peters 1863)

Holotype: ZMB 3175.

Type locality: Caracas, Venezuela.

Distribution: Regions 2, 3, 4, 6. Widely distributed in

northern South America, from Panama and throughout

Colombia, Venezuela, and Guyana to northern Brazil.

Widespread in the lowlands of Venezuela.

Remarks: In Scinax rostratus species group (Faivovich

et al. 2005). Rivero (1961) reported several specimens as

Hyla boulengeri. A complex of species, involving at least

two different taxa only in Venezuela (CBA, unpub. data).

A major review is needed to clarify its taxonomic status

and the whole S. rostratus species group. In northwestern

Venezuela, specimens of S. rostratus should be compared

with Scinax boulengeri (Cope 1887).

Selected references: Ginés (1959); Rivero (1961,

1964a,b, 1967a, 1968g); Tello (1968); Duellman (1970,

1972b, 1997); Fouquette and Delahoussaye (1977);

Duellman and Wiens (1992); Hero and Miyares (1995):

Yustiz (1996); Sefiaris et al. (2014).

Scinax ruber (Laurenti 1768)

Neotype: RMNH 15922B.

Type locality: “America.” Neotype from Paramaribo,

Surinam.

Distribution: Regions 2, 5. Widely distributed through

the Guiana Shield Basin (northeastern Brazil, south and

eastern Venezuela, Guyana, Suriname, French Guiana),

Trinidad and Tobago, and St. Lucia in the Lesser Antilles.

Introduced in Puerto Rico. In Venezuela, eastern and SE

parts, from Peninsula de Paria (type locality of Hyla

robersimoni, a synonym) to the northern versant of Sierra

de Lema. Not known from the uplands of Gran Sabana.

Remarks: Distribution and specimens of this species

historically confused in Venezuela with those of Scinax

x-signatus. Fouquet et al. (2007) showed at least six

cryptic species could be hidden under that name. All

previously mentioned S. ruber from Amazonian Brazil,

Colombia, Ecuador, Peru, and Bolivia, as well as Trans-

Andean, are representatives of other, undescribed taxa.

Since neotype locality 1s Surinam, Scinax ruber sensu

stricto occurs from eastern Venezuela and Trinidad

to northeastern Brazil through the Guianas, being a

Guianan endemic (Hoogmoed 1979b). In Scinax ruber

clade (Faivovich et al. 2005). Barrio-Amoros and Ortiz

(2015) comment on type material of the synonym Hyla

robersimoni Donoso-Barros, 1966.

Selected references: Daudin (1803); Gunther (1858);

Ginés (1959); Rivero (1961, 1964a,b, 1967c, 1968g,

Amphib. Reptile Conserv.

1969a); Heatwole et al. (1965); Donoso-Barros (1966);

Duellman (1972b, 1979a, 1986, 1997): Fouquette and

Delahoussaye (1977); Hoogmoed and Gorzula (1979);

Hoogmoed and Gruber (1983); Gorzula (1985a);

Duellman and Wiens (1992); Barrio-Amoros et al. (2004,

2011b); Fouquet et al. (2007); Sefiaris et al. (2014);

Barrio-Amoros and Ortiz (2015).

Scinax wandae (Pyburn and Fouquette 1971)

Holotype: USNM 192305.

Type locality: “12 km NNE of Villavicencio, Meta,

Colombia, elevation about 580 m.”

Distribution: Regions 1, 3. Llanos of Colombia, and

Amazonian lowlands, eastern slopes of Andes and Upper

Llanos of Venezuela (Amazonas, Bolivar, Apure, and

Barinas States).

Remarks: In Scinax ruber clade, but not assigned to any

Species group (Faivovich et al. 2005).

Selected references: Pyburn and Fouquette (1971);

Lynch and Vargas-Ramirez (2000); Barrio and Fuentes

(2003); Barrio and Chacon-Ortiz (2004); Barrio-Amoros

et al. (2004).

Scinax x-signatus (Spix 1824)

Holotype: ZSM 2494/0 (lost).

Type locality: Bahia, Brazil.

Distribution: Regions 2, 3, 4, 5, 6. Eastern, southeastern,

and southern Brazil, Colombia, Venezuela, Guyana, and

Suriname. There are at least five species in Venezuela

under this name [not three as Barrio-Amoros (2004)

referred]. All inhabit non-heavily forested habitats,

i.e., temporarily flooded savannas and dry forests to

rainforests. Each of the five found in distinct locations:

Maracaibo Lake Basin; Coastal Range; the Llanos region;

Gran Sabana Region; and Isla de Margarita (Ugueto and

Rivas Fuenmayor 2010).

Remarks: The type of Scinax x-signatus is lost

(Hoogmoed and Gruber 1983), so it is not possible to

review the x-signatus group until a neotype is designated.

Selected references: Spix (1824); Rivero (1969a);

Leon-Ochoa (1975); Fouquette and Delahoussaye

(1977); Hoogmoed and Gorzula (1979); Rada (1981);

Hoogmoed and Gruber (1983); Duellman (1986, 1997);

Ramo and Busto (1989, 1990); Duellman and Wiens

(1992); Gorzula and Sefiaris (1998); Barrio-Amoros

et al. (2004, 2011b); Tarano (2010); Ugueto and Rivas

Fuenmayor (2010); Sefiaris et al. (2014).

Genus Sphaenorhynchus Tschudi, 1838

Type species: Hyla lactea Daudin, 1802, by original

designation.

Sphaenorhynchus lacteus (Daudin 1800)

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Catalogue of the amphibians of Venezuela

176B. Scinax rostratus. Cafio Caiman, Apure-Barinas frontier.

Photo: César Barrio-Amoros.

177B. Scinax ruber. Night color. Imataca Forestal Reserve, Delta 178. Scinax wandae. Arassari Rafting Camp, Acequias, Barinas.

Amacuro. Photo: César Barrio-Amoros. Photo: César Barrio-Amoros.

ol ma BS Moa §

179A. Scinax x-signatus. Parque Los Caobos, Caracas. Photo: 179B. Scinax x-signatus. Pagiiey river, Barinas. Photo

César Barrio-Amoros. Barrio-Amoros.

179C. Scinax x-signatus. Arassari Rafting Camp, Acequias river, 179D. Scinax x-signatus. Santa Elena de Uairén, Bolivar. Photo:

Barinas. Photo: César Barrio-Amoros. César Barrio-Amoros.

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Barrio-Amoros et al.

Holotype: MNHNP 4871.

Type locality: Brazil.

Distribution: Region 5. Orinoco and Amazon Basins

(Colombia, Ecuador, Bolivia, Brazil, Venezuela,

Guyana, Suriname, French Guiana), and on the island of

Trinidad (Trinidad and Tobago). In Venezuela, only a few

localities in the south and east (Amazonas, Bolivar, Delta

Amacuro, Monagas, and Sucre states).

Selected references: Rivero (1961, 1969b); Hoogmoed

and Gorzula (1979); Duellman and Lynch (1981);

Duellman (1997); Barrio and Rivero (1999b); Barrio-

Amoros et al. (2011b).

Family Leptodactylidae Werner, 1896 (1838)

Remarks: This family has undergone a massive

change of its member taxa since the widespread use

of molecular data. Prior to 2004, the nomenclature

was quite stable, but Darst and Cannatella (2004)

demonstrated Leptodactylidae to be paraphyletic. After

several revisions of the major clades of Amphibia (Frost

et al. 2006; Grant et al. 2006; Pyron and Wiens 2011;

De Sa et al. 2014), it is now mostly restricted to the old

Leptodactylinae plus Leiuperinae and Paratelmatobiinae.

Subfamily Leptodactylinae Werner, 1896 (1838)

Genus Adenomera Fitzinger, 1867

Type species: Adenomera marmorata Steindachner,

1867, by monotypy.

Remarks: Frost et al. (2006) synonymized Adenomera

with Leptodactylus, which was not widely followed.

Pyron and Wiens (2011) recognized Adenomera and

Lithodytes as separate genera.

Adenomera andreae (Miller 1923)

Holotype: ZSM 136/1911.

Type locality: “Peixeboi (a.d. Bragancabahn), Staat

Para, Brasilien.”

Distribution: Regions 4, 5. Widely distributed in

lowlands east of the Andes, including Colombia,

Venezuela, Ecuador, Peru, Bolivia, Brazil, Guyana,

Suriname, and French Guiana. In Venezuela, common in

lowlands south of Orinoco River.

Remarks: The reported Adenomera andreae from the

eastern slopes of Venezuelan Andes (Barrio-Amoros

2004) are actually A. simonstuarti (see Appendix 1).

Selected references: Heyer (1973, 1977); Rivero et

al. (1986); Barrio-Amorés and Brewer-Carias (2008);

Barrio-Amoros (2010); Barrio-Amoros et al. (2011b);

Fouquet et al. (2014).

Adenomera hylaedactyla (Cope 1868)

Holotype: ANSP 2240.

Amphib. Reptile Conserv.

Type locality: “From the Napo or upper Maranon River,”

Peru.

Distribution: Regions 4, 5. Widely distributed in

lowlands east of the Andes, including Colombia,

Venezuela, Ecuador, Peru, Bolivia, Brazil, Guyana,

Suriname, and French Guiana. In Venezuela, common in

lowlands south of Orinoco River.

Selected references: Ginés (1959); Rivero (1961, 1963c,

1964a-c, 1967a); Heyer (1973, 1977); Rivero et al.

(1986); Duellman (1997); Gorzula and Sefiaris (1998),

Lynch and Vargas-Ramirez (2000); Barrio-Amoros et al.

(2011b); Fouquet et al. (2014); Sefiaris et al. (2014).

Genus Leptodactylus Fitzinger, 1826

Type species: Rana typhonia Latreille, in Sonnini and

Latreille, 1801 (= Rana fusca Schneider, 1799), by

subsequent designation by Fitzinger (1843).

Remarks: Following De Sa et al. (2014), the old phenetic

groups of Leptodactylus established by Heyer are no

longer stable. All species now under L. fuscus, L. latrans,

L. melanonotus, and L. pentadactylus species groups.

Leptodactylus bolivianus

Boulenger, 1898

Lectotype: MSNG 28875A.

Type locality: Barraca, Rio Madidi, Bolivia (by

designation of the Lectotype).

Distribution: Region 4. Widely distributed in Amazon

Region, including Venezuela, Colombia, Peru, Bolivia,

and NW Brazil. In Venezuela, lowlands south of Orinoco

River.

Remarks: Heyer and De Sa (2011) reviewed the L.

bolivianus complex, adscribing the northern Venezuelan

specimens to L. insularum (as did Rivero 1967a, c), and

Barrio-Amoros (1998, 2004, 2009). In the Leptodactylus

latrans species group of De Sa et al. (2014).

Selected references: Many of these apply to L.

insularum sensu Heyer and De Sa, 2011. Aleman (1952);

Ginés (1959): Rivero (1961, 1964a—d, 1967a); Heatwole

et al. (1965); Tello (1968); Staton and Dixon (1977);

Hoogmoed and Gorzula (1979); Rivero et al. (1986);

Yustiz (1996); Duellman (1997); Heyer and De Sa

(2011); De Sa et al. (2014).

Leptodactylus colombiensis Heyer, 1994

Holotype: ICN 7409.

Type locality: “Charala, Virolin (= Inspeccion Policia

Cafiaverales), confluencia del rio Cafiaverales con el rio

Guillermo, vertiente occidental, 1,600—1,700 m, 6°13’N,

73°05’ W,” Santander Department, Colombia.

Distribution: Region 1. Colombia and Venezuela. In

Venezuela, eastern Andean slopes of Tachira State.

Remarks: De Sa et al. (2014) doubt Venezuelan records

(Barrio and Chacon 2001) without explanation. Specimens

July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

179E. Scinax x-signatus. Santa Elena de Uairén, Bolivar. Photo:

César Barrio-Amoros.

180. Sphaenorhynchus lacteus. Male. El Palmar, Bolivar. Photo:

César Barrio-Amoros.

, : m -"

» _

; ‘ ta\ ,

182. Physalaemus cuvieri. Imataca Forestal Reserve, Delta

Amacuro. Photo: César Barrio-Amoros.

CVULA 5638-42 conform to diagnostic characters in

Heyer (1994a). In Leptodactylus melanonotus species

group of De Sa et al. (2014).

Selected references: Heyer (1994a):; Barrio and Chacon

(2001); De Sa et al. (2014).

Leptodactylus diedrus Heyer, 1994

Holotype: UTA A-3726.

Type locality: “Colombia, Vaupés, 1/2 mi NE Timbo,

1°06’N, 70°01? W.”

Distribution: Regions 4, 5. Western Amazonia

Amphib. Reptile Conserv.

179F. Scinax x-signatus. Santa Elena de Uairén, Bolivar. Photo:

César Barrio-Amoros.

fi 5 *

i’ ; * _

Engystomops

eae

183. Physalaemus ephippifer. Santa Elena de Uairén, Bolivar.

Photo: César Barrio-Amoros.

(Colombia, Venezuela, Peru, and Brazil). In Venezuela,

only from extreme south of Amazonas (Neblina) and

Bolivar (Sarisarifiama) States.

Remarks: Vanzolinius discodactylus (Boulenger 1884)

reported by McDiarmid and Paolillo (1988) from base

of Neblina, but specimen later assigned to Leptodactylus

diedrus (Heyer 1997). In Leptodactylus melanonotus

Species group of De Sa et al. (2014).

Selected references: McDiarmid and Paolillo (1988);

Heyer (1994a, 1998); Barrio-Amoros and Brewer-Carias

(2008); Barrio-Amoros et al. (2011b); De Sa et al. (2014).

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

Leptodactylus fragilis (Brocchi 1877)

Type: MNHNP 6316.

Type locality: “Tehuantepec,” Oaxaca, Mexico.

Distribution: Regions 3, 4, 6. Widely distributed:

southern USA (Texas), Central America to northern

Colombia and Venezuela. In Venezuela, lowlands north

of Orinoco River.

Remarks: Probably a complex of species (CBA, unpub.

data). Nomenclatorical history in Heyer and Dubois

(1992) and Heyer (2002). In Leptodactylus fuscus species

group of De Sa et al. (2014).

Selected References: Staton and Dixon (1977); Heyer

(1978, 2002); Ramo and Busto (1989, 1990); Dubois and

Heyer (1992); Tarano (2010); De Sa et al. (2014).

Leptodactylus fuscus (Schneider 1799)

Neotype: MNHNP 680.

Type locality: Not designated. Neotype from “Surinam.”

Distribution: Regions 2, 3, 4, 5, 6. Widespread in

lowlands from Panama through South America to east

of Andes (Colombia, Venezuela, Trinidad and Tobago,

Guyana, Suriname, French Guiana, Brazil, Bolivia,

Paraguay, and northern extreme of Argentina). In

Venezuela, widely distributed in lowlands throughout

country, including Margarita Island.

Remarks: A complex of species. In Leptodactylus fuscus

species group of De Sa et al. (2014).

Selected references: Ginés (1959); Rivero (1964a-d,

1967a); Heatwole et al. (1965); Tello (1968); Bogart

(1974); Staton and Dixon (1977); Hoogmoed and Gorzula

(1979); Péfaur and Diaz De Pascual (1987); Solano

(1987a,b); Ramo and Busto (1989, 1990); Magdefrau et

al. (1991); Manzanilla et al. (1995); Duellman (1997);

Gorzula and Sefiaris (1998); Barrio-Amoroés (2010a);

Tarano (2010); Ugueto and Rivas-Fuenmayor (2010);

Barrio-Amoros et al. (2011b); De Sa et al. (2014); Sefiaris

et al. (2014).

Leptodactylus guianensis Heyer and de Sa, 2011

Holotype: USNM 531509.

Type locality: “Guyana; Rupunini, Iwokrama Forest

Reserve, Sipuruni River, Pakatau Camp, 4°45717”N,

59°01°28” W, 85 m.”

Distribution: Region 5. Lowlands and uplands of Guiana

Shield in Venezuela, Guyana, Suriname, and Brazil. In

Venezuela, only from eastern Bolivar State.

Remarks: Records from eastern part of Venezuelan

Guayana previously assigned to Leptodactylus bolivianus

are actually L. guianensis (e.g., Barrio-Amoros et al.

(2011b) from Triunfo). In Leptodactylus latrans species

group of De Sa et al. (2014).

Selected references: Rivero et al. (1986); Barrio-Amoros

et al. (2011b); De Sa et al. (2014); Sefiaris et al. (2014).

Amphib. Reptile Conserv.

Leptodactylus insularum Barbour, 1906

Lectotype: MCZ 2424.

Type locality: “Saboga island,’ Bahia de Panama,

Panama.

Distribution: Regions 2, 3, 4, 6. From Costa Rica in

Central America through Panama and Colombia (incl.

San Andrés Island) to Venezuela and Trinidad and

Tobago. In Venezuela, common in lowlands north of

Orinoco River.

Remarks: Considered a synonym of Leptodactylus

bolivianus by Heyer (1968) and Savage (2002); Rivero

(1967a,c) found important differences, e.g., one spine on

thumb in L. bolivianus, vs. two in L. insularum. Batrio-

Amoros (1998, 2004) differentiated between insularum

north of Orinoco and bolivianus to the south. Heyer and

De Sa (2011) resolved the question. In Leptodactylus

latrans species group of De Sa et al. (2014).

Selected references: Sexton (1962); Rivero (1967a,c);

Heyer (1968); Bogart (1974); Rivero et al. (1986);

Manzanilla et al. (1995); Savage (2002); Tarano (2010);

Barrio-Amoros et al. (2011b); Heyer and De Sa (2011);

De Sa et al. (2014).

Leptodactylus knudseni Heyer, 1972

Holotype: LACM 72117.

Type locality: “Limoncocha,

Provincia de Napo, Ecuador.”

Distribution: Regions 4, 5. Widely distributed in

Amazon Basin, incl. Bolivia, Peru, Ecuador, Colombia,

Brazil, Venezuela Guyana, Suriname, French Guiana,

and Trinidad and Tobago. In Venezuela, widespread

south of Orinoco River.

Remarks: In Leptodactylus pentadactylus species group

of De Sa et al. (2014).

Selected references: Heyer (1972, 1979, 2005); Hero

and Galatti (1990); Barrio (1996a); Gorzula and Sefiaris

(1998); Barrio-Amoréos et al. (2011b); De Sa et al. (2014);

Sefiaris et al. (2014).

0°24°S, 76°37°W,

Leptodactylus leptodactyloides (Andersson 1945)

Type: NHRM 1945.

Type locality: “Rio Pastaza,” eastern Ecuador.

Distribution: Region 5. Amazon Basin (Colombia,

Ecuador, Peru, Bolivia, Brazil, Venezuela, Guyana,

Suriname, and French Guiana). Only single locality in

Venezuela: Rio Cuyuni, km 69 on road to Santa Elena

de Uairén, Bolivar State. Probably widespread in

southeastern lowlands.

Remarks: Sister taxon of Leptodactylus petersii. In

Leptodactylus melanonotus species group of De Sa et al.

(2014).

Selected references: Heyer (1994); Duellman (1997);

De Sa et al. (2014).

July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

185. Pleurodema brachyops. San Vicente, Apure. Photo: César

Barrio-Amoros.

186. Pseudopaludicola boliviana. Lower Caura river basin, 187A. Pseudopaludicola llanera. Puerto Ayacucho, Amazonas.

Bolivar. Photo: Fernando Rojas-Runjaic. Photo: Zelimir Cernelic.

187B. Pseudopaludicola llanera. Puerto Ayacucho, Amazonas. 188. Pseudopaludicola pusilla. Barrancabermeja, Santander,

Photo: Fernando Rojas-Runjaic. Colombia. Photo: Juan S. Mendoza.

189. Adenomera andreae. Imataca Forestal Reserve, Delta 190A. Adenomera hylaedactyla. Lower Serrania del Cuao,

Amacuro. Photo: César Barrio-Amoros. Amazonas. Photo: Fernando Rojas-Runjaic.

Amphib. Reptile Conserv. July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

7 f hk rial Z = : ae = : : ; : -_ |

190B. Adenomera cf. hylaedactyla. Palmichal, Bejuma, Carabobo.

Photo: César Barrio-Amoros.

, ve % ret .

r eo . r

pe 2

: 1

al] at r

192. Leptodactylus colombiensis. Doradas river valley, Tachira.

Photo: César Barrio-Amoros.

a a

194A. Leptodactylus fuscus. San Vicente, Apure. Photo: César

195. Leptodactylus guianensis. Female. Triunfo, Eastern sector of

Sierra de Lema, Bolivar. Photo: César Barrio-Amoros.

Amphib. Reptile Conserv.

191. Adenomera cf. simonstuarti. San Ramon, Calderas, Barinas.

Photo: César Barrio-Amoros.

193. Leptodactylus fragilis. Pagiey river, Barinas. Photo: César

Barrio-Amoros.

194B. Leptodactylus fuscus. Imataca, Delta Amacuro. Photo:

César Barrio-Amoros.

ke en i

5 igs cit 4 m

196A. Leptodactylus ins.

ularum. Male. Pagtiey river, Barinas.

Photo: César Barrio-Amoros.

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Catalogue of the amphibians of Venezuela

: a - axkie eae ainsi i “2 : “4 . a hol

196B. Leptodactylus ear Female. Pagiiey 1 river, Barinas. 196C. Leptodactylus SE El Vigia, Mérida. Photo: César

Photo: César Barrio-Amoros. Barrio-Amoros.

DM os ies

At 5

197, TEIOORGRTES Wau: Tag Claritas, Bolivar Photo: César 198, eaniedaepis lithonaetes. Raudal del Dantol Aatanal

Barrio-Amoros. Amazonas. Photo: César Barrio-Amoros.

199. Leptodactylus longirostris. Salto Angel, Auyan-tepui, 200. Leptodactylus aff. macrosternum. Pagtiey river, Barinas.

Bolivar. Photo: Fernando Rojas-Runjaic. Photo: César Barrio-Amoros.

Leptodactylus lithonaetes Heyer, 1995 (1995); Lynch and Vargas-Ramirez (2000); Heyer and

Heyer (2001); Heyer and Barrio-Amoros (2009); De Sa

Holotype: AMNH. 100656. et al. (2014).

Type locality: “Venezuela: Amazonas, SW sector Cerro Leptodactylus longirostris Boulenger, 1882

Yapacana, 600 m, 3° 57’N, 67° 00’W.”

Distribution: Regions 4, 5. Western Guiana Shield Lectotype: BM 76.5.26.4.

(Colombia and Venezuela). In Venezuela, restricted to Type locality: “Santarem,” Para, Brazil.

rocky outcrops of extreme southeastern Apure State, Distribution: Region 5. Widespread in Guiana Shield of

Amazonas, and northwestern Bolivar State. Colombia, Venezuela, Guyana, Suriname, and northern

Remarks: Heyer and Barrio-Amorods (2009) describe — Brazil. In Venezuela, restricted to south of the Orinoco

the call. Sister species of Leptodactylus rugosus. In _ River.

Leptodactylus pentadactylus species group of De Sa et Remarks: In Leptodactylus fuscus species group of De

al. (2014). Sa et al. (2014).

Selected references: Donnelly and Myers (1991); Heyer Selected references: Rivero (1971a); Heyer (1978);

Amphib. Reptile Conserv. 94 July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

Crombie and Heyer (1983); Donnelly and Myers (1991),

Magdefrau et al. (1991); Duellman (1997); Gorzula and

Sefiaris (1998); Heyer and Heyer (2001); Barrio-Amoros

and Duellman (2009); Sefiaris et al. (2009, 2014); Barrio-

Amoros et al. (2011b); De Sa et al. (2014).

Leptodactytlus magistris

Mijares-Urrutia, 1997*

Holotype: EBRG 3284.

Terra typica: “Cerro Socop6, cerca de 30 km (por

carretera) al SO de Guajiro, Municipio Mauroa, Estado

Falcon, Venezuela, cerca de 1,250 m.”

Distribution: Region 2. Known only from type locality.

Remarks: Despite recent efforts to find it, Leptodactylus

magistris seems to have suffered a severe decline. No new

information since its description. Considered Critically

Endangered by the Venezuelan Red Book (La Marca et

al. 2015). In Leptodactylus melanonotus species group of

De Sa et al. (2014).

Selected references: Mijares-Urrutia (1997); De Sa et

al. (2014); La Marca et al. (2015).

Leptodactylus mystaceus (Spix 1824)

Lectotipe: ZSM 2504/0.

Type locality: Restricted to “Solimées,” Brazil, by

lectotype designation.

Distribution: Regions 4, 5. Widely distributed in

Amazon Basin and northern portion of the Atlantic Forest

in Brazil, Colombia, Venezuela, Guyana, Suriname,

and French Guiana. In Venezuela, widespread south

of Orinoco River. Localities north of Orinoco must be

reviewed, probably a new species.

Remarks: [n Leptodactylus fuscus species group of De

Sa et al. (2014).

Selected references: Ginés (1959); Rivero (1961,

1964b, 1968e); Roze (1964); Heatwole et al. (1965);

Bogart (1974); Heyer (1978, 1983); Rivero et al. (1986);

La Marca (1992); Duellman (1997); Gorzula and Sefiaris

(1998); De Sa et al. (2014); Sefiaris et al. (2014).

Leptodactylus pentadactylus (Laurent 1768)

Neotype: RMNH 29559.

Type locality: “Suriname, Marowijne, Lelygebergte,

Suralcokamp.”

Distribution: Region 4, 5. Widespread in Amazon

Region of Venezuela, Colombia, Ecuador, Peru, Bolivia,

Brazil, Guyana, Suriname, and French Guiana. In

Venezuela, in lowland forest south of Orinoco River.

Remarks: Vouchers absent from Venezuelan museums;

all pentadactylus-like frogs from southern Venezuela

identified as Leptodactylus knudseni. Heyer’s (2005)

review of the pentadactylus species group does not

mention any voucher from Venezuela. Removed

from previous lists by Barrio-Amords (2009) without

Amphib. Reptile Conserv.

explanation. De Sa et al. (2014) show four Venezuelan

localities in a distribution map, but without vouchers.

In Leptodactylus pentadactylus species group of De Sa

et al. (2014). May be distinguished from L. knudseni

by absence of chest spines in reproductive males of L.

pentadactylus (present in L. knudseni).

Selected references: Rivero (1964b,d, 1969a); Heatwole

et al. (1965); Bogart (1974); Muedeking and Heyer

(1976); Heyer (1979, 2005); Hero and Galatti (1990);

Duellman (1997); Gorzula and Sefiaris (1998); De Sa et

al. (2014).

Leptodactylus petersii (Steindachner 1864)

Holotype: Lost; formerly in the NMW.

Type locality: “Marabitanas,’ Amazonas, Brazil.

Distribution: Regions 4, 5. Widely distributed in

Amazon Region (Venezuela, Colombia, Peru, Bolivia,

Brazil, Guyana, Suriname, and French Guiana) south

to the Cerrado in central Brazil. Two recent confirmed

records for Venezuelain southern part of Amazonas State

and eastern border of Bolivar State.

Remarks: In Leptodactylus melanonotus species

group sensu De Sa et al. (2014). Sister taxon of L.

leptodactyloides. Records of specimens in Venezuelan

museum indicates widespread in lowlands of almost all

the country, but many specimens could be other species of

Leptodactylus melanonotus group; species identification

in this group is very challenging. The map in De Sa et al.

(2014) is misplaced with that of Leptodactylus pascoensis

and only shows two localities in southern Venezuela.

Selected references: Ginés (1959); Rivero (1963c,

1964a-d, 1971b); Heatwole et al. (1965); Heyer (1994);

Duellman (1997); Tarano (2010); De Sa et al. (2014);

Sefiaris et al. (2014).

Leptodactylus poecilochilus (Cope 1862)

Syntype: USNM 4347.

Type locality: “Near Turbo,” Antioquia, Colombia.

Distribution: Regions 2, 3, 6. From Pacific lowlands of

northwestern Costa Rica through Panama and northern

Colombia to northwestern Venezuela.

Remarks: [n Leptodactylus fuscus species group of De

Sa et al. (2014).

Selected references: Rivero (1961, 1963a,c, 1964a,

1971a); Rivero and Esteves (1969); Heyer (1978); La

Marca (1992); Yustiz (1996); De Sa et al. (2014).

Leptodactylus rhodomystax Boulenger, 1884

Lectotype: BMNH 1947.12.17.81.

Type locality: “Yurimaguas, Huallaga River, (Loreto)

Northern Peru.”

Distribution: Region 5. Amazonian Region of Colombia,

Ecuador, Peru, Bolivia, Guyana, Suriname, French

Guiana, and Brazil. In Venezuela, only La Escalera,

July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

i ae ih) a "

Ca ‘d

201. Leptodactylus mystaceus. Lower Caura river basin, Bolivar. 202. Leptodactylus pentadactylus. Yasuni, Ecuador. Photo: César

Photo: Fernando Rojas-Runjaic. Barrio-Amoros.

203A. Leptodactylus petersii. Male. Parque Los Caobos, Caracas. 203B. Leptodactylus petersii. Female. Parque Los Caobos,

Photo: César Barrio-Amoros. Caracas. Photo: César Barrio-Amoros.

py i a a

204. Leptodactylus poecilochilus. Riecito Maché, Sierra de Perija, 205A. Leptodactylus rhodomystax. Juvenile. Las Lajas, Canaima

Zulia. Photo: Fernando Rojas-Runjaic. National Park, Bolivar. Photo: Edward Camargo.

205B. Leptodactylus rhodomystax. Juvenile. Ecuador. Photo: Luis 206A. Leptodactylus riveroi. Capihuara, Casiquiare, Amazonas.

Coloma. Photo: César Barrio-Amoros.

Amphib. Reptile Conserv. 96 July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

Bolivar State (Camargo et al. 2014).

Remarks: First mention in Venezuela is by Rivero

(1961), who was not sure if it was a new species (Rivero

1968e), but decided to use this name. Later Heyer and

Pyburn (1983) described the taxon referred by Rivero

(1968e) as a new species (Leptodactylus riveroi).

Gorzula and Sefiaris (1998) confused a large specimen

of L. guianensis (as L. bolivianus) with L. rhodomystax.

De Sa et al. (2014) indicate a locality in Venezuela, but

no voucher or locality data are mentioned. Camargo et

al. (2014) provide the first voucher for the country. In

Leptodactylus pentadactylus species group of De Sa et

al. (2014).

Selected references: Rivero (1968e); Heyer (1979,

2005); Heyer and Pyburn (1983); Gorzula and Sefiaris

(1998); Camargo et al. (2014); De Sa et al. (2014).

Leptodactylus riveroi Heyer and Pyburn, 1983

Holotype: USNM 232400.

Type locality: “Colombia, Vaupés, Timbo, 01°06’S,

70°01’ W, elevation 170 m.”

Distribution: Region 4. Colombia, Venezuela, and

northern Brazil. In Venezuela, riverine lowlands of

Amazonas State.

Remarks: Leptodactylus rhodomystax was erroneously

reported from Venezuela (Rivero 1961, 1968e) based on

specimens of L. riveroi. In Leptodactylus melanonotus

species group of De Sa et al. (2014).

Selected references: Rivero (1968e); Heyer and Pyburn

(1983); Lima (1992); Gorzula and Sefiaris (1998); Lynch

and Vargas-Ramirez (2000); Barrio-Amoros and Brewer-

Carias (2008); De Sa et al. (2014).

Leptodactylus rugosus Noble, 1923

Holotype: AMNH 1169.

Type locality: “near Kaieteur Falls, British Guiana.”

Distribution: Region 5. Venezuela and Guyana. In

Venezuela, in lowlands and uplands of eastern Guiana

shield (Bolivar State) including summits of some tepuis.

Remarks: In Leptodactylus pentadactylus species group

of De Sa et al. (2014).

Selected references: Ginés (1959); Rivero (1961,

1964b,d, 1968d); Heatwole et al. (1965); Heyer (1979,

1995); Hoogmoed and Gorzula (1979); Donnelly and

Myers (1991); Magdefrau et al. (1991); Duellman (1997);

Gorzula and Sefiaris (1998); Heyer and Thompson

(2000); Barrio-Amoros and Duellman (2009); Heyer and

Barrio-Amoros (2009); De Sa et al. (2014); Sefiaris et al.

(2014).

Leptodactylus sabanensis Heyer, 1994*

Holotype: KU 166559.

Type locality: “Venezuela; Bolivar; km 127, El Dorado-

Santa Elena de Uairen road, 1,250 m, 6°00’N, 61°30’ W.”

Amphib. Reptile Conserv.

Distribution: Region 5. Gran Sabana in eastern

Venezuela and adjacent Lavrado in Roraima, Brazil.

Remarks: In the Leptodactylus melanonotus species

group of De Sa et al. (2014).

Selected references: Heyer (1994); Duellman (1997);

Gorzula and Sefiaris (1998); De Sa et al. (2014); Sefiaris

et al. (2014).

Leptodactylus turimiquensis Heyer, 2005

Holotype: AMNH 70667

Type locality: “Caripito, Monagas, Venezuela, ~ 100 m,

10°08’N, 63°06’ W.”

Distribution: Region 2. Endemic from northeastern

Venezuela, from Macizo de Turimiquire (Anzoategui,

Monagas and Sucre States) to Peninsula de Paria.

Remarks: Heyer (2005) described this species based

on a population previously assigned to Leptodactylus

labyrinthicus. In pentadactylus species group of De Sa

et al. (2014).

Selected references: Spix (1824); Gunther (1858);

Heyer (1979); Péfaur and Sierra (1995); Gorzula and

Sefiaris (1998).

Leptodactylus validus Garman, 1888

Lectotype: MCZ A-71920.

Type locality: “Kingston, St. Vincent,” Lesser Antilles.

Distribution: Regions, 2, 3, 4, 5, 6. Lesser Antilles,

Trinidad and Tobago, Venezuela, Guyana, Suriname, and

French Guiana. Widely distributed in lowland Venezuela,

north and south of Orinoco River.

Remarks: Includes all previous references for

Leptodactylus podicipinus, L. wagneri, and L.

pallidirostris from Venezuela. Synonymy by Yanek et

al. (2006). In me/anonotus species group of De Sa et al.

(2014).

Selected references: Ginés (1959); Rivero (1961, 1963c,

1964a-d, 1968e); Heatwole et al. (1965); Rivero and

Esteves (1969); Bogart (1974); Hoogmoed and Gorzula

(1979); Rivero et al. (1986); Donnelly and Myers (1991),

Heyer (1994); Gorzula and Sefiaris (1998); Yanek et al.

(2006); Camargo et al. (2009); De Sa et al. (2014).

Genus Lithodytes Fitzinger, 1843

Type species: Hylodes lineatus Dumeéril and Bibron,

1841 (= Rana lineata Schneider, 1799), by original

designation.

Remarks: Frost et al. (2006) synonymized Lithodytes

within Leptodactylus, but not widely followed. Pyron

and Wiens (2011) and de Sa et al. (2014) recognize

Adenomera and Lithodytes as valid genera.

Lithodytes lineatus (Schneider 1799)

Holotype(s): “Musei Lampiani” (= the “collection de

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Catalogue of the amphibians of Venezuela

206B. Leptodactylus riveroi. Nifial, Casiquiare, Amazonas. Photo:

Fernando Rojas-Runjaic.

208. ep ALCA is anne Chivaton, Gran Span Piet

César Barrio-Amoros.

207. emedacnis rugosus. EI Playén, an river, Bolivar

Photo: César Barrio-Amoros.

209. Leptodactylus turimiquensis. Las Melenas, Peninsula de

Paria, Sucre. Photo: César Barrio-Amoros.

210. Leptodactylus validus: Apure river, Api Beis! César

Barrio-Amoros.

Lampi” according to Daudin 1802: 105, and Daudin

1803, 8: 188).

Type locality: Not stated.

Distribution: Regions 1, 4, 5, 6. East of Andes from

northwestern Venezuela through Guyana, Suriname,

French Guiana, Brazil, Colombia, Ecuador, and Peru

to northern Bolivia. In Venezuela, disjunct populations

south of Orinoco River, eastern slopes of Andes and

Maracaibo Basin (west of Andes).

Remarks: Fouquet et al. (2007) found genetic divergence

Amphib. Reptile Conserv.

211A. Pecans Sp: al. fasaniiae Mérida, Bias ce

Barrio-Amoros.

in two populations of Lithodytes from Brazil and Peru.

Disjunct populations of Lithodytes in Venezuela may

represent undescribed species.

Selected references: Ginés (1959); Rivero (1961,

1964b); Lynch (1979b); Barrio-Amoros (1999d); Barros

and Barrio (2001); Sefiaris et al. (2009, 2014); Barrio-

Amoros et al. (2011b).

Subfamily Leiuperinae Bonaparte, 1850

Remarks: Grant et al. (2006) placed several genera into

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Barrio-Amoros et al.

family Leiuperidae, from a Leptodactylidae sensu Jato.

Pyron and Wiens (2011) ranked the taxon as a subfamily,

which 1s supported by Faivovich et al. (2012).

Genus Engystomops Jiménez de la Espada, 1872

Type species: Engystomops petersi Jiménez de la Espada,

1872, by monotypy.

Engystomops pustulosus (Cope 1864)

Holotype: USNM 4339 (lost).

Type locality: “New Grenada, on the River Truando,”

Colombia.

Distribution: Regions 1, 2, 3, 4, 5, 6. From Mexico

through Central America to Colombia, Venezuela,

Trinidad and Tobago, and Guyana in northern South

America. Widespread in lowlands in northern Venezuela.

Remarks: May hide a complex of species (Wynn and

Heyer 2002). At least two clear clades distinguishable,

one from Southern Mexico to Costa Rica, another from

Panama to northern South America (Weigt et al. 2005).

Call has been studied in many localities (Ryan et al. 2007).

Selected references: Lutz (1927); Ginés (1959); Rivero

(1961, 1963a, 1964a,c,d); Heatwole et al. (1965); Tello

(1968); Rivero and Esteves (1969); Lynch (1970);

Staton and Dixon (1977); Hoogmoed and Gorzula

(1979); Cannatella and Duellman (1984); Yustiz

(1996); Duellman (1997); Gorzula and Sefiaris (1998);

Nascimento et al. (2005); Weigt et al. (2005); Ryan et al.

(2007); Tarano (2010).

Genus Physalaemus Fitzinger, 1826

Type species: Physalaemus cuvieri Fitzinger, 1826, by

monotypy.

Remarks: A major review of this genus in Venezuela is

needed.

Physalaemus cuvieri Fitzinger, 1826

Type: Not stated.

Type locality: Brazil.

Distribution: Region 5. Venezuela, Brazil, Paraguay,

and Argentina. In Venezuela, only from one locality in

Bolivar State: Hato Terecay, 16 km N of El Manteco

(savannah). Herein a second population is reported from

Reserva Forestal Rio Grande, Sierra de Imataca, Delta

Amacuro State (MHNLS 20220-20223).

Remarks: In Physalaemus cuvieri species group of

Louren¢o et al. (2015). These authors noted this name

represents a complex of species. Gorzula and Sefiaris

(1999) indicated specimens referred for Venezuela as

Physalaemus cuvieri may be a related but undescribed

species.

Selected references: Gorzula and Sefiaris (1999);

Louren¢o et al. (2015).

Amphib. Reptile Conserv.

Physalaemus ephippifer (Steindachner 1864)

Holotype: NHMW; not traced.

Type locality: Restricted to Belém, Para State, Brazil, by

Bokerman (1966).

Distribution: Region 5. Venezuela, Guyana, Suriname,

French Guiana, and northeastern Brazil. In Venezuela,

only from a few localities in Bolivar State: Calceta

de Perro (rainforest), Las Claritas, and Santa Elena

de Uairen. Confused with P. fischeri and probably

most references to Physalaemus fischeri from eastern

Venezuela correspond to P. ephippifer.

Remarks: In Physalaemus cuvieri species group of

Nascimento et al. (2005).

Selected references: Gorzula and Sefiaris (1998);

Nascimento et al. (2005).

Physalaemus fischeri (Boulenger 1890)

Holotype: BMNH 1947.2.18.28.

Type locality: “Venezuela.”

Distribution: Regions 1, 2, 3, 4, 5. Llanos of Colombia

and Venezuela. Widespread in Northern Venezuela

(Cojedes, Guarico, Barinas, and Apure States), and

scattered to south of Orinoco River (Amazonas and

Bolivar States).

Remarks: Physalaemus enesefae Heatwole, Solano

and Heatwole, 1965 is a junior synonym of P. fischeri.

However, P. fisheri lacks a specific type locality and

is likely a species complex (Tarano and Ryan 2002;

Louren¢o et al. 2015). Its complex call is well-studied

(Tarano 2001, 2002, 2010; Tarano and Herrera 2003;

Tarano and Ryan 2002). In Physalaemus cuvieri species

group of Louren¢o et al. (2015).

Selected references: Boulenger (1890); Parker (1927);

Rivero (1961, 1967a); Heatwole et al. (1965); La Marca

(1992); Duellman (1997); Gorzula and Sefiaris (1998);

Tarano (2002, 2003, 2010); Barrio-Amoros et. al.

(2011b); Sefiaris et al. (2014).

Genus Pleurodema Tschudi, 1838

Type species: Pleurodema bibroni Tschudi, 1838, by

monotypy.

Pleurodema brachyops (Cope 1869)

Syntypes: ANSP 2260-2264.

Type locality: “Magdalene River, New Grenada” (=

Colombia).

Distribution: Regions 2, 3, 6. From Panama in Central

America through northern Colombia, Venezuela, and

Guyana to northern Brazil, plus Margarita Island and

the Netherlands Antilles (Curacao, Aruba, and Bonaire).

In Venezuela, widespread in lowlands north of Orinoco

River, northern Amazonas State, and eastern Bolivar

State south of Orinoco.

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Catalogue of the amphibians of Venezuela

“af ~ . ‘ , he . es ee a |

a ae a a =

211B. Leptodactylus sp. 1. El Tao, La Azulita, Mérida. Photo:

7, 3

— € = 2 BA

212B. Lithodytes lineatus. San Isidro, Barinas. Photo: Daniel

Ouihua.

214. Chiasmocleis hudsoni.

Fernando Rojas-Runjaic.

Selected references: Lutz (1927); Ginés (1959);

Rivero (1961, 1963a,c, 1964b,d, 1969a); Roze (1964):

Tello (1968); Leon-Ochoa and Donoso-Barros (1970);

Duellman and Veloso (1977); Staton and Dixon (1977),

Hoogmoed and Gorzula (1979); Ramo and Busto (1989,

1990); Barrio (1996a); Yustiz (1996); Gorzula and

Sefiaris (1998); Molina (2004 “2002”); Tarano (2010);

Ugueto and Rivas Fuenmayor (2010); Faivovich et al.

(2012).

Genus Pseudopaludicola Miranda-Ribeiro, 1926

Type species: Liuperus falcipes Hensel, 1867, by

Amphib. Reptile Conserv.

—

212A. Lithodytes lineatus. Cuyuni river, Bolivar. Photo: César

Barrio-Amoros.

iy as ay * Ac ees

213. Adelastes hylonomos. Paratype MBUCV 6185. Cerro de |

Neblina base camp. Photo: Roy McDiarmid.

215. Ctenophryne geayi. Imataca Forestal reserve, Delta Amacuro.

Photo: César Barrio-Amoros.

monotypy.

Pseudopaludicola boliviana Parker, 1927

Holotype: BMNH 1927.8.1.1.

Type locality: “Sta. Cruz, Bolivia.”

Distribution: Region 4. Colombia, Venezuela, Guyana,

Suriname, Brazil, Bolivia, Paraguay, and Argentina. In

Venezuela, south of Orinoco, but few precise localities

known. Besides Castillos de Guayana, Delta Amacuro

State (Gorzula and Sefiaris 1998), herein reported from

Capihuara, Casiquiare, Amazonas State, where it was

very abundant on river sand beaches in December 1998.

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

Remarks: This species has a disjunct distribution,

widely separated by the Amazon Basin. De la Riva et

al. (2000) suggested this greatly disjunct distribution

likely indicates at least two distinct species. Myers and

Donnelly (2001) presumed that Venezuelan populations

correspond to two species. In Pseudopaludicola pusilla

species group of Lynch (1989).

Selected references: Lynch (1989); Gorzula and Sefiaris

(1998); Lynch and Vargas-Ramirez (2000); Myers and

Donnelly (2001).

Pseudopaludicola llanera Lynch, 1989

Holotype: ICN 13576.

Type locality: “Puerto Gaitan, Departamento Meta,

Colombia.”

Distribution: Region 4. Northeastern Colombia and

southern Venezuela (Amazonas and Bolivar States);

eastern slopes of Andes in Merida State.

Remarks: In Pseudopaludicola pusilla species group of

Lynch (1989).

Selected references: Lynch (1989); Gorzula and Sefiaris

(1998); Lynch and Vargas-Ramirez (2000); Myers and

Donnelly (2001); Barrio and Chacon (2002).

Pseudopaludicola pusilla (Ruthven 1916)

Holotype: UMMZ 48305.

Type locality: “Fundacion, (Sierra de Santa Marta),

Colombia.”

Distribution: Region 6. Magdalena River Valley and

Caribbean lowlands of Colombia, and Maracaibo Lake

Basin in Venezuela.

Remarks: In Pseudopaludicola pusilla species group of

Lynch (1989).

Selected references: Ginés (1959); Rivero (1961, 1963a,

1964b,d); Lynch (1989).

Family Microhylidae Giinther, 1858

Remarks: Two recent phylogenetic systematics papers

on Microhylidae (De Sa et al. 2012; Peloso et al. 2016)

resulted in substantial changes in the taxonomy of the

group, both at subfamily and generic ranks. Both authors

found well resolved Gastrophryninae and Otophryninae;

the latter erected a new subfamily (Adelastinae). Most

generic changes proposed by De Sa et al. (2012) were

corroborated by later authors, except for the arrangement

of Syncope and Chiasmocleis. De Sa et al. (2012)

transferred several species of Chiasmocleis to Syncope,

whereas Peloso et al. (2014) placed Syncope in synonymy

of Chiasmocleis.

Subfamily Adelastinae Peloso, Frost, Richards,

Rodrigues, Donnellan, Matsui, Raxworthy, Biju,

Lemmon, Lemmon and Wheeler, 2016

Amphib. Reptile Conserv.

Genus Adelastes Zweifel, 1986

Type species: Adelastes hylonomos Zweifel, 1986, by

original designation.

Adelastes hylonomos Zweifel, 1986

Holotype: AMNH 123696.

Type locality: “Near the Neblina Base Camp on the

Rio Baria, 140 m elevation, 00°49’50”N, 66°09’40”W,

Rio Negro Department, Amazonas Federal Territory,

Venezuela.”

Distribution: Region 4. Venezuela, Brazil, and Guyana.

Known from type locality at the southern border of

Venezuelan Amazonas, from the proximity of Rio

Daraha, tributary of the Rio Negro, municipality of Santa

Isabel do Rio Negro, State of Amazonas, northern Brazil

(0°23°57.58’S, 064°47712.98”W), and from Guyana, at

plateau above Meamu River.

Remarks: Peloso et al. (2016) place Ade/astes in its own

subfamily Adelastinae. However, we believe it should

be relegated as part of Otophryninae, being clustered

basally with this last subfamily (Peloso et al. 2016: 136).

Call was recently described by De Almeida et al. (2014)

based on a single male.

Selected references: Zweifel (1986); McDiarmid and

Paolillo (1988); De Almeida et al. (2014); Peloso et al.

(2016).

Subfamily Gastrophryninae Fitzinger, 1843

Genus Chiasmocleis Méhely, 1904

Type species: Engystoma albopunctatum Boettger, 1885,

by monotypy.

Chiasmocleis hudsoni Parker, 1940

Type: BMNH 1939.1.1.3.

Type locality: “New River, British Guiana (750 feet).”

Distribution: Region 4. Colombia, Brazil, Venezuela,

Guyana, Suriname, and French Guiana. In Venezuela,

southern half of Amazonas State.

Remarks: Briefly transferred to Syncope by de Sa et al.

(2012) but again transferred to Chiasmocleis by Peloso

et al. (2014).

Selected references: Zweifel (1986); Barrio-Amoros

and Schargel (2003); De Sa et al. (2012); Peloso et al.

(2014, 2016).

Genus Ctenophryne Mocquard, 1904

Type species: Glossostoma aterrimum Gunther, 1900, by

monotypy.

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Catalogue of the amphibians of Venezuela

216A. Elachistocleis ovalis. Male calling. Chururu, Tachira.

Photo: César Barrio-Amoros.

rat

Jig f Spe : le ; { he we jt es Re ; S,, = = fe ;

216C. Elachistocleis aff. ovalis. San Diego de Cabrutica,

Anzoategui. Photo: César Barrio-Amoros.

Barrio-Amoros.

Ctenophryne geayi Mocquard, 1904

Holotype: MNHNP 1903.84.

Type locality: “La riviere Sarare en Colombie.”

Distribution: Region 4. Northern South America, to east

of Andes, in Colombia, Ecuador, Peru, Brazil, Venezuela,

Guyana, and Suriname. In Venezuela, two localities

in eastern Venezuela: km 13 Cuyuni, Bolivar State

(Duellman 1997), and Sierra de Imataca, Delta Amacuro

State (MHNLS 20230, reported herein).

Remarks: The type locality is “Riviere Sarare en

Colombie” (= Sarare River, Colombia). This river is born

in eastern side of Cordillera Oriental de Colombia and

flows through cloud forest and foothill rainforest with

Amazonian elements to Los Llanos Region. Though

Mocquard did not establish exact locality, current authors

Amphib. Reptile Conserv.

216B. Elachistocleis ovalis.

Barrio-Amoros.

216D. Elachistocleis aff. ovalis. San Diego de Cabrutica,

Anzoategui. Photo: César Barrio-Amoros.

217. Elachistocleis pearsei. Perija, Zulia. Photo: Fernando Rojas-

Runjaic.

estimate it must be in immediate forested foothills of the

Andes, where many Amazonian elements are known,

both in Colombia and Venezuela. However, it is strange

that no additional specimens have been reported from

that area in Venezuela and Colombia (in Colombia mostly

because of public order problems in the upper Llanos).

Lynch (2006) mentions its occurrence from Villavicencio

area in Colombia, but without voucher specimens.

Selected references: Carvalho (1954); Zweifel and

Myers (1989); Duellman (1997); Lynch (2006).

Genus Elachistocleis Parker, 1927

Type species: Not clearly stated: either Rana gibbosa

Linnaeus, by original designation, or Rana ovalis

Schneider, 1799, by subsequent designation of Dumeril

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Barrio-Amoros et al.

and Bibron, 1841 (sensu Frost 2019).

Remarks: Review of the genus is needed to clarify

chaotic taxonomic status of several populations

throughout its distribution. Duellman (1997) assigned

to Elachistocleis bicolor all forms with an immaculate

venter, whereas Lavilla et al. (2003) restricted E. bicolor

to southern South America. Similarly, E. ovalis is not the

species with spotted venter, as Schneider (1799) clearly

states, but one with “inferne flavidus” (yellow belly). De

Sa et al. (2012) placed Relictivomer into Elachistocleis.

Elachistocleis ovalis (Schneider 1799)

Type: Unknown.

Type locality: Not assigned.

Distribution: Regions 2, 3. Name Elachistocleis ovalis

historically applied to populations from Panama, Brazil,

Colombia, Venezuela, Ecuador, Peru, and Bolivia. In

Venezuela, present north of the Orinoco in open lowlands.

Remarks: Name Elachistocleis ovalis \ong-applied

to the Elachistocleis with a uniform yellow or orange

ventral pattern in northern Venezuela. Caramaschi (2010)

considered this name a nomen dubium (name of unknown

or doubtful application), and species inquirenda (species

of doubtful identity needing further investigation).

Current authors agree that a complex of species is hidden

under that name, and even knowing that the nomen ovalis

is not valid, but must be used while awaiting a definitive

solution and new available names for this long-standing

taxonomic conundrum.

Selected references: Lutz (1927); Carvalho (1954);

Ginés (1959); Rivero (1961, 1964b-d); Bogart and

Nelson (1976); Staton and Dixon (1977); Hoogmoed and

Gorzula (1979); Rada (1981a); Gremone et al. (1986);

Ramo and Busto (1989, 1990); Yustiz (1996); Gorzula

and Sefiaris (1998); Barrio and Durant (2000); Lavilla et

al. (2003); Caramaschi (2010); Tarano (2010).

Elachistocleis pearsei (Ruthven 1914)

Holotype: UMMZ 45571.

Type locality: “Vicinity of Fundacion, (Sierra de Santa

Marta), Colombia.”

Distribution: Panama, northern Colombia, and

Maracaibo Lake Basin in Venezuela, in open areas.

Remarks: Previously in monotypic genus Relictivomer

Carvalho, 1954, which was synonymized with

Elachistocleis by De Sa et al. (2012) based on its

phylogenetic position.

Selected references: Lutz (1927); Carvalho (1954);

Ginés (1959); Rivero (1961, 1964b-d); Bogart and

Nelson (1976); Staton and Dixon (1977); Hoogmoed and

Gorzula (1979); Rada (1981a); Gremone et al. (1986);

Ramo and Busto (1989, 1990); Yustiz (1996); Gorzula

and Sefiaris (1998); Barrio and Durant (2000); Lavilla

et al. (2003); Infante-Rivero et al. (2006b); De Sa et al.

(2012).

Amphib. Reptile Conserv.

Elachistocleis surinamensis (Daudin 1802)

Type: Unknown.

Type locality: “Surinam.”

Distribution: Regions 4, 5. Venezuela, Trinidad,

Guyana, Suriname, and French Guiana. In Venezuela,

present south of Orinoco River, including Orinoco Delta,

usually in open areas.

Remarks: All populations of E/achistocleis in Venezuela

with a gray venter, and orange or yellow spots, are

assigned to this species, although it must be a species

complex (Lavilla et al. 2003). A comprehensive review

of the genus and populations currently assigned to E.

surinamensis 1s needed to clarify taxonomic status of

Venezuelan populations.

Selected references: Rivero et al. (1986); Gorzula and

Sefiaris (1999); Sefiaris et al. (2014).

Genus Hamptophryne Carvalho, 1954

Type species: Chiasmocleis boliviana Parker, 1927, by

original designation.

Hamptophryne boliviana (Parker 1927)

Type: BMNH 1927.8.1.1.

Type locality: “Buena Vista, Santa Cruz, Bolivia.”

Distribution: Region 5. North and western portions of

Amazon Basin, in Venezuela, Colombia, Ecuador, Peru,

Bolivia, Brazil, Guyana, Suriname, and French Guiana.

Known to date from single locality in Venezuela: Reserva

Forestal Rio Grande, between Rio Grande and El Palmar,

Estado Bolivar (Paolillo 1986).

Remarks: Barrio-Amoros (2004) confirmed the identity

of the only voucher of this species from Venezuela (at

Berkeley, US). Image 219 shows a second specimen

from a nearby locality: Imataca, Delta Amacuro.

Selected references: Carvalho (1954); Bogart and

Nelson (1976); Paolillo (1986).

Subfamily Otophryninae

Wassersug and Pyburn, 1987

Remarks: Status of subfamily discussed in Wild (1995).

De Sa et al. (2012) and Peloso et al. (2016) reviewed

the taxonomy of Microhylidae and both recognize a

monophyletic Otophryninae containing Ofophryne and

Synapturanus. De Sa et al. (2012) found Otophryninae

well embedded into the family whereas Peloso et al.

(2016) found support for a sister taxon relationship of

Otophryninae, Gastrophryninae, and Adelastinae, thus

providing evidence for the monophyly of New World

microhylids. Current authors believe Ade/astes (currently

in Adelastinae) should be considered as in Otophryninae.

Genus Otophryne Boulenger, 1900

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Catalogue of the amphibians of Venezuela

: i : = —

= ae . , = 7

218A. Elachistocleis surinamensis. Kavanayen, Gran Sabana,

F i ‘ ae 5 : ? Bed * . rn — . s ?

219. Hamptophryne boliviana. Imataca Forestal Reserve, Bolivar-

Delta Amacuro border. Photo: Luis Merlo.

221. Otophryne steyermarki. Ayanganna tepui, Guyana. Photo:

Amy Lathrop-Ross MacCulloch.

Type species: Otophryne robusta Boulenger, 1900, by

original designation.

Otophryne pyburni Campbell and Clarke, 1998

Holotype: UTA-A3814.

Type locality: “Rainforest at 213 m elevation at Wacara,

Vaupés, Colombia (...) 01°09’N, 69°55’ W.”

Distribution: Region 4. Guiana Shield Region. Records

from Brazil, French Guiana, Guyana, Suriname, and

Venezuela. In Venezuela, at least three localities in

Amazonas state.

Selected references: Rivero (1967a); Nelson (1971);

Campbell and Clarke (1998); Barrio (1999e).

Amphib. Reptile Conserv.

kaa eee

EE. , . cx ol

Soe

220. Otophryne robusta. La Escalera, Sierra de Lema, Bolivar.

Photo: Hinrich Kaiser.

“.

- hehe a3 ee PE.

e: tu <.* * » has

PTs EP git ei it ha") as

222. Synapturanus mirandaribeiroi. Piedra de la Virgen, northern

slopes of La Escalera, Sierra de Lema, Bolivar. Photo: Eric Smith.

Otophryne robusta Boulenger, 1900

Type: BMNH 1899.3.25.18.

Type locality: “Foot on Mt. Roraima, 3,500 ft,” Guyana.

Distribution: Region 5. Gran Sabana, Bolivar State, in

eastern Venezuela, as well as Western Guyana.

Remarks: Boulenger (1900) stated specimens he

described from Roraima collected by Quelch and

Macconnell, were implicitly from Guyana. See further

comments on same type locality under accounts

of Oreophrynella macconnelli and _ Pristimantis

marmoratus. Could be in Venezuela, according to Kok

et al. (2018).

Selected references:

Boulenger (1900); Carvalho

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

(1954); Ginés (1959); Rivero (1961, 1964b, 1967a,b),;

Nelson (1971); Bogart et al. (1976); Gorzula (1985b,

1992); Gremone et al. (1986); Wassersug and Pyburn

(1987); Campbell and Clarke (1998); Gorzula and

Sefiaris (1998); MacCulloch et al. (2008); Cole et al.

(2013); Sefiaris et al. (2014).

Otophryne steyermarki Rivero, 1967

Holotype: FMNH 74031.

Type locality: “Forest along rio Tirica..., 5,900 ft, falls

below summit camp, Chimanta tepui massif, Venezuela.”

Distribution: Region 5. Known from three different

tepuis in Venezuela: Chimanta, Roraima, and Jaua

(Bolivar State), and Mount Ayanganna in Guyana.

Remarks: Previously considered endemic of the

Chimanta massif in the Venezuelan Guayana. Barrio

(1999f) and Gorzula and Sefiaris (1999) reported

additional localities in Venezuela, whereas MacCulloch

et al. (2008) provided records from Guyana, suggesting

a wider distribution. See redescription and data on

morphological variation in MacCulloch et al. (2008).

Selected references: Rivero (1967b); Gorzula (1985b,

1992); Campbell and Clarke (1998); Gorzula and Sefiaris

(1998); Barrio (1999f); MacCulloch et al. (2008); Sefiaris

et al. (2014).

Genus Synapturanus Carvalho, 1954

Type species: Synapturanus mirandaribeiroi Nelson and

Lescure, 1975

Synapturanus mirandaribeiroi

Nelson and Lescure, 1975

Holotype: MZUSP 49981.

Type locality: “Kanashen (a Waiwai Indian village

and mission) in the Upper Essiquibo River, Rupununi

District, Guayana.”

Distribution: Region 5. Colombia, Venezuela, Guyana,

Suriname, French Guiana, and Brazil. Only two localities

in Venezuela: Eastern slopes of Cerro Santa Rosa,

Serrania del Supamo (Barrio and Brewer-Carias 1999)

and Piedra de la Virgen, northern slopes of La Escalera,

Sierra de Lema, Bolivar (Image 222).

Selected references: Nelson and Lescure (1975); Pyburn

(1975); Barrio and Brewer-Carias (1999); Barrio-Amoros

et al. (2011b).

Synapturanus salseri Pyburn, 1975

Holotype: UTA A-4011.

Type locality: “Timbo, Vaupés,” Colombia.

Distribution: Regions 4, 5. Colombia, Venezuela,

Guyana, and Brazil. In Venezuela, only two localities at

northern and southern parts of Amazonas State.

Selected references: Pyburn (1975); Zweifel (1986);

Amphib. Reptile Conserv.

McDiarmid and Paolillo (1988); Sefiaris et al. (2003,

2014).

Family Phyllomedusidae Giinther, 1859 “1858”

Remarks: Recently elevated from subfamily to family

level by Duellman et al. (2016).

Genus Agalychnis Cope, 1864

Type species: Agalychnis callidryas Cope, 1862, by

original designation.

Remarks: Faivovich et al. (2005) passed the

Phyllomedusa buckleyi group sensu Cannatella (1980) to

Hylomantis, until Faivovich et al. (2010) extended the

definition of Agalychnis in order to include Hylomantis

+ Pachymedusa.

Agalychnis medinae (Funkhouser 1962)*

Holotype: EBRG 37.

Type locality: “The biological station ‘Henri Pittier’,”

Estado Aragua, Venezuela.

Distribution: Region 2. Endemic to Venezuela. Known

only from four localities (Rancho Grande in Aragua,

near Beyuma in Carabobo, Cerro Zapatero, and Sierra de

Aroa in Yaracuy) in the Coastal Range of Venezuela, but

apparently extinct from type locality.

Remarks: Named medinae (as in original description)

but later emended to medinai (since described after a

masculine patronym, and thereafter ended in -1). However,

article 31.1.1 of the Code admit a genitive ending in -ae

even for masculine names if they end in -a (as Medina).

Considered as Endangered by the current Venezuelan

Red Book (Rojas-Runjaic and Sefiaris 2015f). See

distribution map in Rojas-Runyaic et al. (2014).

Selected references: Funkhouser (1962); Rivero

(1967c); Duellman (1968, 1969, 1979b); Cannatella

(1980); Manzanilla et al. (1995); Proy (2000); Barrio-

Amoros (2001c, 2006c, 2009b, 2013); Lotzkat et al.

(2007); Barrio-Amoros and Torres (2010); Rojas-Runyjaic

et al. (2014); Rojas-Runjaic and Sefiaris (2015f).

Genus Callimedusa

Duellman, Marion and Hedges, 2016

Type species: Phy/lomedusa perinesos Duellman, 1973.

Remarks: Genus created by Duellman et al. (2016) to

accommodate the former Phyllomedusa perinesos group

plus P. atelopoides and P. tomopterna.

Callimedusa tomopterna (Cope 1868)

Syntypes: USNM 6651 (two specimens; lost).

Type locality: “Rio Napo, or Upper Amazon, below the

mouth of the former,’ Departamento Loreto, Peru.

Distribution: Regions 4, 5. Colombia, Ecuador, Peru,

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Pent re aE Re ede AP gre pent SE ants

223. Synapturanus salseri. Cuao, Amazonas. Photo: César

Barrio-Amoros.

224B. Agalychnis medinae. Amplexus. Cerro Zapatero, Nirgua

Massif, Yaracuy. Photo: Sebastian Lotzkat.

226. Pithecopus hypochonadrialis. Imataca Forestal Reserve, Delta

Amacuro. Photo: César Barrio-Amoros.

Bolivia, Brazil, Venezuela, Guyana, Suriname, and

French Guiana. In Venezuela, south of Orinoco River,

in states of Amazonas, Bolivar, and Delta Amacuro. See

list of localities and distribution map in Barrio-Amoros

(2009b).

Remarks: Divergence among samples of this species

from the upper Amazon and Guianas suggest at least

two species may be hidden within the name Callimedusa

tomopterna (Faivovich et al. 2010), but a thorough

revision is needed to assess this hypothesis.

Selected references: Funkhouser (1957); Duellman

Amphib. Reptile Conserv.

224A. Agalychnis medinae. El Silencio, Sierra de Aroa, Yaracuy.

Photo: Fernando Rojas-Runjaic.

225. Callimedusa tomopterna. Imataca Forestal Reserve, Delta

Amacuro. Photo: César Barrio-Amoros.

F a i

227. Phyllomedusa bicolor. Surroundings of Puerto Ayacucho,

Amazonas. Photo: César Barrio-Amoros.

(1968, 1974b, 1997); Cannatella (1980); Barrio and

Rivero (1999a); Barrio-Amoros (2009b); Faivovich et al.

(2010).

Genus Pithecopus Cope, 1866

Type species: Phyllonedusa azurea Cope, 1862, by

original designation.

Remarks: Genus recently resurrected of the synonymy

of Phyllomedusa by Duellman et al. (2016) to content the

old Phyllomedusa hypochondrialis group.

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Barrio-Amoros et al.

Pithecopus hypochondrialis (Daudin 1800)

Holotype: Unknown (but see Frost 2015).

Type locality: “Surinam.”

Distribution: Regions 3, 4, 5. Colombia, Brazil,

Venezuela, Guyana, Suriname, and French Guiana.

Widespread in lowland Venezuela in open areas, north

and south of Orinoco River. See list of localities and

distribution map in Barrio-Amoros (2009b).

Remarks: In Phyllomedusa hypochondrialis group of

Faivovich et al. (2010).

Selected references: Funkhouser (1957); Rivero (1961,

1964b); Heatwole et al. (1965); Duellman (1968, 1974b,

1997); Pyburn and Glidewell (1971); Hoogmoed and

Gorzula (1979); Rivero-Blanco and Dixon (1979); Ramo

and Busto (1989, 1990); Gorzula and Sefiaris (1998);

Rivas and Manzanilla (1999); Barrio-Amoroés (2009b);

Barrio-Amoros et al. (2010, 2011b); Faivovich et al.

(2010); Sefiaris et al. (2014).

Genus Phyllomedusa Wagler, 1830

Type species: Rana bicolor Boddaert, 1772 by monotypy.

Phyllomedusa bicolor (Boddaert 1772)

Holotype: Unknown.

Type locality: “Surinam” (fide Funkhouser, 1957).

Distribution: Regions 4, 5. Colombia, Peru, Bolivia,

Brazil, Venezuela, Guyana, Suriname, and French

Guiana. Widespread in southern Venezuela, Amazonas

and Bolivar States. See list of localities and distribution

map in Barrio-Amoros (2009b).

Remarks: Not assigned to any phylogenetic or phenetic

group by Faivovich et al. (2010).

Selected references: Funkhouser (1957); Ginés (1959);

Rohl (1959); Rivero (1961, 1964b); Heatwole et al.

(1965); Duellman (1968, 1974b, 1997); Gorzula and

Sefiaris (1998); Barrio-Amoros (2009b); Barrio-Amoros

et al. (2010, 2011b); Faivovich et al. (2010).

Phyllomedusa neildi Barrio-Amoros, 2006*

Holotype: MBUCV 6684.

Type locality: “Vicinity of Murucusa, Municipio Petit

(11°02'N, 69°35'W), 550 m asl., spurs of Sierra de San

Luis, Estado Falcon, Venezuela.”

Distribution: Region 2. NW Venezuela in Falcon and

Lara States.

Remarks: Phyllomedusa neildi appears as a sister species

to P. trinitatis in Fatvovich et al. (2010), suggesting it

could be conspecific, despite striking morphological

differences in size and call structure (Barrio-Amoros

2006b). This can be due to a clinal variation or adaptation

to xeric situations. However, direct comparison of

continental and Trinidadian populations of P. trinitatis

must be done (in progress by CBA and collaborators).

Amphib. Reptile Conserv.

Selected references: Barrio-Amorés (2006b, 2009b);

Faivovich et al. (2010).

Phyllomedusa tarsius (Cope 1868)

Holotype: USNM 6652; lost.

Type locality: “Rio Napo, or Upper Amazon, below the

mouth of the former” (Loreto, Peru).

Distribution: Regions 1, 3. Widely distributed in Amazon

Basin, in southwestern Colombia, Ecuador, Peru, Brazil,

with isolated populations in Cordillera Oriental of

Colombia and Venezuela. The Guyanan Phyllomedusa

tarsius (Forlani et al. 2012) must be compared with P.

trinitatis (see discussion in Barrio-Amorés 2009b).

In Venezuela, known only from a few localities in

eastern foothills of the Andes. See list of localities and

distribution map in Barrio-Amoros (2009b).

Remarks: We really doubt presence of Phyllomedusa

tarsius sensu stricto in Venezuela. Previous reports from

southeastern Venezuela were meant to be P. trinitatis

(Barrio-Amoros 2009b) or are very distinct from P.

tarsius sensu stricto, aS individuals from the Andean

piedmont reported by La Marca (1996) and Markezich

(1998) are much smaller and the iris pattern is much

less reticulated than on Ecuadorian (near type locality)

specimens (Image 229b). This could indicate a putative

new species of the farsius group in western Venezuela.

Selected references: Funkhouser (1957); Duellman

(1968, 1974b, 1997); La Marca (1996b); Barrio-Amoros

(2006b, 2009b); Sefiaris et al. (2014).

Phyllomedusa trinitatis Mertens, 1926

Holotype: SMF 2633.

Type locality: “Port of Spain,” Trinidad.

Distribution: Region 2. Trinidad and Venezuela. In

Venezuela, distributed across north of the country, to the

east of Falcon State. Not known from Los Llanos.

Remarks: Barrio-Amords (2009b) compared those

populations called Phyllomedusa tarsius from the

Venezuelan Guayana (reported by Duellman and

Trueb 1986) with P. trinitatis, and found no significant

morphological differences. Thus, the Guianan

populations are referred as P. trinitatis (see contrary view

in Sefiaris et al. 2014), pending more investigation of the

Andean piedmont populations named tarsius. Recently

reported P. tarsius from Guyana (Forlani et al. 2012)

must be compared to P. trinitatis. See comments under P.

neildi and P. tarsius. See list of localities and distribution

map in Barrio-Amoros (2009b).

Selected references: Mertens (1926); Funkhouser (1957,

1962); Ginés (1959); Rivero (1961, 1964a, 1969c);

Heatwole et al. (1965); Kenny (1966, 1969); Duellman

(1968, 1974b); Tello (1968); Mijares-Urruitia and

Arends (1993); Manzanilla et al. (1995); Barrio-Amoros

(2006b,c, 2009b); Faivovich et al. (2010); Forlani et al.

(2012); Sefiaris et al. (2014).

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Catalogue of the amphibians of Venezuela

228. Phyllomedusa_neildi. . Murucusa, Falcon. Photo: César

Barrio-Amoros.

_ RE =, ra —

par “4 i 2 SE . = ns Wa ol

229B. Phyllomedusa tarsius. Yasuni National Park, Ecuador.

Photo: Martin Bustamante.

230B. Phyllomedusa trinitatis. Cerro el Volcan, Caracas, Miranda.

Photo: Charles Brewer-Carias.

Phyllomedusa vaillanti Boulenger, 1882

Holotype: BM 1947.2.22.34.

Type locality: “Santarem, Brasil.”

Distribution: Region 4. Amazon Basin (Brazil,

Colombia, Ecuador, Peru, Bolivia, Venezuela) and

Guianas (Guyana, Suriname, and French Guiana). In

Venezuela, only reported from one locality (base of

Cerro La Neblina) in extreme southern Amazonas State.

Remarks: Unassigned to any species group.

Selected references: Funkhouser (1957); Duellman

(1968, 1974b); McDiarmid and Paolillo (1988); Barrio-

Amoros (2009b).

Amphib. Reptile Conserv.

229A. Phyllomedusa tarsius. Caparo Forestal Reserve, Barinas.

Photo: César Barrio-Amoros.

230A. Phyllomedusa trinitatis. Cerro el Volcan, Caracas, Miranda.

Photo: César Barrio-Amoros.

230C. Phyllomedusa trinitatis. Imataca Forestal Reserve, Delta

Amacuro. Photo: César Barrio-Amoros.

ns, *!

Phyllomedusa venusta Duellman and Trueb, 1967

Holotype: KU 96150.

Type locality: One km west-southwest of the junction

of the Rio Mono and the Rio Tuira, Darien Province,

Panama, 130 m.

Distribution: Region 6. Southern Panama, and Caribbean

lowlands of Colombia, entering the Magdalena River

Valley; in Venezuela, known from the Maracaibo Lake

Basin.

Remarks: Species in the Phyllomedusa tarsius group

(sensu Barrio-Amoros 2006 and Faivovich et al. 2010)

are very difficult to distinguish among them. Barrio-

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Barrio-Amoros et al.

Amoroés (2009b) comment on the possibility that P

venusta is conspecific with P. trinitatis, as the only

reliable character to differentiate them is the length of

the parotoid glands, a variable character influenced by

reproductive or hormonal condition.

Selected References: Duellman and Trueb (1967);

Barrio-Amoros (2006b, 2009b); Infante-Rivero et al.

(2006a); Faivovich et al. (2010).

Family Pipidae Gray, 1825

Remarks: Pyron and Wiens (2011) and Blackburn and

Wake (2011) refrain from using subfamilies within

Pipidae.

Genus Pipa Laurenti, 1768

Type species: Pipa Americana Laurenti, 1768 (= Rana

pipa Linnaeus, 1758), by monotypy.

Pipa arrabali \zecksohn, 1976

Holotype: EI 5311 (destroyed by fungus, Peixoto and

Gomes 2007).

Type locality: “Vila Amazonia, Municipio de Parintins,

Estado do Amazonas, Brasil.”

Distribution: Venezuela, Guyana, Suriname, and Brazil.

In Venezuela, known only from a few localities in eastern

Bolivar State.

Remarks: Specimens referred previously as Pipa aspera

by La Marca (1992) are in fact P. arrabali (Trueb and

Cannatella 1986). Both species very similar, differing only

by narrow or wide separation of the nostrils sensu Trueb

and Cannatella (1986). Both taxa are mostly sympatric,

so validity of P. arrabali is doubtful, it could be a junior

synonym of P. aspera despite the morphological support

offered by Trueb and Masserin (2001).

Selected references: Trueb and Cannatella (1986);

Duellman (1997); Peixoto and Gomes (2007); Barrio-

Amoroés and Duellman (2009); Barrio-Amoros et al.

(2011b); Sefiaris et al. (2014).

Pipa parva Ruthven and Gaige, 1923

Holotype: UMMZ 57443.

Type locality: “Sabana de Mendoza (Trujillo State),

Venezuela.”

Distribution: Northeastern Colombia and Northeastern

Venezuela. In Venezuela, natural populations restricted

to Maracaibo Lake Basin; introduced population

established in Lago de Valencia, Carabobo State (Royero

and Hernandez 1996).

Selected references: Barbour (1923); Ruthven and

Gaige (1923); Lutz (1927); Ginés (1958); Rivero (1961);

Trueb (1984); Trueb and Cannatella (1986); Barrio

(1996a); Royero and Hernandez (1996); Barrio and

Fuentes (2000b).

Amphib. Reptile Conserv.

Pipa pipa (Linnaeus 1758)

Syntypes: Specimens figured in Seba, 1734 (see http://

linnaeus.nrm.se/zool/herp/madamph.html.en)

Type locality: “Surinam.”

Distribution: Amazonian Region, in Colombia, Ecuador,

Peru, Bolivia, Brazil, Venezuela, Trinidad, Guyana,

Suriname, and French Guiana. In Venezuela, widespread

in Orinoco Basin, from eastern foothills of the Andes

at Barinas State, Llanos of Monagas, Delta Amacuro,

Bolivar, and Amazonas.

Selected references: Dunn (1948); Ginés (1958); Rohl

(1959); Rivero (1961); Trueb and Cannatella (1986);

Péfaur and Diaz De Pascual (1987); La Marca (1992);

Gorzula and Sefiaris (1998).

Family Ranidae Rafinesque-Schmaltz, 1814

Subfamily Raninae Rafinesque-Schmaltz, 1814

Genus Lithobates Fitzinger, 1843

Type species: Rana palmipes Fitzinger, 1843 (= Rana

palmipes Spix, 1824), by designation.

Lithobates catesbeianus (Shaw 1802)

Holotype: Illustrated by Shaw (1802: 106, plate 33).

Type locality: “Many parts of North America...

Carolina, Virginia.” Attempts to restrict type locality are

not accepted by Fouquette and Dubois (2014).

Distribution: Southern Canada, eastern North America,

and northern Mexico. Introduced 1n numerous countries

of Americas (southern and western Mexico, Colombia,

Ecuador, Venezuela, Guyana, Brazil, Paraguay,

Argentina, Chile) several Caribbean islands (Cuba,

Puerto Rico, Jamaica, Hispaniola), Europe (Netherlands,

France, Belgium, Italy, Spain, Greece), and Asia

(Malaysia, Indonesia, Japan, Thailand, Korea, and

Taiwan). In Venezuela, known introduced population in

the surroundings of La Carbonera, Mérida State.

Remarks: Since its introduction, species was the subject

of a population control program by ULA (Universidad de

los Andes), IVIC (Instituto Venezolano de Investigaciones

Cientificas), and MARN (Ministerio del Ambiente y

los Recursos Naturales), without continuation or much

SUCCESS.

Selected references: = Barrio-Amoros = (2001c);

Hanselmann et al. (2004); Lampo et al. (2004); Barrio-

Amoros et al. (2011).

Lithobates palmipes (Spix 1824)

Syntypes: ZSM, including ZSM 963/0, lost.

Type locality: “In aquis stagnantibus fluminis

Amazonum” (= stagnant waters of the Amazon River),

Brazil.

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—_ “ ee

231. Phyllomedusa vaillanti. Yasuni, Ecuador. Photo: César

Barrio-Amoros.

Oe | tii AEM Oe

f a \l * «at 3.

; Eoariapr. ae:

{ i « i

i 1 » ae

Bg » i ‘ me ' ! +

233. Pipa arrabali. La Laja, Sierra de Lema, Bolivar. Photo:

César Barrio-Amoros.

235A. Pipa pipa. Capihuara, Casiquiare, Amazonas. Photo: César

Barrio-Amoros.

Distribution: Northern South America, including

Colombia, Venezuela, Guyana, Suriname, Brazil,

Ecuador, Peru, and Bolivia. Widespread in Venezuela

except in Los Llanos.

Remarks: Venezuelan specimens (from Cerro de la

Neblina, the southernmost point of the country) used in

a phylogeny by Hillis and Wilcox (2005) stand out from

Lithobates palmipes. The geographically closest available

name for such populations is Ranula gollmeri Peters,

1859 (Hillis and Wilcox 2005). However, Lithobates

palmipes must be a species complex, so more intensive

Amphib. Reptile Conserv.

8 £

232. Phyllomedusa venusta. La Orchila, Sierra de Perija, Zulia.

Photo: Fernando Rojas-Runjaic.

vd . ' ~ i, “

234. Pipa parva. Female. El Vigia, Mérida. Photo: César Barrio-

Amoros.

We “

235B. Pipa pipa. Cafio Morichal Largo, Monagas. Photo: Eduardo

Asens.

sampling would be desired to apply that name, given that

L. gollmeri 1s from Caracas, and few species are proven

to inhabit both far north and far south Venezuela, in very

different habitats. On the other hand, Lithobates vaillanti

(Brocchi 1877) is known to occur on the Colombian side

of Serrania de Perija, and Venezuelan specimens from

western Zulia State could be mistaken for it.

Selected references: Spix (1824); Lutz (1927); Ginés

(1959); Rohl (1959); Rivero (1961, 1964a,b, 1967a,

1971a); Heatwole et al. (1965); Tello (1968); Hoogmoed

and Gorzula (1979); Gremone et al. (1986); Rivero et

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

al. (1986); Hillis and De Sa (1988); Duellman (1997);

Gorzula and Sefiaris (1998); Barrio-Amoros and Brewer-

Carias (2008); Barrio-Amoros et al. (2011b); Sefiaris et

al. (2014).

Family Strabomantidae

Hedges, Duellman and Heinicke, 2008

Incertae sedis:

Genus Dischidodactylus Lynch, 1979

Type species: Elosia duidensis Rivero, 1968, by original

designation.

Remarks: Hedges et al. (2008) tentatively placed

Dischidodactylus into Strabomantidae: Strabomantinae,

due to only one morphological character (possession of

expanded terminal disks with circumferential grooves).

Because of external similarities with Ceuthomantis, Padial

et al. (2014) placed Dischidodactylus in the subfamily

Ceuthomantinae. However, the phylogenetic position of

genus Dischidodactylus has never been evaluated using

molecular data, and it differs greatly from Ceuthomantis

and all other Terraranae in external morphology. Further

research should clarify relationships. Meanwhile, this

genus is left as incertae sedis into a comprehensive

Strabomantidae.

Dischidodactylus colonnelloi Ayarzagiiena, 1985*

Holotype: MHNLS 9378.

Type locality: “Cima del Tepui Marahuaca (Marahuaca-

tepui), Estado Amazonas, Venezuela. 2,550 m.”

Distribution: Region 5. Endemic to the summit of

Cerro Marahuaca, a tepui in Amazonas State, southern

Venezuela.

Selected references: Ayarzagtiena (1985).

Dischidodactylus duidensis (Rivero 1968)*

Holotype: AMNH 23190.

Type locality: “Summit at Vegas Falls, 4,600 feet (1,400

m), Mt. Duida, Venezuela.”

Distribution: Region 5. Endemic to Cerro Duida, a tepui

in Amazonas State, southern Venezuela.

Selected references: Rivero (1968a, 1971b); Lynch

(1979a); McDiarmid and Donnelly (2005).

Subfamily Pristimantinae Ohler and Dubois, 2012

Genus Pristimantis Jiménez de la Espada, 1870

Type species: Pristimantis galdi Jiménez de la Espada,

1870, by monotypy.

Remarks: Heinicke et al. (2007) split long-established

genus Eleutherodactylus, formerly in _ family

Leptodactylidae (Lynch 1971, 1981; Frost 1985;

Amphib. Reptile Conserv.

Duellman 1993; Lynch and Duellman 1997) and placed

afterwards in family Brachycephalidae by Frost et al.

(2006) containing three major genera: Eleutherodactylus

for mainly Caribbean species, Craugastor Cope, 1862 for

mainly Central American species, and Pristimantis for

almost all South American (and some Central American)

species. Hedges et al. (2008) offered a more comprehensive

view of the so-called “eleutherodactylines” and erected the

new unranked taxon Terrarana (emended as Terraranae by

Heinicke et al. 2018) for four families (Brachycephalidae,

Craugastoridae, Eleutherodactylidae, and Strabomantidae).

As of this writing, the phylogenetic arrangement remains

in flux. Most South American species formerly in

Eleutherodactylus are now members of Pristimantis,

within Craugastoridae (after Padial et al. 2014). The

most recent approach (Heinicke et al. 2018) recovers

Tachiramantis as part of Craugastoridae, and Pristimantis

into Strabomantidae. Mucubatrachus La Marca, 2007

and Paramophrynella La Marca, 2007 are considered

synonyms of Pristimantis sensu (Hedges et al. 2008;

Barrio-Amoros et al. 2013; Padial et al. 2014).

Selected references: Lutz (1927); Lynch (1976); La

Marca (1992); Barrio-Amorés (1998, 2004, 2009);

Heinicke et al. (2007); Hedges et al. (2008); Barrio-

Amoréos et al. (2013); Padial et al. (2014).

Pristimantis abakapa

Rojas-Runjaic, Salerno, Sefiaris and Pauly, 2013*

Holotype: MHNLS 20544.

Type locality: Abakapa-tepui, Macizo de Chimanta,

Parque Nacional Canaima, Gran Sabana municipality,

Bolivar State, Venezuela (5°11731.2”N, 62°18°56.6”W;

elevation 2,245 m).

Distribution: Region 5. Endemic to Abakapa-tepui in

the Chimanta massif, Venezuelan Guayana Region.

Remarks: In non-monophyletic Pristimantis unistrigatus

Species group sensu Rojas-Runyaic et al. (2013). Not

assigned to any species group by Padial et al. (2014).

Selected references: Rojas-Runjaic et al. (2013); Sefiaris

et al. (2014).

Pristimantis ameliae Barrio-Amoros, 2011*

Holotype: CVULA 7118.

Type locality: San Javier del Valle, 8°56’43”N,

70°25’54°W, elevation 2,500 m, Mérida State, Venezuela.

Distribution: Region 1. Endemic to eastern versant of

Sierra de la Culata of the Venezuelan Andes.

Remarks: Not assigned to any species group (Padial et

al. 2014).

Selected references: Barrio-Amoros (2011).

Pristimantis anolirex (Lynch 1983)

Holotype: KU 168626.

Type locality: “18.5 km (by road) S_ Chitaja,

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Catalogue of the amphibians of Venezuela

236. OBES rane La Gato Mend Photo:

César Barrio-Amoros.

237. GEES Sarees Arassari Rafting ‘Canips Acequias,

Barinas. Photo: César Barrio-Amoros.

238. Dischidodactylus colonnelloi. Holotype MHNLS 9378, Gal of the

right side, inside right hand from below. Photo: César Barrio-Amoros.

239, Pristimantis Bianabe Abekapacoul Chimanté Tain Gani

Sabana, Bolivar. Photo: Fernando Rojas-Runjaic.

240. Pristimantis cf. ameliae. Way to Paramo Los Conejos, 2500

masl. Photo: Daniel Quihua.

Departamento de Norte de Santander, Colombia, 2,850

Distribution: Region 1. Cordillera Oriental of the

Colombian Andes and Paramo de Tama, paramos in SW

Tachira State in the Venezuelan Andes.

Remarks: La Marca (1995d) reported this species

from Venezuela. In non-monophyletic Pristimantis

unistrigatus species group sensu Hedges et al. (2008),

but not assigned to any species group by Padial et al.

(2014).

Selected references: Lynch (1983); La Marca (1995d).

Pristimantis anotis (Walker and Test 1955)*

Amphib. Reptile Conserv.

241. Pristimantis anolirex. Paramo Tama, Norte de Santander,

Colombia. Photo: Aldemar A. Acevedo.

Holotype: UMMZ 109876.

Type locality: Rancho Grande, 1,090 m, Aragua State,

Venezuela.

Distribution: Region 2. Endemic to Venezuela.

Apparently restricted to surroundings of type locality.

Remarks: A severe decline in this species 1s presumed,

as not seen since 1974 (Barrio-Amoros 2006c). In non-

monophyletic Pristimantis unistrigatus species group

sensu Lynch and Duellman (1997) and Hedges et al.

(2008); not assigned to any species group by Padial et

al. (2014).

Selected references: Walker and Test (1955); Ginés

(1959); Rivero (1961, 1964a); Manzanilla et al. (1995),

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Barrio-Amoros et al.

Barrio-Amoros (2006c); Kaiser et al. (2015).

Pristimantis aureoventris

Kok, Means and Bossuyt, 2011

Holotype: IRSNB 4152.

Type locality: Summit of Wei-Assiput-tepui, Cuyuni-

Mazaruni District, Guyana (5°13’05”N, 60°42715”’W,

2,210 m).

Distribution: Region 5. Guyana Shield, in western

Guyana and eastern Venezuela. In Venezuela, known

from Kukenan-tepui (Jablonski et al. 2017) and here

reported from summit of Mt. Roraima, Bolivar State.

Remarks: Listed by Sefiaris et al. (2014), who anticipated

its presence in Venezuela (Canaima National Park) but

without any voucher specimen. Jablonski et al. (2017)

reports it for first time in Venezuela from the summit of

Kukenan-tepui, in the eastern chain of tepuis, Bolivar

State. Two vouchers are added here from the summit of

Mt. Roraima (MCNC 7196 and EBRG 200), at 2,800

m. Detailed comparison fails to distinguish them from

the original description of Pristimantis aureoventris,

from a nearby tepui (Wei-Assipu-tepui) in Guyana, and

the Guyanan side of Mt. Roraima. The photographs by

R.W. McDiarmid and Thomas Marent (Images 243A and

243B) are almost identical to those of Kok et al. (2011;

see Fig. 4C, E). Not assigned to any species group (Padial

et al. 2014).

Selected references: Kok et al. (2011); Sefiaris et al.

(2014); Jablonski et al. (2017).

Pristimantis auricarens (Myers and Donnelly 2008)*

Holotype: EBRG 2725.

Type locality: Summit of Auyan-tepui, AMNH-

TERRAMAR Camp 2, 1,750 m, Bolivar State, Venezuela

(5°54’N, 62°29°W).

Distribution: Region 5. Apparently endemic to Auyan-

tepui in Venezuelan Guyana Shield.

Remarks: In Pristimantis unistrigatus species group

according to original description (Myers and Donnelly 2008);

not assigned to any species group by Padial et al. (2014).

Selected references: Myers and Donnelly (2008);

Sefiaris et al. (2014).

Pristimantis avius (Myers and Donnelly 1997)*

Holotype: AMNH 131481.

Type locality: “North base of Pico Tamacuari, 1,160-

1,200 m elevation. Sierra Tapirapecd, Amazonas,

Venezuela (1°13’N, 64°42’W).”

Distribution: Region 5. Endemic to Sierra Tapirapeco,

Venezuelan Amazonas. Possibly present in Brazilian side

of Tapirapeco.

Remarks: In non-monophyletic Pristimantis unistrigatus

species group according to Myers and Donnelly (1997)

and Hedges et al. (2008); not assigned to any species

Amphib. Reptile Conserv.

group by Padial et al. (2014).

Selected references: Myers and Donnelly (1997).

Pristimantis bicumulus (Peters 1863)*

Syntypes: ZMB 4899 (two specimens).

Type locality: Caracas, Venezuela.

Distribution: Region 2. Cordillera de la Costa,

Venezuela.

Remarks: Not assigned to any species group (Hedges et

al. 2008; Padial et al. 2014).

Selected references: Lutz (1927); Ginés (1959); Rivero

(1961, 1964a); Tello (1968); Lynch and La Marca (1993);

Manzanilla et al. (1995); Barrio-Amoros (2006c); Sefiaris

and Rojas-Runjaic (2015c).

Pristimantis boconoensis (Rivero and Mayorga 1973)*

Type: UPRM 4932.

Type locality: “Paramo Guaramacal, Bocono, Estado

Trujillo, Venezuela, 9,400 feet” (2,865 m).

Distribution: Region 1. Subpdramo and paramo habitats

in Trujillo State, Venezuelan Andes.

Remarks: Not assigned to any species group (Hedges et

al. 2008; Padial et al. 2014).

Selected references: Rivero and Mayorga (1973); Lynch

(1976); Duellman (1979b); Rivero (1982b); La Marca

(2007).

Pristimantis briceni (Boulenger 1903)*

Syntypes: MCZ 3888 and 7601, NHMW 22871 (3

specimens); UMMZ 46471.

Type locality: “Merida (Merida), Venezuela, at an altitude

(elevation) of 1,600 metres.” Elevation may be in error, as

it is known from 2,600 to 3,400 m; 1,600 m Is the elevation

of Merida, from where the specimens were shipped.

Distribution: Region 1. Sierra de la Culata in subpdramo

and paramo habitats in Venezuelan Andes of Merida State.

Remarks: Considered type species of genus

Mucubatrachus La Marca, 2007. Barrio-Amoros et al.

(2013) show this species is phylogenetically embedded 1n

Pristimantis. Not assigned to any species group (Hedges

et al. 2008; Padial et al. 2014).

Selected references: Boulenger (1903); Lutz (1927);

Ginés (1959); Rivero (1961, 1982c, 1983); Lynch (1976);

La Marca (2007); Barrio-Amoros et al. (2013).

Pristimantis cantitans (Myers and Donnelly 1996)*

Holotype: EBRG 3005.

Type locality: “Summit of Cerro Yavi, 2,150 m,

Amazonas, Venezuela.”

Distribution: Region 5. Endemic to Cerros Yavi and

Yutajé, two tepuis in northern Amazonas State.

Remarks: In non-monophyletic Pristimantis unistrigatus

species group according to Lynch and Duellman (1997),

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Catalogue of the amphibians of Venezuela

242. Pristimantis anotis. Rancho Grande, Henri Pittier National

Park. Photo: William. E. Duellman.

F Faas be" ky

a a Ne = es

i *

243B. Pristimantis aureoventris. Summit of Rora

Photo: Thomas Marent.

ima, Venezuela.

244A. Pristimantis bicumulus. Rancho Grande, Henri Pittier

National Park. Photo: Javier Mesa.

Myers and Donnelly (2001), and Hedges et al. (2008) not

assigned to any species group by Padial et al. (2014).

Selected references: Myers and Donnelly (1996, 2001).

Pristimantis colostichos (La Marca and Smith 1982)*

Holotype: UMMZ 173044.

Type locality: “Paramo de los Conejos at the intersection

of Quebrada Las Gonzales with the trail Manzano Alto-

Las Gonzales, 2.5 hours on foot from the water pipeline

Amphib. Reptile Conserv.

243A. Pristimantis aureoventris. Summit of Roraima, Venezuela.

Photo: Roy McDiarmid.

243C. Pristimantis aureoventris. Slopes of northern Roraima,

Guyana. Photo: Bruce Means.

244B. Pristimantis bicumulus. Nirgua Massif, Yaracuy. Photo:

Sebastian Lotzkat.

known as “Las Canalejas,” Serrania del Norte, Estado

Merida, Venezuela.”

Distribution: Region |. Known only from type locality

in Sierra de la Culata, Mérida State, Venezuelan Andes.

Remarks: In non-monophyletic Pristimantis unistrigatus

Species group according to Hedges et al. (2008); not

assigned to any species group by Padial et al. (2014).

Selected references: La Marca and Smith (1982).

Pristimantis culatensis (La Marca 2007)*

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Barrio-Amoros et al.

Holotype: ULABG 912.

Type locality: “Paramo La Culata, 2,870 m (8°44’33’N,

71°04’20”"W), Municipio Libertador, Estado Merida,”

Venezuela.

Distribution: Region 1. Known only from type locality

in Sierra de la Culata, Mérida, Venezuelan Andes.

Remarks: Described without comparison to its apparently

closest relatives from the same area, Pristimantis briceni

and P. colostichos. Not assigned to any species group (La

Marca 2007; Hedges et al. 2008; Padial et al. 2014).

Selected references: La Marca (2007).

Pristimantis conservatio

Barrio-Amoros, Heinicke and Hedges, 2013*

Holotype: CVULA 7174.

Type locality: Cloud forest immediately above hamlet of

Los Alcaravanes at Aguas Blancas, 1,640 m, 8°56’43”N,

70°25°54"W; Ramal de Calderas, Barinas State,

Venezuela.

Distribution: Region 1. Endemic from cloud forests in

Venezuelan Andes. Known only from eastern versant

of Cordillera de Mérida in Barinas State. Probably also

present in continuous cloud forest in Trujillo State.

Remarks: Not assigned to any species group Padial et

al. (2014).

Selected references: Barrio-Amoroés (2010a); Barrio-

Amoros et al. (2013).

Pristimantis fasciatus

Barrio-Amoros, Rojas-Runjaic and Infante, 2007*

Holotype: MHNLS 18466.

Type locality: Kunana, cuenca del rio Negro, Sierra

de Perija, Municipio Machiques de Perija, Zulia State,

Venezuela (10°02’N, 72°47’W; 1,094 m).

Distribution: Region |. Only known from three localities

at eastern versant of Sierra de Perija, at 800—1,200 m.

Remarks: Considered Endangered according to

Venezuelan Red Book (Royas-Runyjaic and Sefiaris

2015g). In nonmonophyletic Pristimantis unistrigatus

group according to original description (Barrio-Amoros

et al. 2007), but not assigned to any species group by

Padial et al. (2014).

Selected references: Barrio-Amoros et al. (2007); Rojas-

Runjaic and Sefiaris (2015g).

Pristimantis flabellidiscus (La Marca 2007)*

Holotype: ULABG 2883.

Type locality: Los Aranguren, 2,860 m, (ca. 8°35’59’N,

70°55’28”W), southern versant of Sierra Nevada National

Park, Rangel Municipality, Merida State, Venezuela.

Distribution: Region 1. Known only from type locality.

Remarks: Not assigned to any species group (Hedges et

al. 2008; Padial et al. 2014).

Selected references: La Marca (2007).

Amphib. Reptile Conserv.

Pristrimantis geminus Kaiser, Barrio-Amoros, Rivas-

Fuenmayor, Steilein and Schmidt, 2015*

Holotype: USNM unnumbered (field number CMD

341).

Type locality: Southern slopes of Cerro Humo

(10.7073°N, 62.6284°W), elevation 750 m, Sucre State,

Venezuela.

Distribution: Region 2. Apparently restricted to

Peninsula de Paria.

Remarks: In nonmonophyletic Pristimantis unistrigatus

Species group according to the original description

(Kaiser et al. 2015).

Selected references: Kaiser et al. (2015).

Pristimantis ginesi (Rivero 1964)*

Holotype: MHNLS 250, sensu Frost (1985) and La

Marca (1992).

Type locality: Laguna Mucubaji, Rangel Municipality,

Merida State, Venezuela.

Distribution: Region |. Paramos of the Sierra Nevada,

Merida State.

Remarks: Considered type species of genus

Paramophrynella by La Marca (2007). Using molecular

data, Barrio-Amoros et al. (2013) demonstrated this

species is phylogenetically embedded in Pristimantis.

Selected references: Rivero (1964d, 1967c, 1982b,

1983); Rivero and Mayorga (1973); Lynch (1976); La

Marca (1991b “1994,” 2007); Barrio-Amoros et al.

(2013).

Pristimantis gryllus

Barrio-Amoros, Guayasamin and Hedges, 2012

Holotype: CVULA 8343.

Type locality: “Cloud forest along the road from

Estanques to Paramo la Tosta, 8.4333N, 71.5000W,

1,320 m, Estado Merida, Venezuela.”

Distribution: Region 1. Cloud forest and perianthropic

situations in Andes of Tachira and Merida States in

Venezuela, recently reported for Norte de Santander in

Colombia (Acevedo-Rincon et al. 2014).

Remarks: In Pristimantis unistrigatus species group

according to original description (Barrio-Amoros et al.

2012), but not assigned to any species group by Padial

et al. (2014).

Selected references: Barrio-Amoros et al. (2012);

Acevedo-Rincon et al. (2014).

Pristimantis guaiquinimensis

(Schliiter and Rodder 2007)*

Holotype: SMNS 8004.1.

Type locality: Guaiquinima tepui, Bolivar State,

Venezuela (5°44’N, 63°38’ W, elevation 980 m).

Distribution: Region 5. Endemic to Guaiquinima, a

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Catalogue of the amphibians of Venezuela

Photo: Juan E. Garcia.

247. Pristimantis colostichos. Paramo Los Conejos, 3,000 m asl.

Photo: Daniel Quihua.

7 % al

249. Pristimantis fasciatus. Male paratype LS 18470.

Kunana, Rio Negro basin, Serrania de Perija, Zulia. Photo:

Fernando Rojas-Runjaic.

tepui in Bolivar State, Venezuelan Guayana.

Remarks: In Pristimantis unistrigatus species group

according to original description (Schliiter and Rodder

2007) and Hedges et al. (2008), but not assigned to any

species group by Padial et al. (2014). Eleutherodactylus

tepuiensis Schliter and R6dder, 2007 is a junior synonym

Amphib. Reptile Conserv.

246. Pristimantis briceni. Cabafia del Cura, Sierra de la Culata,

Merida. Photo: César Barrio-Amoros.

248. Pristimantis conservatio. Female holotype CVUI A 7174.

Los Alcaravanes, Calderas, Barinas. Photo: César Barrio-Amoros.

250. Pristimantis geminus. Paratype CVULA 7443. Las Melenas,

southern slope of Cerro El Humo, Peninsula de Paria, Sucre.

Photo: César Barrio-Amoros.

(Kok and Barrio-Amoros 2013).

Selected references: Schliiter and Rodder (2007); Kok

and Barrio-Amoros (2013).

Pristimantis hoogmoedi Kaiser, Barrio-Amor6os, Rivas-

Fuenmayor, Steilein and Schmidt, 2015*

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Barrio-Amoros et al.

Holotype: CVULA 7433.

Type locality: “Southern slopes of Cerro Humo

(10.7073°N, 62.6284°W), elevation 750 m, Estado

Sucre, Venezuela.”

Distribution: Region 2. Apparently endemic to southern

versant of Peninsula de Paria in northeastern Venezuela.

Remarks: Most similar to Pristimantis anotis. Not

assigned to any species group according to original

description (Kaiser et al. 2015). Apparently suffering

some decline (G. Rivas, pers. comm. ).

Selected references: Kaiser et al. (2015).

Pristimantis imthurni Kok, 2013*

Holotype: IRSNB 4165.

Type locality: Summit of Ptari-tepui, Bolivar State,

5°46’09’”N, 61°49°02”W, 2,471 m elevation.

Distribution: Region 5. Endemic to Ptari, a tepui in

Bolivar State, Venezuela.

Remarks: Not assigned to any species group (Padial et

al. 2014).

Selected references: Kok (2013); Sefiaris et al. (2014).

Pristimantis incertus (Lutz 1927)*

Holotype: AL-MNRJ 1190.

Type locality: Mamo, near La Guaira, Vargas State,

Venezuela.

Distribution: Region 2. Restricted to central Cordillera

de la Costa in Aragua, Carabobo, Distrito Capital,

Miranda, and Vargas States, and western Cordillera de la

Costa in Nirgua massif, Yaracuy State, 20—1,500 m asl.

Remarks: Hylodes anonymus was first mentioned by

Lutz (1927: 40) as a new species, but description lacked

designation of a type specimen. Later in same paper,

two illustrations appear on Plate 13 (Image 256A) but

associated text refers to Hylodes incertus, here assumed to

be a lapsus and unjustified emendation for H. anonymus.

These names appear alternatively in recent checklists (La

Marca 1992; Barrio-Amoros 1998, 2004, 2009), but none

are identified with identifiable specimens or vouchers.

Hylodes anonymus is therefore a nomen nudum under

Article 12 of the Code (ICZN 1999), whereby names

without illustration or diagnosis coined prior to 1931

are not valid for the purposes of nomenclature. The case

of Hylodes incertus is different. Since Adolpho Lutz’s

material was stored at Museum of the Universidade

Federal do Rio de Janeiro, Brazil (MNRJ), the current

number assigned to the holotype of H. incertus is AL-

MNRJ 1190. Having received photos of the specimen

thanks to Dr. José Pombal and, despite the complete

fading of the color (Image 256B), it clearly belongs to a

member of the Pristimantis conspicillatus species group.

Members of this group are easily distinguished by finger

I being equal or longer than finger II, clearly visible in

Image 256C. Pristimantis terraebolivaris (Rivero 1961)

is the only species in Cordillera de la Costa in the P.

Amphib. Reptile Conserv.

conspicillatus species group and is abundant in the central

sector of the Cordillera de la Costa in Aragua, Carabobo,

Miranda, and Yaracuy States and Distrito Capital.

Mamo, near La Guaira (Vargas State), the type locality

of H. incertus, is at 85 m asl. The previously known

altitudinal range of P. terraebolivaris was 800—1,500 m

(Barrio-Amoros 2004); at MHNLS specimens are from

20-1,500 m. Comparing the photos and illustration of

Hylodes incertus with specimens and the description

of Pristimantis terraebolivaris (Rivero 1961) indicate

those are conspecific and Hylodes incertus Lutz, 1927

is a senior synonym of Eleutherodactylus terraebolivaris

Rivero, 1961; the color pattern of H. incertus holotype

corresponds with the stripped morph recognized by

Barrio-Amoros (2006c). Despite the poor conservation

status of the H. incertus holotype, it does match with

Pristimantis terraebolivaris in having the head longer

than wide; long snout (longer than eye diameter),

acuminate in dorsal view; canthus rostralis straight and

transversely angular; loreal region vertical, finger I equal

to finger II, fingers and toes with lateral keels; tip of toe

II not reaching the middle subarticular tubercle of toe

IV, and tip of toe V just passing the middle subarticular

tubercle of toe IV. No other Pristimantis species in the

Cordillera de la Costa matches with this combination of

characters. Thus, all previous references to Pristimantis

terraebolivaris must be now attributed to Pristimantis

incertus. In Pristimantis conspicillatus species group

(Hedges et al. 2008; Padial et al. 2014). Barrio-Amoroés

(2006c) showed some chromatic variation of the species.

Selected references: All references mentioning

Eleutherodactylus or Pristimantis terraebolivaris. Lutz

(1927); Rivero (1961, 1964a,d); Heatwole (1963a); Tello

(1968); Lynch (1976, 1979b); Barrio-Amorés (1996a,

2006c); Lotzkat (2007).

Pristimantis jabonensis (La Marca 2007)*

Holotype: ULABG 2813.

Type locality: “Paramo El Jabon, 3,100 m, Municipio

Carache, Estado Trujillo, Venezuela.”

Distribution: Region 1. Known only from type locality.

Remarks: Not assigned to any species group (Hedges et

al. 2008; Padial et al. 2014).

Selected references: La Marca (2007).

Pristimantis kareliae (La Marca 2005)*

Holotype: ULABG 3376.

Type locality: “Quebrada la Corcovada,

aproximadamente 8 km de Laguna Mucubaji, en via

Apartaderos-Santo Domingo, 3,000 m snm., 8°48’42”N,

70°47°46”W, Municipio Cardenal Quintero, Estado

Merida, Venezuela.”

Distribution: Region 1.

surroundings of type locality.

Remarks: Material assigned to this name must be

Apparently restricted to

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Catalogue of the amphibians of Venezuela

to: César

Fe, et eS Ti oa rey Sa aie gd ee? pale gn

251. Pristimantis ginesi. Paramo Mucubaji, Mérida. Pho

Barrio-Amoros.

* 4,

252B. Pristimantis gryllus. Male. Matamula, Tachira. Photo:

César Barrio-Amoros.

‘ . os a

We : —_ i ] ns cats

254. Pristimantis hoogmoedi. Female, paratype CVULA 7434.

Las Melenas, southern slope of Cerro El Humo, Peninsula de

Paria, Sucre. Photo: César Barrio-Amoros.

compared with Pristimantis paramerus, a very similar

microsympatric species, as original description makes

no mention of any possible similarity. In Pristimantis

unistrigatus species group according to Hedges et al.

(2008); not assigned to any species group by Padial et

al. (2014).

Selected references: La Marca (2005).

Pristimantis jamescameroni Kok, 2013*

Holotype: IRSNB 4160.

Amphib. Reptile Conserv.

252A. Pristimantis gryllus. Female holotype CVULA 8343. Road from

Estanques to Paramo La Tosta, Mérida. Photo: César Barrio-Amoros.

253. Pristimantis guaiquinimensis. Holotype SMNS 8004.1,

dorsal view. Guaiquinima-tepui, Bolivar. Photo: Dennis Rédder.

oy ie, ee eS. .

= rt a 2 Wie

255. Pristimantis imthurmi. Ptari tepu

Kok.

1, Bolivar. Photo: Philippe

Type locality: Summit of Aprada-tepui, Bolivar State,

5°24’42”N, 62°27°00”W, 2,570 m elevation.

Distribution: Region 5. Endemic to Aprada, a tepui in

Bolivar State, Venezuela.

Remarks: Not assigned to any species group (Padial et

al. 2014).

Selected references: Kok (2013); Sefiaris et al. (2014).

Pristimantis lancinii (Donoso-Barros 1965)*

Holotype: USNM 165604.

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Barrio-Amoros et al.

Type locality: “Apartaderos, Estado Mérida,” Venezuela.

Distribution: Region 1. Pdramos in Sierra Nevada,

Merida State.

Remarks: Described briefly by Donoso-Barros (1965)

stating type locality as: “Venezuela ... de las altas

montafias de Meérida;” type locality restricted later by

Donoso-Barros (1968). Frost (2019) erroneously states

the species was originally described as Eupsophus (see

Barrio-Amoros and Ortiz 2015). Not assigned to any

species group by Hedges et al. (2008) or Padial et al.

(2014).

Selected references: Donoso-Barros (1965, 1968);

Rivero (1967c, 1982b); Rivero and Mayorga (1973);

Lynch (1976); Duellman (1979b); La Marca and Smith

(1982); La Marca (1991b “1994,” 2007); Barrio-Amoros

et al. (2013); Barrio-Amoros and Ortiz (2015).

Pristimantis lassoalcalai

Barrio-Amoros, Rojas-Runjaic and Barros, 2010*

Holotype: MHNLS 18898.

Type locality: “Cerro Las Antenas, elevation 1,827

m, 10°19°40.0°N, 72°35’27.0”°W, Sierra de Perija,

Municipio Rosario de Perija, Estado Zulia, Venezuela.”

Distribution: Region |. Known only from type locality

on Venezuelan side of Sierra de Perija, at 1,827—1,950 m

asl. Probably also present on Colombian side of Sierra

de Periya.

Remarks: Considered Vulnerable (VU) according

to Venezuelan Red Book (Rojas-Runjaic and Sefiaris

2015g). In Pristimantis unistrigatus species group

according to original description (Barrio-Amoros et al.

2010), but not assigned to any species group by Padial

et al. (2014).

Selected references: Barrio-Amoros et al. (20101);

Rojas-Runyaic and Sefiaris (2015g).

Pristimantis longicorpus Kaiser, Barrio-Amor6os,

Rivas-Fuenmayor, Steilein and Schmidt, 2015*

Holotype: CVULA 7444.

Type locality: “Southern slopes of Cerro Humo

(10.7073°N, 62.6284°W), elevation 750 m, Estado

Sucre, Venezuela.”

Distribution: Region 2. Known only from type locality

in Peninsula de Paria, northeastern Venezuela.

Remarks: In Pristimantis unistrigatus species group

according to original description (Kaiser et al. 2015).

Selected references: Kaiser et al. (2015).

Pristimantis marahuaka

(Fuentes and Barrio-Amoros 2004) *

Holotype: MBUCV 6637.

Type locality: Summit of Cerro Marahuaka (03°55’N,

65°27’W), ca. 2,450 m asl, Amazonas State, Venezuela.

Distribution: Region 5. Known only from type locality

Amphib. Reptile Conserv.

in Venezuelan Pantepui.

Remarks: In Pristimantis unistrigatus species group

according to original description (Fuentes and Barrio-

Amoros 2004) and Hedges et al. (2008), but not assigned

to any species group by Padial et al. (2014).

Selected references: Fuentes and Barrio-Amoros (2004).

Pristimantis marmoratus (Boulenger 1900)

Holotype: BMNH 1947.2.16.92.

Type locality: “Foot of Mt. Roraima, 3,500 feet (1,066

m),” Guyana.

Distribution: Region 5. Lowlands and uplands in

Guayana Shield in Venezuela, Guyana, Suriname, and

French Guiana. In Venezuela, restricted to Gran Sabana

in eastern Bolivar State, 600—1,800 m asl.

Remarks: Type locality (same as for Oreophrynella

macconnelli, see its account) 1s not clear and subject

of debate. Kok et al. (2018) suggest it is in Venezuela,

contrary to the opinion herein (see Remarks on O.

macconnelli), and provide a thorough morphological

description, vocalization, natural history, and

phylogenetic relationships. Probably a species complex.

In Pristimantis unistrigatus species group according

to Hedges et al. (2008) and Kok et al. (2018), but not

assigned to any species group by Padial et al. (2014).

Selected references: Boulenger (1900); Ginés (1959);

Rivero (1961, 1964b); Lynch (1976, 1980); Sefiaris et al.

(2014); Kok et al. (2018).

Pristimantis melanoproctus (Rivero 1984)

Holotype: UPR-M 4407.

Type locality: From km 15 Delicias a Rubio, elevation

1,800 m, Tachira State, Venezuela.

Distribution: Region 1. Venezuela and Colombia. In

Venezuela, known only from Andean cloud forests in

Tachira State.

Remarks: Not easily differentiated from Pristimantis

mondolfii, and original author was unable to discriminate

between them unequivocally. Acevedo et al. (2014)

provided new insights to differentiate them. Not assigned

to any species group by Hedges et al. (2008) or Padial

et al. (2014). By its close resemblance to Zachiramantis

lentiginosus, might be a member of genus Zachiramantis

(Heinicke et al. 2015).

Selected references: Rivero (1982c); Acevedo et al.

(2014).

Pristimantis memorans (Myers and Donnelly 1997)

Holotype: AMNH 131466.

Type locality: “Ridge N Pico Tamacuari, 1,270 m

elevation, Sierra Tapirapecd, Amazonas, Venezuela

(1°13’N, 64°42’W).”

Distribution: Region 5. Known from type locality and

one additional locality, a distance of 350 m on Brazilian

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Catalogue of the amphibians of Venezuela

256A. Pristimantis incertus. Holotype of Hylodes incertus,

Original drawing in Lutz (1927).

256B. Pristimantis incertus. Dorsal view of the holotype of

Fylodes incertus, AL-MNRJ 1190.

256C. Pristimantis incertus. Left hand of the holotype of Hylodes

incertus, AL-MNRJ 1190.

256E. Pristimantis incertus. Panemedl pase? reat Zapatero,

Yaracuy. Photo: Sebastian Lotzkat.

side of Tapirapeco (Caramaschi and Niemeyer 2005c).

Remarks: In Pristimantis unistrigatus species group

according to original publication (Myers and Donnelly

1997) and by Hedges et al. (2008), but not assigned to

any species group by Padial et al. (2014).

Selected references: Myers and Donnelly (1997);

Caramaschi and Niemeyer (2005d).

Pristimantis mondolfii (Rivero 1984)

Holotype: UPR-M 6082.

Amphib. Reptile Conserv.

256D. Pristimantis incertus. Male. Plain ohage Cerro El Volcan,

Miranda. Photo: César Barrio-Amoros.

= si ee ee ea ms

256F, USE incertus. Stripped phase. Cerro “Zapatero,

Yaracuy. Photo: Sebastian Lotzkat.

Type locality: Matamula, Municipio Delicias, elevation

1,120 m, Tachira State, Venezuela.

Distribution: Region 1. Colombia and Venezuela. In

Venezuela, known only from the Andes of Tachira State.

Remarks: Acevedo et al. (2014) found it in Colombia and

provided new morphological traits to differentiate it from

Pristimantis melanoproctus. In Pristimantis unistrigatus

Species group according to original publication (Rivero

1984) and Hedges et al. (2008), but not assigned to

any species group by Padial et al. (2014). By its close

resemblance to Tachiramantis lentiginosus, might be a

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Barrio-Amoros et al.

member of genus Zachiramantis (Heinicke et al. 2015).

Selected references: Rivero (1982c); Acevedo et al.

(2014).

Pristimantis muchimuk Barrio-Amoros, Mesa, Brewer-

Carias and McDiarmid, 2010*

Holotype: MHNLS 19652.

Type locality: Northern face of Churi-tepui, 5°16’45”N,

62°00’56”W, 2,325 m asl, Bolivar State, Venezuela.

Distribution: Region 5. Apparently endemic on Churi-

tepui since not observed on nearby tepuis such as

Abakapa (Rojas-Runyjaic et al. 2013).

Remarks: Described by Barrio-Amoros et al. (2010f)

from a single specimen. Variation and an augmented

diagnosis given by Rojas-Runyaic et al. (2013). Not

assigned to any species group by Padial et al. (2014).

Selected references: McDiarmid and Donnelly (2005);

Barrio-Amoros et al. (2010f); Rojas-Runjaic et al.

(2013); Sefiaris et al. (2014).

Pristimantis nicefori (Cochran and Goin 1970)

Holotype: USNM 147012.

Type locality: “Paramo del Almorzadero, Santander,

Colombia.”

Distribution: Region 1. Colombian and Venezuelan

Andes. In Venezuela, known only from Paramo de Tama,

Tachira State.

Remarks: Lynch (1994) rediagnosed it and related it

to Phrynopus sensu lato. In Pristimantis unistrigatus

species group according to Lynch and Duellman (1997)

and Hedges et al. (2008), but not assigned to any species

group by Padial et al. (2014).

Selected references: Lynch (1981, 1994); La Marca

Geo):

Pristimantis nubisilva Kaiser, Barrio-Amoros, Rivas-

Fuenmayor, Steilein and Schmidt, 2015*

Holotype: CVULA 7430.

Type locality: “Southern slopes of Cerro Humo,

Peninsula de Paria, Estado Sucre, Venezuela (10.7073°N,

62.6284°W), elevation ca. 750 m.”

Distribution: Region 2. Endemic to Peninsula de Paria

in northeastern Venezuela.

Remarks: In Pristimantis unistrigatus species group

according to original publication (Kaiser et al. 2015).

Selected references: Kaiser et al. (2015).

Pristimantis paramerus (Rivero 1984)*

Holotype: UPR-M 2926.

Type locality: “Paramo de Santo Domingo, region de

Mucuchies, Edo. Mérida, Venezuela, 3,330 m.”

Distribution: Region 1. Restricted to paramos in Sierra

Amphib. Reptile Conserv.

Nevada (Mucubaji, Sai Sai, Los Granates), Mérida State.

Remarks: In Pristimantis unistrigatus species group

according to original publication (Rivero 1984), but not

assigned to any species group by Hedges et al. (2008) or

Padial et al. (2014).

Selected references: Rivero (1982b); La Marca (2005).

Pristimantis pariagnomus Kaiser, Barrio-Amoros,

Rivas-Fuenmayor, Steilein and Schmidt, 2015*

Holotype: MHNLS 14456.

Type locality: “Southern slopes of Cerro Humo

(10.7073°N, 62.6284°W), elevation 780 m, Estado

Sucre, Venezuela.”

Distribution: Region 2. Only known from type locality in

Peninsula de Paria, Sucre State, northeastern Venezuela.

Remarks: Described based on a single specimen; no

other specimens known to date as vouchers; however,

individual in photograph (Image 269) is a second

individual of the species, and additional specimens have

been observed (G. Rivas, pers. comm.). In Pristimantis

unistrigatus species group according to original

publication (Kaiser et al. 2015).

Selected references: Kaiser et al. (2015).

Pristimantis pedimontanus (La Marca 2004)*

Holotype: ULABG 3221.

Type locality: Parque Nacional Chorro del Indio, 16.5

km in road San Cristobal-Macanilla, approx. 8 km SE of

San Cristobal, San Cristobal municipality, Tachira State,

Venezuela.

Distribution: Region 1. Eastern slopes of Cordillera de

Merida, Andes of Venezuela.

Remarks: Assigned to Pristimantis conspicillatus

Species group in original publication (La Marca 2004)

and maintained in this group by Hedges et al. (2008).

More recently, removed from P. conspicillatus species

group and left unassigned to any species group by Padial

et al. (2014).

Selected references: La Marca (2004).

Pristimantis pleurostriatus (Rivero 1984)*

Holotype: UPR-M 4971.

Type locality: San Eusebio, La Carbonera, 2,316 m,

Merida State, Venezuela.

Distribution: Region 1. Endemic to

Cordillera de Mérida. Apparently

surroundings of type locality.

Remarks: Barrio-Amoros et al. (2013) redescribed

this previously poorly-known species using fresh

material and placed it into a phylogenetic context. In

Pristimantis unistrigatus species group according to

original publication (Rivero 1984), but not assigned to

any species group by Hedges et al. (2008) or Padial et

Venezuelan

restricted to

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Catalogue of the amphibians of Venezuela

ee ee ae

*, eae

257. Pristimantis jamescameroni. Aprada tepui, Bolivar. Photo:

Philippe Kok.

259. Pristimantis lassoalcalai. Female paratype MHNLS 18877.

Cerro Las Antenas, Sierra de Perija, Zulia. Photo: Pablo Velozo.

. * 7s 4

261. Pristimantis marahuaka. Cerro Marahuaka (tepui) Summit,

Amazonas. Photo: Charles Brewer-Carias.

al. (2014).

Selected references: Rivero (1982c, 1983); Barrio-

Amoros et al. (2013).

Pristimantis pruinatus (Myers and Donnelly 1996)*

Holotype: EBRG 3006.

Type locality: “Summit of cerro Yavi, 2,150 m,

Amazonas, Venezuela.”

Distribution: Region 5. Endemic to Cerro Yavi, a tepui

in northern Amazonas State.

Remarks: In subgenus Pristimantis but not assigned

Amphib. Reptile Conserv.

260. Pristimantis longicorpus. Male holotype CVULA 7444.

Southern slopes of Cerro Humo, Peninsula de Paria, Sucre,

Venezuela. Photo: César Barrio-Amoros.

]

262A. Pristimantis aff. marmoratus. El Pefion, Auyan-tepul,

Bolivar. Photo: César Barrio-Amoros.

to any species group (Hedges et al. 2008; Myers and

Donnelly 1996; Padial et al. 2014).

Selected references: Myers and Donnelly (1996).

Pristimantis pulvinatus (Rivero 1968)*

Holotype: MCZ 64741.

Type locality: “Paso del Danto, region of La Escalera

around 1,400 m above San Isidro, road from El] Dorado

to Sta. Elena de Uairén, Estado Bolivar, Venezuela.”

Distribution: Region 5. Venezuela and Guyana. In

Venezuela, in surroundings of La Escalera and Gran

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Barrio-Amoros et al.

262B. Pristimantis marmoratus. Chivaton, Gran Sabana, Bolivar.

Photo: César Barrio-Amoros.

264A. Pristimantis cf. mondolfii. Matamula, Tachira. Photo:

César Barrio-Amoros.

265A. Pristimantis muchimuk. Female holotype MHNLS 19652.

Churi-tepui, Chimanta massif, Bolivar. Photo: Javier Mesa.

Sabana, eastern Bolivar State. Has been confused with P.

marmoratus (Kok et al. 2018).

Remarks: In subgenus Pristimantis but not assigned

to any species group (Hedges et al. 2008; Padial et al.

2014). Probably a species complex (P. Kok and CBA, in

prep.).

Selected references: Rivero (1968c); Lynch (1976);

Duellman (1997); Myers and Donnelly (1997); Gorzula

and Sefiaris (1998); Barrio-Amords and Duellman

(2009); Sefiaris et al. (2014); Kok et al. (2018).

Amphib. Reptile Conserv.

263. Pristimantis melanoproctus. Tama National Park, Norte de

Santander, Colombia. Photo: Aldemar A. Acevedo.

264B. Pristimantis mondolfii. Tama National Park, Norte de

Santander, Colombia. Photo: Aldemar. A. Acevedo.

a ae

7 ~ . a - .

‘ ~~. |

, as = . |

265B. Pristimantis muchimuk. Churi-tepui, Chimanta massif,

Bolivar. Photo: Fernando Rojas-Runjaic.

Pristimantis reticulatus (Walker and Test 1955)*

Holotype: UMMZ 109872.

Type locality: “Slope of Pico Periquito +/- 1,275 m,

Rancho Grande, Estado Aragua, Venezuela.”

Distribution: Only known from type locality and

surroundings at elevations over 1,000 m in central

Cordillera de la Costa of Aragua State.

Remarks: Region 2. Apparently, specimens last collected

in 1971. In subgenus Pristimantis but not assigned to any

species group (Hedges et al. 2008; Padial et al. 2014).

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Catalogue of the amphibians of Venezuela

266. Pristimantis nicefori. Paramo Tama, Tama National Park,

Norte de Santander, Colombia. Photo: Aldemar A. Acevedo.

. 8 et a eee

268A. Pristimantis paramerus. Quebrada del Padre, way to

paramo Los Granates, Mérida. Photo: César Barrio-Amoros.

: = ir

267. Pristimantis nubisilva. Amplexus. Southern slopes of Cerro

Humo, Peninsula de Paria, Sucre. Photo: César Barrio-Amoros.

he]

268B. Pristimantis paramerus. Quebrada del Padre, way to

paramo Los Granates, Mérida. Photo: César Barrio-Amoros.

269. Pristimantis pariagnomus. Cerro el Humo, Peninsula de

Paria, Sucre. Photo: Mayke de Freitas.

Selected references: Walker and Test (1955); Ginés

(1959); Rivero (1961, 1964a); Lynch (1976); Lynch and

La Marca (1993); Barrio-Amoros (2006c).

Pristimantis rhigophilus (La Marca 2007)*

Holotype: ULABG 1915.

Type locality: “Paramo de Guaramacal, 3,100 m

(9°13’53.9"N, 70°11°8.7’W), en la via Laguna de los

Cedros-Guaramacal, Municipio Bocono, Estado Trujillo,

Venezuela.”

Amphib. Reptile Conserv.

270A. Pristimantis pleurostriatus. Calling male. Estancia La

Bravera, way from Mérida to La Azulita, 2,200 m asl, Mérida.

Photo: César Barrio-Amoros.

Distribution: Known only from type locality in

Venezuelan Andes.

Remarks: In subgenus Pristimantis but not assigned to

any species group (Hedges et al. 2008; Padial et al. 2014).

Selected references: La Marca (2007).

Pristimantis rivasi

Barrio-Amoros, Rojas-Runjaic and Barros, 2010

Holotype: MHNLS 18445.

Type locality: “Cerro Las Antenas, elevation 1,670 m,

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Barrio-Amoros et al.

10°20°N, 72°35’ W, Sierra de Perija, Municipio Rosario

de Perija, Estado Zulia, Venezuela.”

Distribution: Region 1. Known only from eastern

(Venezuelan) versant of Sierra de Perija, in cloud forest

at 1,438—1,933 m asl. Recently reported from Colombian

side of Perija (Meza-Joya 2016).

Remarks: In Pristimantis unistrigatus species group

according to original publication (Barrio-Amoros et al.

2010), but not assigned to any species group by Padial

et al. (2014).

Selected references: Barrio-Amoroés et al. (20101);

Meza-Joya (2016).

Pristimantis riveroi (Lynch and La Marca 1993)*

Holotype: AMNH 70599.

Type locality: “Rancho Grande,

Venezuela.”

Distribution: Region 2. Known only from type locality

in central Cordillera de la Costa in Aragua State.

Remarks: In Pristimantis unistrigatus species group

according to Hedges et al. (2008), but not assigned to

any species group by Padial et al. (2014).

Selected references: Lynch and La Marca (1993);

Barrio-Amoros (2006c).

Estado Aragua,

Pristimantis rozei (Rivero 1961)*

Holotype: MBUCV 2018.

Type locality: “Curucuruma, Estado Aragua, Venezuela.”

Distribution: Region 2. Known only from surroundings

of type locality in central Cordillera de la Costa, Aragua

State.

Remarks: Perhaps conspecific with Pristimantis

reticulatus, according to Lynch and La Marca (1993). In

P. unistrigatus species group according to Hedges et al.

(2008), but not assigned to any species group by Padial

et al. (2014). Reported P. rozei from Nirgua Massif in

Yaracuy State by Lotzkat (2007) considered here to be P.

cf. stenodiscus, due to characteristic shape of finger disks

(Image 274).

Selected references: Rivero (1961, 1964a); Lynch

(1976, 1979b); La Marca (1992); Lynch and La Marca

(1993); Lotzkat (2007).

Pristimantis sarisarinama

Barrio-Amoros and Brewer-Carias, 2008*

Holotype: EBRG 4668.

Type locality: “Sima Mayor, Sarisarifiama-tepui, Estado

Bolivar, Venezuela (4°41’N, 64°13’W), elevation 1,100

Distribution: Region 5. Apparently endemic to

Sarisarifiama, a tepui in Bolivar State, southern

Venezuela.

Remarks: In Pristimantis unistrigatus species group

Amphib. Reptile Conserv.

according to original publication (Barrio-Amoros and

Brewer-Carias 2008), but not assigned to any species

group by Padial et al. (2014).

Selected references: Barrio-Amoros and Brewer-Carias

(2008).

Pristimantis stenodiscus (Walker and Test 1955)*

Holotype: UMMZ 109866.

Type locality: “Pico Periquito, Rancho Grande, Estado

Aragua, Venezuela.”

Distribution: Region 2. Only known from vicinity of

type locality in central Cordillera de la Costa. Records

from Nirgua Massif, Yaracuy State, must be confirmed.

Remarks: There is no recent information about the

species. In subgenus Pristimantis but not assigned to any

species group (Hedges et al. 2008; Padial et al. 2014).

Sebastian Lotzkat sent photos taken in Nirgua Massif,

Yaracuy (Lotzkat 2007), and the reported P. rozei in that

thesis is herein considered to be P. cf. stenodiscus, due

to characteristic shape of finger disks (Image 276), but

should be better confirmed with voucher specimens.

Selected references: Walker and Test (1955); Ginés

(1959); Rivero (1961); Lynch (1976); Barrio-Amor6és

(2006c); Lotzkat (2007).

Pristimantis telefericus (La Marca 2005)*

Holotype: ULABG 2232.

Type locality: From path between third and fourth

stations of Teleférico de Mérida, 3,500 m, Libertador

Municipality, Mérida State, Venezuela.

Distribution: Region 1. Apparently microendemic

around type locality in Sierra Nevada de Mérida, in

paramo habitat at 3,400—3,500 m asl.

Remarks: Appears to have suffered a decline, as intensive

search by CBA in March-April 2012 (nearly 20 hours)

resulted in no positive observations. In Pristimantis

unistrigatus species group according to Hedges et al.

(2008) but not assigned to any species group by Padial

et al. (2014).

References: La Marca (2005).

Pristimantis thyellus (La Marca 2007)*

Holotype: ULABG 2137.

Type locality: “Paramo El Batallon, 3,125 m (aprox.

8°10’°48"N, 71°54’08”W), Parque Nacional Juan Pablo

Pefialoza, Municipio Guaraque, Estado Mérida, cerca del

limite con el Estado Tachira, Venezuela.”

Distribution: Region 1. Apparently restricted to paramo

El Batallon, 2,900-—3,800 m asl. Expected in Tachira

State due to continuous habitat.

Remarks: In subgenus Pristimantis but not assigned to

any species group (Hedges et al. 2008; Padial et al. 2014).

Selected references: La Marca (2007).

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270B. Pristimantis pleurostriatus. Juvenile. Estancia La Bravera,

way from Mérida to La Azulita, 2,200 m asl, Mérida. Photo: César

Barrio-Amoros.

271B. Pristimantis cf. pulvinatus. Male. La Laja, Sierra de Lema,

Bolivar. Photo: Alan Highton.

272A. Pristimantis rivasi. Female holotype MHNLS 18445. Cerro

Las Antenas, Sierra de Perija, Zulia. Photo: Tito Barros.

Pristimantis tubernasus (Rivero 1984)*

Holotype: UPR-M 4349.

Type locality: “Boca de Monte, Camino a Pregonero,

2,393 m, Edo. Mérida, Venezuela.”

Distribution: Region 1. Colombia and Venezuela. In

Venezuela, distributed in the Andes of Tachira, Barinas,

and Merida States.

Remarks: Eleutherodactylus pulidoi Rivero, 1984 was

considered a junior synonym of P. tubernasus by Lynch

(2003). In P. unistrigatus species group of Hedges et al.

(2008) but not assigned to any species group by Padial

et al. (2014).

Amphib. Reptile Conserv.

271A. Pristimantis pulvinatus. Paso del Danto, La Escalera region,

Bolivar. Photo: César Barrio-Amoros.

272B. Pristimantis rivasi. Male. Cerro Las Antenas, Sierra de

Perija, Zulia. Photo: Fernando Rojas-Runjaic.

Selected references: Rivero (1982c); Ruiz-Carranza et

al. (1996); Lynch (2003).

Pristimantis turik

Barrio-Amoros, Rojas-Runjaic and Infante, 2007*

Holotype: MBLUZ 155.

Type locality: Cueva del Agua (lower gallery in the

Northern Wall cave), Mesa Turik, Rosario de Perija

Municipality, Sierra de Perija, Zulia State, Venezuela

(10°24’N, 72°42’W; 1,700 m asl).

Distribution: Region 1. Only known from type locality;

could be present on Colombian side of Sierra de Periya.

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Barrio-Amoros et al.

273A. Pristimantis riveroi. Male. Nirgua Massif, Yaracuy. Photo:

Sebastian Lotzkat.

Lotzkat.

5S dae ec ea we eee Bowe te |

276. Pristimantis cf. stenodiscus. Nirgua Massif, Yaracuy. Photo:

Sebastian Lotzkat.

ot "

| ia

278. Pristimantis tubernasus. San Isidro, Barinas. Photo: César

Barrio-Amoros.

Amphib. Reptile Conserv.

re % a a

273B. Pristimantis riveroi. Amplexus. Rancho Grande, Henri

Pittier National Park, Aragua. Photo: Andrés Gonzalez.

21S; Pristimantis sarisarinama. Paratopotype EBRG 4674. Sima

Mayor Sarisarifiama-tepui, Bolivar. Photo: César Barrio-Amoros.

5 cc

277. Pristimantis thyellus. Paramo Batallon y la Negra, Tachira.

Photo: Liz del Valle.

on. a ge

mea fF ao

279A. Pristimantis vanadisae. Female, pattern plain. Estancia La

Bravera, way from Mérida to La Azulita, 2,200 m asl, Mérida.

Photo: César Barrio-Amoros.

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Catalogue of the amphibians of Venezuela

~ im,

‘i | | 3 +

279B. Pristimantis vanadisae. Pattern plain. Estancia La Bravera,

way from Mérida to La Azulita, 2,200 m asl, Mérida. Photo: César

Barrio-Amoros.

wv f a

279D. Pristimantis vanadisae. Juvenile, spotted pattern. Estancia

La Bravera, way from Mérida to La Azulita, 2,200 m asl, Mérida.

Photo: César Barrio-Amoros.

280B. Pristimantis vilarsi. Female. Tobogan de la Selva, near

Puerto Ayacucho, Amazonas. Photo: Zelimir Cernelic.

Remarks: Known only from single specimen. In

Pristimantis unistrigatus species group according to

original publication (Barrio-Amoros et al. 2007) but not

assigned to any species group by Padial et al. (2014).

Selected references: Barrio-Amoros et al. (2007).

Pristimantis turumiquirensis (Rivero 1961)*

Holotype: AMNH 22557.

Type locality: La Trinidad, Mount Turumiquire, in cave

at approximate elevation of 1,830 m, between Sucre and

Monagas States, Venezuela.

Distribution: Region 2. Apparently endemic to

Amphib. Reptile Conserv.

279C. Pristimantis vanadisae. Female, pattern dorsoconcolor.

Estancia La Bravera, way from Mérida to La Azulita, 2,200 m asl,

Mérida. Photo: César Barrio-Amoros.

3 ‘t “s i

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“4 ts

2 Yl ‘

y ie ;

ja ek : i

280A. Pasamanhs

vilarsi. Male. Tobogan de la Selva, near Puerto

Ayacucho, Amazonas. Photo: César Barrio-Amoros.

281. Pristimantis yukpa. Female. Cerro Las Antenas. Photo:

Fernando Rojas-Runjaic.

Turimiquire, a massif in eastern part of Coastal Range,

northwestern Venezuela.

Remarks: Considered Endangered (EN) by Venezuelan

Red Book (Sefiaris and Rojas-Runjaic 2015d). In

Pristimantis unistrigatus species group of Hedges et al.

(2008) but not assigned to any species group by Padial

et al. (2014).

Selected references: Rivero (1961, 1964a); La Marca

(1992); Sefiaris and Rojas-Runyaic (2015d).

Pristimantis vanadise (La Marca 1984)*

Holotype: CVULA 2805.

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Barrio-Amoros et al.

Type locality: “Stream in cloud forest above

Truchicultura La Mucuy, 2,350 m, Sierra Nevada de

Merida, Estado Mérida, Venezuela.”

Distribution: Region 1. In Andes of Mérida State.

Remarks: Barrio-Amoros et al. (2013) described four

chromotypes, placed it in a phylogenetic context, and

made an unjustified amendment, spelling species name

as vanadisae. In Pristimantis unistrigatus species group

according to original publication (La Marca 1984) and

Hedges et al. (2008); not assigned to any species group

by Padial et al. (2014).

Selected references: Rivero (1982c); La Marca (1984b,

1985a, 1991b “1994”): Barrio-Amoros et al. (2013).

Pristimantis vilarsi (Melin 1941)

Syntypes: NHMG 491 (two specimens).

Type locality: Taracua, Rio Uaupés, (Amazonas State),

Brazil.

Distribution: Regions 4, 5. Upper Amazonian Region

of Brazil, Peru, Colombia, and Venezuela. In Venezuela,

widespread south of Orinoco River.

Remarks: A redescription of the species and map of

its distribution appeared in Barrio-Amoroés and Molina

(2006). Heyer and Barrio-Amoros (2009) described the

call. Eleutherodactylus stegolepis Schliter and Rodder,

2007 is a junior synonym (Kok and Barrio-Amoros

2013). In Pristimantis conspicillatus species group of

Hedges et al. (2008) and Padial et al. (2014).

Selected references: Melin (1941); Rivero (1961, 1964b,

1967a); Lynch (1975a, 1976, 1979b, 1980); Rivero et

al. (1986); La Marca (1992); Barrio (1996a); Gorzula

and Sefiaris (1998); Barrio-Amoros and Molina (2006);

Schliiter and R6dder (2007); Heyer and Barrio-Amoros

(2009); Barrio-Amoros et al. (2011b); Kok and Barrio-

Amoréos (2013).

Pristimantis yaviensis (Myers and Donnelly 1996)*

Holotype: EBRG 3017.

Type locality: “Summit of Cerro Yavi, 2,150 m,

Amazonas, Venezuela.”

Distribution: Region 5. Only known from Cerro Yavi

and Cerro Yutajé, two tepuis in northern Amazonas State.

Remarks: In Pristimantis unistrigatus species group

according to original publication (Myers and Donnelly

1996) and Hedges et al. (2008).

Selected references: Myers and Donnelly (1996, 2001);

Gorzula and Sefiaris (1998).

Pristimantis yukpa

Barrio-Amoros, Rojas-Runjaic and Infante, 2007*

Holotype: MHNLS 18525.

Type locality: Kiriponsa, an indigenous Yukpa

community along the Tokuko River, Machiques de Perija

Amphib. Reptile Conserv.

Municipality, Sierra de Perija, Zulia State, Venezuela

(09°51’°N, 72°53’ W; 1,005 m asl).

Distribution: Region 1. Widely distributed along eastern

(Venezuelan) versant of Sierra de Perija, 600-1,200 m

asl. Recently reported from Colombian side of Perija

(Meza-Joya 2016).

Remarks: Barrio-Amoros et al. (2012) show P. gryllus

as vicariant species of P. yukpa in Cordilleras Oriental

and de Mérida. Both species are members of uwnistrigatus

species group sensu stricto (nested with P. unistrigatus),

Padial et al. (2014) did not assign it to any species group.

Selected references: Barrio-Amordos et al. (2007, 2012);

Meza-Joya (2016).

Pristimantis yuruaniensis Rodder and Jungfer, 2008*

Holotype: ZFMK 87278.

Type locality: Yuruani-tepui, Gran Sabana Municipality,

Bolivar State, Venezuela (5°19°31”N, 60°50’40°W;

elevation 2,300 m.

Distribution: Region 5. Restricted to summit of Yuruani,

a tepui in southeastern Bolivar State.

Remarks: Considered Vulnerable (VU) according

to Venezuelan Red Book (Sefiaris and Rojas-Runjaic

2015b). Not assigned to any species group by Padial et

al. (2014) or Rodder and Jungfer (2008), but considered

in “unistrigatus” species group by Kok et al. (2018).

Selected references: R6dder and Jungfer (2008); Sefiaris

et al. (2014); Sefiaris and Rojas-Runjaic (2015b).

Pristimantis yustizi

(Barrio-Amoros and Chacon 2004)*

Holotype: CVULA 2150.

Type locality: Cerro Alto, La Soledad, Barinas State,

Venezuela, elevation 1,500 m.

Distribution: Region 1. Eastern slopes of Andes of

Tachira, Barinas, Lara, and Portuguesa States.

Remarks: In Pristimantis unistrigatus species group of

Hedges et al. (2008) but not assigned to any species group

by Padial et al. (2014). Most likely actually a member of

P. conspicillatus group (B.S. Hedges and CBA, in prep.).

Selected references: Barrio-Amoros and Chacon 2004;

Barrio-Amoros 2010a.

Pristimantis zeuctotylus (Lynch and Hoogmoed 1977)

Holotype: RMNH 17701.

Type locality: “West slope, Vier Gebroeders Mountain,

Sipaliwini, Nickerie District, Suriname.”

Distribution: Region 4. Widely distributed in Amazonia,

from southern Colombia through Venezuela, Brazil,

Guyana, and Suriname to French Guiana. In Venezuela,

only known from a few localities in southern Amazonas

State.

Remarks: Barrio-Amoros and Molina (2006) doubted

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Catalogue of the amphibians of Venezuela

282. Pristimantis yuruaniensis. Female. Summit of Yuruani-tepul,

Bolivar. Photo: César Barrio-Amoros.

Pie os - = . r “’

284. Pristimantis zeuctotylus. Camp Angouleme, French Guiana.

Photo: P.J.R. Kok.

; a . : ~& : a 3 ve San

286. Pristimantis sp. 2. Sub paramo La Motus, Mérida. Photo:

César Barrio-Amoros.

the validity of this taxon, which is distinguished from

Pristimantis vilarsi only by absence of tarsal fold (present

on P. vilarsi) and rounded palmar tubercle (variable in P.

vilarsi). Araujo de Oliveira et al. (2017) show a maximum

likelihood tree in which P. zeuctotylus from Suriname

is presented as sister to P. vilarsi from Colombia; this

should be confirmed with specimens from type localities

of both species. In Pristimantis conspicillatus species

group of Hedges et al. (2008) and Padial et al. (2014).

Selected references: Lynch and Hoogmoed (1977);

Lynch (1980); Barrio-Amoros and Molina (2006).

Subfamily Strabomantinae

Hedges, Duellman and Heinicke, 2008

Amphib. Reptile Conserv.

283. Pristimantis yustizi. Female. Calderas, Barinas. Photo: César

Barrio-Amoros.

LS ee

e r sr L-. Fee See

285. Pristimantis sp. 1. Paramo Los Granates, Mérida. Photo:

César Barrio-Amoros.

: ee -

Aa f: a os "

he t pre

287. Pristimantis sp. 3. Paramo Pifiango, Mérida. Photo: César

Barrio-Amoros.

Genus Strabomantis Cope, 1862

Type species: Strabomantis biporcatus Peters, 1863, by

monotypy.

Strabomantis biporcatus Peters, 1863*

Lectotype: ZMB 3330B, assigned by Savage and Myers

(2002).

Type locality: “Veragua,’ in error for “northern

Venezuela.”

Distribution: Region 2. Distributed throughout

Cordillera de la Costa (Fig. 1 in Barrio-Amoros and

Kaiser 2008).

Remarks: Long-known as Eleutherodactylus cornutus

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Barrio-Amoros et al.

maussi (Rivero 1961) or Eleutherodactylus maussi (La

Marca 1882; Barrio-Amoros 1998). Reviewed by Savage

and Myers (2002), who determined the name biporcatus

(originally stated from Veragua, Panama) applied to

four specimens collected in Northern Venezuela, then

Eleutherodactylus biporcatus. Crawford and Smith (2005)

transferred E. biporcatus to genus Craugastor. Hedges

et al. (2008) subsequently recovered Strabomantis and

included biporcatus as its type species. Barrio-Amoros

and Kaiser (2008) commented on phenotypic variation

and distribution. In Strabomantis biporcatus species

group of Hedges et al. (2008) and Padial et al. (2014).

Selected references: Boettger (1893); Lutz (1927);

Ginés (1959); Rivero (1961, 1964a, d); Heatwole (1962);

Mertens (1967); Lynch (1975, 1976); La Marca (1992);

Manzanilla et al. (1996); Savage and Myers (2002);

Rivero and Mijares (2004); Barrio-Amoros (2006c);

Barrio-Amoros and Kaiser (2008).

Order Urodela Duméril, 1805

Remarks: Herein follows Dubois and Raffaelli (2012) in

the rationale for using the taxon nomina Urodela Dumeéril,

1805 instead of Caudata Fischer von Waldheim, 1813.

Family Plethodontidae Gray, 1850

Subfamily Hemidactyliinae Hallowel, 1856

Remarks: Herein follows most comprehensive and

symmetric taxonomy by Vieites et al. (2011).

Genus Bolitoglossa Dumeril, Bibron and Dumeril, 1854

Type species: Salamandra platydactylus Gray, 1831, by

monotypy.

Remarks: All members of Venezuelan Bolitoglossa

pertain to subgenus Eladinea Miranda Ribeiro, 1937

(Parra-Olea et al. 2004).

Bolitoglossa borburata Trapido, 1942*

Holotype: USNM 115509.

Type locality: “Valle del Rio Borburata, Estado

Carabobo, Venezuela, 1,200 m.”

Distribution: Region 2. Endemic from Venezuela.

Restricted to Cordillera de la Costa (Carabobo, Aragua,

and Yaracuy States).

Remarks: Probably a complex of species; specimens

from Yaracuy assigned tentatively to Bolitoglossa

borburata by Barrio and Fuentes (1999b) can represent a

different taxon. Unfortunately, seems to have vanished, at

least from central Cordillera de la Costa. In Bolitoglossa

adspersa species group of Parra-Olea et al. (2004).

Selected references: Trapido (1942); Rohl (1959);

Brame and Wake (1963); Hanken and Wake (1982);

Amphib. Reptile Conserv.

Manzanilla et al. (1995); Rodriguez and Rojas-Suarez

(1995); Barrio (1999a); Barrio and Fuentes (1999b),;

Barrio-Amoros (2006c).

Bolitoglossa guaramacalensis

Schargel, Garcia-Pérez and Smith, 2002*

Holotype: MCNG-A 2121.

Type locality: “Bocond-Guaramacal road, Quebrada el

Pollo (9°13’N, 70°10’W), south slope of the Macizo de

Guramacal, 2,400 m, Trujillo, Venezuela.”

Distribution: Region 1. Endemic to Venezuela.

Apparently restricted to Guaramacal massif (Trujillo

State), in eastern versant of Venezuelan Andes.

Remarks: In Bolitoglossa adspersa species group

according to original publication (Schargel et al. 2002)

and by Parra-Olea et al. (2004).

Selected references: Schargel et al. (2002).

Bolitoglossa leandrae Acevedo, Wake, Marquez, Silva,

Franco and Amézquita, 2013

Holotype: MCNUP 62.

Type locality: “San Antonio, Parque Nacional Natural

Tama (PNNT) (7.153092, -72.227306) 600 m elevation,

Departamento de Norte de Santander, Colombia.”

Distribution: Region 1. Endemic from foothills of Tama

massif between Colombia and Venezuela. Reported from

Venezuelan side by Barrio-Amoros et al. (2015).

Selected references: Acevedo et al. (2013); Barrio-

Amoros et al. (2015).

Bolitoglossa mucuyensis Garcia-Gutiérrez, Escalona,

Mora, Diaz de Pascual and Fermin, 2013*

Holotype: ULABG 1772.

Type locality: “Parque Nacional Sierra Nevada, La

Mucuy, 2,225 m, Santos Marquina municipality, Mérida

State, Venezuela.”

Distribution: Region 1. Endemic from Venezuela. Only

known from type locality, La Mucuy, 1,800-2,250 m.

Probably more extended through cloud forests of western

versant of Sierra Nevada de Merida.

Selected references: Garcia-Gutiérrez et al. (2013).

Bolitoglossa orestes Brame and Wake, 1962*

Holotype: BMNH 1905.5.31.103.

Type locality: “Culata, 9,810 feet (3,000 meters)

Cordillera de Mérida, Estado de Mérida, Venezuela.”

9,810 feet corresponds to 2,990 m.

Distribution: Region 1. Sierra de la Culata in Mérida

State, Venezuela. Type locality and distribution referred

to Bolitoglossa spongai is summed for B. orestes. Locality

“La Mucuy” in Sierra Nevada de Merida, given in Barrio-

Amoros and Fuentes Ramos (1999: 16) for B. spongai

was an editorial error; currently B. orestes is only known

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Catalogue of the amphibians of Venezuela

288A. Strabomantis biporcatus: Female. N

Photo: Sebastian Lotzkat.

+ a » tia ~s a Doni + ?

| AS \9

a, “eo

. ee

oe eentae

289. Bolitoglossa cf. altamazonica. Doradas river valley. Photo:

César Barrio-Amoros.

aR J aj - “¥

Y et I a

“ a if ' 1 | ue be i

291. Bolitoglossa guaramacalensis. Guaramacal National Park.

Photo: Juan E. Garcia-Pérez.

from Sierra de la Culata, separated from B. mucuyensis

by the deep valley of Chama River. Populations from

La Mucuy later described as B. mucuyensis (Garcia-

Gutiérrez et al. 2013).

Remarks: Fermin et al. (2012) recently passed

Bolitoglossa spongai Barrio-Amordos and Fuentes-

Ramos, 1999 to synonymy of B. orestes, based on genetic

and morphological data. Therefore, all data in literature

for B. spongai (Rodriguez and Royas-Suarez 2008;

Barrio-Amoros et al. 2010e) refers now to B. orestes.

Amphib. Reptile Conserv.

. fe 3

S baa,

Humo, Peninsula de Paria, Sucre. Photo: Hinrich Kaiser.

290. Bolitoglossa borburata. Nirgua Massif, Yaracuy. Photo:

Sebastian Lotzkat.

wil *

292: Bolitoglossa leandrae. Female. Rio Frio, Tachira. Photo:

César Barrio-Amoros.

Two works on natural history and ecological traits of B.

orestes: Cadenas et al. (2009) based on a population at

Monte Zerpa, north of Mérida, and Barrio-Amoros et al.

(2010e) under name B. spongai, about a population from

cloud forests around Paramo El Tambor, Mérida State.

The Venezuelan Red Book considers it Vulnerable (VU)

[Barrio-Amoros and Rivas 2015].

Selected references: Brame and Wake (1962, 1963);

Péfaur and Diaz de Pascual (1982); Rodriguez and

Rojas-Suarez (1995, 2008); Barrio-Amords (1998,

2004); Barrio-Amoros and Fuentes-Ramos (1999, 2004);

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Barrio-Amoros et al.

Cadenas et al. (2009); Barrio-Amoros et al. (2010e);

Fermin et al. (2012); Barrio-Amoros and Rivas (2015).

Bolitoglossa tamaense Acevedo, Wake, Marquez, Silva,

Franco and Amézquita, 2013

Holotype: MCNUP 50.

Terra typica: “La Asiria de Belén, Parque Nacional

Natural Tama (PNNT) (7.319278, -72.374778) at 2,700

m elevation, Departamento de Norte de Santander,

Colombia.”

Distribution: Region 1. Endemic from cloud forest of

Tama massif between Colombia and Venezuela. Reported

from Venezuelan side by Barrio-Amoros et al. (2015).

Selected references: Acevedo et al. (2013); Barrio-

Amoros et al. (2015).

Order GYMNOPHIONA Miller, 1832

Family Caeciliidae Rafinesque, 1814

Genus Caecilia Linnaeus, 1758

Type species: Caecilia tentaculata Linnaeus, 1758, by

subsequent designation of Dunn, 1942.

Caecilia flavopunctata Roze and Solano, 1963*

Holotype: MBUCV 5358.

Type locality: “Albarico, Yaracuy, Venezuela.”

Distribution: Region 2. Endemic to Venezuela. Known

only from surroundings of type locality.

Selected references: Roze and Solano (1963); Taylor

(1968); Nussbaum and Wilkinson (1989); Rivas (2009).

Caecilia subnigricans Dunn, 1942

Holotype: ANSP 4921 (not 4821 as given in description,

sensu Taylor, 1968).

Type locality: “Magdalena River, Colombia.”

Distribution: Regions 1, 2. Magdalena Valley in

Colombia, Andes of Venezuela (Tachira and Barinas),

and Falcon State.

Selected references: Dunn (1942); Ginés (1959); Taylor

(1968); Péfaur and Diaz de Pascual (1987); Nussbaum

and Wilkinson (1989); Péfaur et al. (1992); Lynch (2000).

Caecilia tentaculata Linnaeus, 1758

Type: NHRM (lost).

Type locality: “America.”

Distribution: Regions 1, 2, 4. South America east of

Andes, in Colombia, Venezuela, Guyana, Suriname,

French Guiana, Brazil, Ecuador, and Peru. Apparently

widespread in lowland Venezuela, known from Estados

Amazonas, Tachira, and Yaracuy (Barrio-Amoros 1998).

Selected references: Dunn (1942); Taylor (1968); Péfaur

Amphib. Reptile Conserv.

et al. (1987, 1992); Nussbaum and Wilkinson (1989);

Maciel and Hoogmoed (2011).

Family Rhinatrematidae Nussbaum, 1977

Genus Epicrionops Boulenger, 1883

Type species: Epicrionops bicolor Boulenger, 1883, by

monotypy.

Epicrionops niger (Dunn 1942)

Neotype: MBUCV 5360.

Type locality: “Arundabara, British Guiana, elevation

2,200 feet;” locality of neotype is “El Dorado, Bolivar,

Venezuela.”

Distribution: Region 5. South-central Guayana and

Southeastern Venezuela. In Venezuela, a few localities in

Bolivar State (Barrio-Amoros 1998).

Selected references: Dunn (1942); Taylor (1968);

Nussbaum and Wilkinson (1989); Péfaur et al. (1992);

Sefiaris et al. (2014).

Family Shiphonopidae Bonaparte, 1850

Genus Microcaecilia Taylor, 1968

Type species: Dermophis albiceps Boulenger, 1882, by

original designation.

Microcaecilia rabei (Roze and Solano 1963)

Holotype: MBUCV 5126.

Type locality: “Al pie de Cerro Lema, Rio Chicanan,

Bolivar, Venezuela.”

Distribution: Region 5. Venezuela and Suriname. In

Venezuela, a few localities in Bolivar State and Delta

Amacuro, south of Orinoco River.

Selected references: Roze and Solano (1963); Taylor

(1968); Nussbaum and Wilkinson (1989); Péfaur et al.

(1992); Maciel and Hoogmoed (2011b); Sefiaris et al.

(2014).

Genus Siphonops Wagler, 1828

Type species: Caecilia annulata Mikan, 1820, by

monotypy.

Siphonops annulatus (Mikan 1820)

Lectotype: RMNH 2419.

Type locality: “Sebastianopolis (Rio de Janeiro), Brasil.”

Distribution: Region 1. Widely distributed east of Andes

in Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil,

and French Guiana. In Venezuela, eastern slopes of the

Andes in Barinas and Portuguesa States.

Selected references: Dunn (1942); Ginés (1959); Rohl

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Catalogue of the amphibians of Venezuela

293. Bolitoglossa mucuyensis. La Mucuy, Mérida. Photo: Ariel

Espinosa-Blanco.

294B. Bolitoglossa orestes. La Bravera, way from Mérida to La

Azulita, Mérida, near type locality of the synonym B. spongai.

Photo: César Barrio-Amoros.

295. Bolitoglossa tamaense. Matamula, Tachira. Photo: Andrés

Chacon.

(1959); Taylor (1968); Péfaur and Diaz de Pascual

(1982); Nussbaum and Wilkinson (1989); Péfaur et al.

(1992); Lynch (1999); Maciel and Hoogmoed (201 1a).

Family Tiphlonectidae Taylor, 1968

Genus Nectocaecilia Taylor, 1968

Type species: Chthonerpeton petersii Boulenger, 1882,

by original designation.

Nectocaecilia petersii (Boulenger 1882)

Amphib. Reptile Conserv.

294A. Bolitoglossa orestes. Sierra de la Culata, Mérida. Photo:

César Barrio-Amoros.

294C. Bolitoglossa orestes. La Bravera, way from Mérida to La

Azulita, Mérida. Photo: César Barrio-Amoros.

a 23

iid fig saad ‘i

Seek

296. Eanes niger. ABiazil Pe ae TET ae

Holotype: BMNH 1946.9.5.68.

Type locality: “Upper Amazon.”

Distribution: Region 4. Colombia, Venezuela, and

Brazil. In Venezuela, southern half of Amazonas State.

Selected references: Dunn (1942); Taylor (1968);

Nussbaum (1977); Nussbaum and Wilkinson (1989);

Gorzula and Sefiaris (1998); Maciel and Hoogmoed

(2011).

Genus Potomotyphlus Taylor, 1968

Type species: Caecilia kaupii Berthold, 1859, by original

designation.

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Barrio-Amoros et al.

297. Microcaecilia naaBe Imataca Forestal Reserve, ‘Bolivar-Delta

Amacuro boundary. Photo: Bf ernando iat ag

Va —— F are :

f ef a rt ate 7 ag oe.

ed f-, ‘ i

- 4

299. Potomotyphlus kaupii. Caroni river, Bolivar. Photo: Fernando

Rojas-Runjaic.

300B. Zyphlonectes natans. Puerto Concha, Zulia. Photo: Alan

Highton.

Potomotyphlus kaupii (Berthold 1859)

Holotype: ZFMK 27684.

Type locality: “Angostura,’ Ciudad Bolivar, Venezuela.

Distribution: Regions 3, 4, 5. Orinoco and Amazons

Basins, in Venezuela, Colombia, Ecuador, Peru, Brazil,

and French Guiana. In Venezuela, northern Amazonas

and Bolivar States, and Delta of the Orinoco (Delta

Amacuro).

Amphib. Reptile Conserv.

298, Siphonops annulatus. aannies Barinas. Photo: César

Barrio-Amoros.

~~ i +

300A. ephichertes ae Rea eure Zulia. Photo: Cesar

Barrio-Amoros.

Remarks: CBA saw a photo of a fisherman in San

Vicente, Rio Apure (a major tributary of the Orinoco)

which obtained one specimen fishing and used it as bait.

Selected references: Taylor (1968); Nussbaum and

Wilkinson (1989); Péfaur et al. (1992); Maciel and

Hoogmoed (201 1a).

Genus Typhlonectes Peters, 1880

Type species: Caecilia compressicauda Dumeril and

Bibron, 1841, by subsequent designation of Dunn (1942).

Typhlonectes compressicauda Dumeéril et Bibron, 1841

Lectotype: MNHNP 4269.

Type locality: “Cayenne,” French Guiana.

Distribution: Region 4. Amazon Region, in Colombia,

Venezuela, Guyana, French Guiana, Brazil, and Ecuador.

In Venezuela, only Puerto Ayacucho (Gorzula and Cerda,

1979; Barrio-Amoros 1998).

Selected references: Roze (1963); Roze and Solano

(1963); Taylor (1968); Gorzula and Cerda (1979),

Nussbaum and Wilkinson (1989); Péfaur et al. (1992);

Wilkinson (1996a, 1996b); Lynch (1999); Maciel and

Hoogmoed (201 1a).

July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

Typhlonectes natans (Fischer 1880)

Syntypes: ZMB 9522-23 (two specimens).

Type locality: “Cauca, Nebenfluss des Magdalenenstroms

in Neu-Granada” = Cauca River, tributary of Magdalena

River, Colombia.

Distribution: Region 6. Western and northern Colombia,

and Maracaibo Lake Basin in northwestern Venezuela.

Remarks: 7yphlonectes venezuelensis Fuhrmann, 1914

and Chthonerpeton haydeeae Roze, 1963 are junior

synonyms of this species.

Selected references: Roze (1963); Roze and Solano

(1963); Taylor (1968); Lancini (1969); Nussbaum and

Wilkinson (1989); Péfaur et al. (1992); Wilkinson

(1996a,b); Lynch (1999).

Species which may occur in Venezuela

Five of the species predicted to be in the country from

Barrio-Amoros (1998) first approach to the Venezuelan

amphibian fauna were posteriorly reported to occur

in Venezuela: Rhaebo glaberrimus, Leptodactylus

colombiensis, Phytotriades auratus, Elachistocleis

pearsei, and one species of Cryptobatrachus, C.

remotus (Chacon-Ortiz et al. 2001; Barrio-Amoros and

Chacon-Ortiz 2001; Infante-Rivero et al. 2006b, 2009;

Rivas and De Freitas 2015). Some taxa predicted later

by Barrio-Amoros (2004) have lasted in recent reports,

like Yepuihyla exophthalma, Amazophrynella minuta,

and Phyllomedusa venusta (Barrio-Amoros et al. 2010;

Rojas-Runjaic et al. 2013; Infante-Rivero et al. 2006a).

Many more predicted species have either failed detection,

or do not truly occur in Venezuela. Below are some taxa

which so far remain only as “predicted” or “expected”

for the Venezuelan fauna, especially those known from

close localities in Guyana, Brazil, and Colombia.

The Andes. Biogeographically, some taxa known

from nearby Colombia are expected in the Venezuelan

Andes, in the Cordillera de Mérida, the Tama massif,

or the Serrania de Perija. Pristimantis cuentasi Lynch,

2003 and P. reclusus Lynch, 2003 are known from

paramo over 3,000 m in shared habitat with Venezuela

(Serrania de Perija), but not yet reported from the

country. A/lobates ignotus Anganoy-Criollo, 2012 is an

aromobatine also likely to be found on the Venezuelan

side of Perija. Strabomantis ingeri (Cochran and Goin

1963) has been observed in Colombia very close to the

Venezuelan border in the Cordillera Oriental (Ramirez-

Pinilla 2004). Pristimantis douglasi Lynch, 1996 could

be present in the Tama cloud forests or even in Sierra

de Perija. Pristimantis frater (Werner 1899) has been

recently reported from the Colombian side of the

Tama massif (Acevedo et al. 2014). Another species

which maybe present on the Venezuelan side of Tama

is Cryptobatrachus conditus Lynch, 2008. Hyloscirtus

callipeza Duellman, 1989 is known from extreme NE

Amphib. Reptile Conserv.

Cordillera Oriental in Colombia, and could also be

present on its Venezuelan side or in Perija. Another

high Andean species is Niceforonia nana (Goin and

Cochran 1963), known to occur on the western versant

of The Cordillera Oriental in Departamento Norte de

Santander (http://www. batrachia.com/orden-anura/

craugastoridae/niceforonia-nana/); it could occur as well

on the Venezuelan side of Tama or Perija. Populations

of Lithobates palmipes from the Maracaibo Lake Basin

could prove to represent L. vaillanti (Brocchi 1877), as

these species are practically indistinguishable. Moreno-

Arias et al. (2010) mention Bolitoglossa biseriata Tanner,

1962 from La Jagua de Ibirico, Departamento Cesar

at 1,450 m in Serrania de Perija, Colombia. A. Acosta

identifies it as B. /ozanoi Acosta and Restrepo, 2001

(http://www. batrachia.com/orden-caudata/bolitoglossa/

bolitoglossa-biseriata/) so this species could be present

on the Venezuelan side as well. Moreno-Arias et al.

(2010) also mention Pristimantis taeniatus Boulenger,

1912 and P. viejas Lynch et Rueda-Almonacid, 2000

as present on the Colombian side of Perija, but this

statement is not followed or corroborated by A. Acosta

(http://www. batrachia.com).

The Guiana Shield. In Guyana, even on the other

side of Roraima tepui, several described species have

not yet reported from Venezuela. Among them are

Myersohyla kanaima (Duellman and Hoogmoed 1992),

Tepuihyla warreni (Duellman and Hoogmoed 1992),

Trachycephalus hadroceps (Duellman and Hoogmoed

1992), Stefania evansi (Boulenger 1904), S. roraima

Duellman and Hoogmoed, 1984, and S. woodleyi Rivero,

1968. The recently described Boana diabolica (Fouquet,

Martinez, Zeidler, Curtois, Gauchier, Blanc, Dias-Lima,

Marques-Souza, Rodrigues and Kok 2016) could also

be present in southeastern Venezuela. Adenomera lutzi

Heyer, 1975 is present along the Pakaraima mountains

in Guyana, which are continuous with eastern Gran

Sabana of Venezuela (Kok et al. 2007). Chiasmocleis

shudikarensis Dunn, 1949 was described from the Upper

Essequibo River, and is widely distributed in the Guianas

and northern Amazonia. Other Guiana shield inhabitants

that may be present in Venezuela include:

Rhinella martyi

Fouquet, Gaucher, Blanc and Vélez-Rodriguez, 2007

Fouquet et al. (2007) and Reynolds and MacCulloch

(2012) show a locality in western Guyana (Baramita)

near the Venezuelan border. As the R. margaritifera group

requires further study, the presence of many described

and undescribed species under this name would not be

surprising. Avila-Pires et al. (2010) synonymized this

name into R. margaritifera, which does not resolve the

paraphyly that this name hides. Lavilla et al. (2013)

considered this action premature, and Frost (2018) still

counted it as valid.

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

Ameerega hahneli (Boulenger 1884)

Reported on different occasions from Venezuela (Lotters

et al. 2007) with no vouchers or photos to sustain the

claims; however, it could be present in the southern

and easternmost parts of the country. Reynolds and

MacCulloch (2012) report it from Baramita, Guyana,

very near the Venezuelan border. Specimens from

Baramita, however, should be compared with Ameerega

picta (guayanensis).

Dendrobates nubeculosus Jungfer and Bohme, 2004

Only known from its type locality, on the Essequibo

River, Guyana. May be an arboreal inhabitant that has

been completely overlooked. As many colleagues have

commented (see Guayasamin et al. 2006 for a resumé),

the next realm for exciting discoveries is the exploration

of the rainforest high canopy.

Adelophryne patamona MacCulloch, Lathrop, Kok,

Minter, Khan and Barrio-Amoros, 2008

This recently described minute species has _ been

confused with Adelophryne gutturosa, a common species

in Venezuela, but is only known from a few specimens

and localities due to the difficulty of its collection in leaf

litter.

Leptodactylus myersi Heyer, 1995

Reported from rocky outcrops in French Guiana,

Suriname, Guyana, and the neighboring Brazilian state

of Roraima, this species seems not to be present in

Venezuela, probably due to niche competition with L.

rugosus, a similar-sized inhabitant of the same habitat

from eastern Venezuela.

Dendropsophus counani Fouquet, Del Orrico, Ernst,

Blanc, Martinez, Vacher, Rodrigues, Ouboter, Jairam

and Ron, 2015

This recently described species of the D. parviceps

group 1s known from the lowlands of the Guiana Shield,

including localities in Guyana approximately 100 km

from the eastern Venezuelan border, and could be present

in the lowlands of the Cuyuni River Basin.

Boana liliae (Kok 2006)

Reported only from Kaieteur National Park in Guyana.

However, as many other species known until recently

from that area of Guyana, it could also be present in Sierra

de Lema or other parts of the Gran Sabana in Venezuela.

Osteocephalus oophagus Jungfer and Schiesari, 1995

Amphib. Reptile Conserv.

Known from nearby Colombia in Departamento de

Guania (Lynch and Vargas-Ramirez 2000) and Guyana

(Kok and Kalamandeen 2008; Jungfer et al. 2013). It

is similar to Osteocephalus taurinus in shape and iris

coloration, but is much smaller. Examining O. taurinus

or O. leprieurii samples in Venezuelan collections may

lead to the recognition of this species in Venezuela.

Elachistocleis surumu Caramaschi, 2010

Recently Caramaschi (2010) described from Pacaraima

in Brazil, very close to Santa Elena de Uairén and the

Gran Sabana. Perhaps those Elachistocleis sp. mentioned

by Duellman (1997) could represent this species.

Lysapsus laevis (Parker 1935)

An aquatic hylid that could be present in the southeastern

region of Venezuela, close to Brazil and Guyana (Cole et

al. 2013).

Furthermore, many different species of caecilians

have been described and are known from Guyana

(Caecilia gracilis Shaw, 1802, Caecilia pressula Taylor,

1968, Oscaecilia zweifeli Taylor, 1968, Rhinatrema shiv

Gower, Wilkinson, Sherratt and Kok, 2010, Caecilita

iwokramae Wake and Donnelly, 2010, Microcaecilia

iyob Wilkinson and Kok, 2010) (see Cole et al. 2013

and references therein). Caecilians are poorly studied

amphibians and a much larger diversity could be present

in Venezuela.

The Amazon. The Amazonian Region, 1s one of the

less explored areas in Venezuela. Several widespread

Amazonian species are expected to occur inside

Venezuelan borders, as recently discovered with

Amazophrynella minuta (Rojas-Runjaic et al. 2013).

Other such species include Teratohyla midas (Lynch

et Duellman 1973), which is distributed throughout

the upper, middle, and lower Amazon from 1,000 m

in the eastern piedmont of Ecuador to French Guiana.

Such distribution, paralleled by HAyalinobatrachium

iaspidiense and H. mondolfii, make its presence likely in

the southern or southeasternmost corners of Venezuela.

Pipa snethlageae Miller, 1914 has been mentioned from

nearby Colombia at Departamento de Guainia by Lynch

and Vargas-Ramirez (2000).

The Maracaibo Basin. Zulia state in Northwestern

Venezuela is connected with northern Colombia

through the adjacent lowlands and foothills of the

Serrania de Perija. Several species known from the

neighboring country could be present in the Venezuelan

side as well. Scinax boulengeri (Cope 1887) extends

from Central America to the Colombian Caribbean. It

should be examined if the reported S. rostratus from

the Maracaibo Lake Basin does not correspond to this

July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

species. Similarly, its presence would not be a surprise

in the area of Craugastor raniformis (Boulenger 1896),

Scinax elaeochrous (Cope 1876), Dendropsophus

ebracattus Cope, 1874, or Dendropsophus phlebodes

Stejneger, 1906. The complex of Rhinella granulosa has

been a difficult group of bufonids to assess (Gallardo

1965; Narvaes and Rodrigues 2009; Pereyra et al. 2015).

The recently described Central American (Panamanian)

populations as R. centralis Narvaes and Rodrigues, 2009

can be more widely expanded through open habitats in

northern Colombia and NW Venezuela. Specimens from

Zulia State recognized as R. beebei should be closely

compared with R. centralis. Dendrobates truncatus

(Cope 1861) is known from localities in the Magdalena

River Valley close to the western versant of Serrania de

Perija, and may be present in its Venezuelan counterpart.

Elachistocleis panamensis (Dunn, Trapido and Evans

1948) is distributed along the Magdalena River Valley,

extending to the dry areas of La Guajira, and thus could

also be present in Venezuelan territory.

Conservation

Amphibians are facing an extinction crisis. Globally

at least 32% of all described species are currently

threatened with extinction (IUCN 2016), a number

that may be underestimated because amphibian faunas

are poorly-known in many regions. This percentage of

threatened amphibians is noticeably higher than other

vertebrate groups that have been comprehensively

assessed, specifically mammals (23%) and birds (12%)

[Baillie et al. 2004; IUCN 2016]. The factors driving the

global amphibian population declines are diverse, but are

predominantly anthropogenic and intrinsically linked to

human population growth (Gascon et al. 2007). Habitat

loss is the number one threat to amphibians, followed

by pollution. The emerging disease chytridiomycosis,

caused by Batrachochytrium dendrobatidis (Bd), and

climate change are also important threats, and have been

linked to rapid enigmatic amphibian declines, especially

in relatively cool, wet, diverse habitats in alpine,

temperate, and tropical areas (Hof et al. 2011; Catenazzi

2015; Berger et al. 2016).

There have been several attempts to assess the

extinction risk for Venezuelan amphibians, direct or

indirectly (Vial and Saylor 1993; La Marca 1995;

Rodriguez and Rojas-Suarez 1995, 1999, 2008; Barrio-

Amoros 2001; Young et al. 2004; Molina et al. 2009;

Barrio-Amoros and Torres 2010). More recently,

Rodriguez et al. (2015) re-assessment of the extinction

risk of Venezuelan fauna mentioned that 12% of the

amphibian species are threatened. About half of the

species of amphibians (47%) in this assessment are

reported as Least Concern, 13% as Near Threatened,

12% as threatened (Vulnerable, Endangered, or Critically

Endangered), and 23% as Data Deficient. Only one

species 1s considered extinct, the harlequin toad Ate/opus

Amphib. Reptile Conserv.

vogli, and about 5% of the Venezuelan amphibian fauna

is still under evaluation.

Based on these results, the most threatened group of

amphibians are the harlequin toads, Ate/opus, followed

by the aromobatines of the genera “Prostherapis,”

Aromobates, and Mannophryne, and the land frogs of the

genus Pristimantis. Additionally, some of the species of

these groups are Near Threatened or In Evaluation, thus

they will possibly become categorized as threatened in

the future. Also, the high proportions of species in the

Data Deficient category (23%) can significantly mask the

real number of amphibians in risk, and this is especially

noticeable in the families Craugastoridae, Dendrobatidae,

and some Hylidae. All of these threatened amphibians

are Venezuelan endemics, and in most cases, they have a

small geographic distribution range in medium elevation

or highlands of the Andes, Coastal Ranges, or the

Pantepul.

Acknowledgements.—The present work is based on

preceding versions which tried to give a glimpse of the

Venezuelan amphibian fauna (Barrio-Amoroés 1998,

2004) and all people acknowledged in those lists must

be recognized here again. Herein we produced a more

comprehensive annotated list, including images of all

possible amphibians present in Venezuela. Such an

enormous task was not possible without continuous

conversations with colleagues, constructive discussions,

and help in obtaining difficult papers and photographs.

We herein thank the following people, in alphabetical

order (by first name): Alan Highton, Aldemar A. Acevedo,

Allan Markezich, Amelia Diaz de Pascual, Amy Lathrop,

Andreas Schluter, Andrés Chacon, Andrés Gonzalez,

Andrés Orellana, Antoine Fouquet, Ariel Espinosa,

Arlene Cardozo, Blair Hedges, Brad Wilson, Brice

Noonan, Bruce Means, Carmen Ferreira, César Molinart,

Charles Brewer-Carias, Daniel Calcafio, Daniel Cuentas,

Daniel Quihua, Danté Fenolio, David Blackburn, Denis

Torres, Dennis Rédder, Diego A. Flores, Eduard Filella,

Eduardo Asens, Edward Camargo, Edwin Infante-Rivero,

Eliécer Gutiérrez, Eric Smith, Erik Arrietat, Esteban

Lavilla, Francisco Lopez, Franklin Rojas-Suarez, Gabriel

Ugueto, Gilson Rivas, Hinrich Kaiser, Jaime Bautista,

Jaime Péfaur, Javier Garcia, Javier Mesa, Jesus Salas,

Joan Garcia-Porta, John D. Lynch, Jorge Bravo, Jose Luis

Vieira, Jose Manuel Padial, Jose Pombal, José Rosado,

Juan Carlos Ortiz, Juan Carlos Santos, Juan David

Jiménez, Juan E. Garcia-Pérez, Juan M. Guayasamin,

Juan M. Renyifo; Juan Pablo Diasparra, Juan S. Mendoza,

Julian Faivovich, Liz del Valle, Luis Coloma, Luis Merlo,

Luis Scottt, Manuel Gonzalez, Margarita Lampo, Mark

Moffett, Matthew Heinicke, Mayke de Freitas, Mercedes

Salazar, Miguel Molinari, Moisés Escalona, Oscar Lasso-

Alcala, Oswaldo Fuentes, Pablo Velozo, Patricia Salerno,

Pau Cardellach, Pat Gutiérrez-Cardenas, Philippe Kok,

Rafael Morillo, Rainer Schulte, Ramon Rivero, Roger

Manrique, Ross MacCulloch, Roy McDiarmid, Salvador

July 2019 | Volume 13 | Number 1 | e180

Barrio-Amoros et al.

Carranza, Santiago Castroviejo-Fisher, Saul Gutiérrezy,

Sebastian Lotzkat, Stefan Lotters, Tito R. Barros, Tom

Evenue, Ulisses Caramashi, Walter Schargel, William

E. Duellman, Zaida Tarano, and Zelimir Cernelic, who

contributed in some way with the final publication of

this work. Miguel A. Alonso-Zarazaga responded some

concerns about nomenclature.

We are finally deeply grateful for four anonymous and

five known reviewers which made great contributions

by improving the English, making points about the

content, and editing the whole text. Among them are

Hinrich Kaiser, Pedro Peloso, Philippe Kok, Ted Kahn,

and Ross MacCulloch. Thanks also to Craig Hassapakis

who encompassed the whole project for more than

three years and to Tyson Terry and Michael Grieneisen,

the managing editors of this particular paper, for their

patience and good nature.

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César L. Barrio Amords is a graduate in Geography and History

(Anthropology) from the Universitat de Barcelona, Spain, and was co-

founder and Director of Fundacion AndigenA in Venezuela from 1999

to 2009. As a member of several exploration teams, César has produced

more than 200 publications and described 51 new taxa of amphibians

and reptiles, mainly from Venezuela. His main interests are taxonomy of

Terraranae, Arboranae, and Dendrobatoids, as well as some reptile groups;

biogeography of the Pantepui based on herpetofauna; and natural history

of amphibians at the Pacific versant of Central America. César is currently

Photo: Lucas Bustamante.

Fernando J.M. Rojas-Runjaic is a researcher at the Museo de Historia

Natural La Salle in Caracas, Venezuela, as well as curator of the amphibian,

reptile, and arachnid collections. Fernando has a B.Sc. in Biology

from the Universidad del Zulia (Venezuela), and a Master’s degree in

Biodiversity and Conservation of Tropical Areas from the Universidad

Internacional Menéndez Pelayo (Spain). While currently enrolled in the

Ph.D. program in Zoology at the Pontificia Universidade Catoélica do Rio

Grande do Sul (Brazil), Fernando’s research interests are broad and include

evolution, phylogenetic systematics, taxonomy, diversity, biogeography,

and conservation of amphibians, reptiles, and scorpions. Photo: Santi

Castroviejo Fisher.

J. Celsa Sefiaris is a Biology graduate from the Universidad Central

de Venezuela (UCV), with a Ph.D. from Universidad de Santiago de

Compostela on Venezuelan centrolenids. She was director of the Museo de

Historia Natural La Salle from 1991 to 2012, and is currently a researcher

at the Laboratorio de Ecologia y Genética de Poblaciones del Instituto

Venezolano de Investigaciones Cientificas (IVIC) as well as a professor

of Herpetology at the UCV and IVIC. She is the author of three books and

89 papers on Venezuelan herpetofauna, with two genera and 37 species

described, and has served as editor of 12 scientific books. She is a founding

member of PROVITA, one of the most important NGOs in Venezuela,

and has coordinated six expeditions in the Guayanan region of Venezuela.

Since 2005, she has been co-president for Venezuela of the Amphibian

Specialist Group (ASG), IUCN. Photo: J. Celsa Sefiaris.

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Barrio-Amoros et al.

Appendix 1. Species known to inhabit Venezuela but not yet described or correctly identified.

Several species of amphibians known to be in Venezuela lack proper names. Some are undescribed (Ate/opus sp

1, Pristimantis sp 4); some are incorrectly identified (even for a long time), e.g., Allobates aff. marchesianus, or

the Osteocephalus taurinus complex, with at least two different candidate species identified in Venezuela (Jungfer

et al. 2013); others include a new Adenomera sp. Q (Fouquet et al. 2014), and the five new candidate species of

Stefania reported by Kok et al. (2016), which all deserve proper descriptions and names. Certain taxa, like Rhinella cf.

proboscidea are documented by a photograph, although without vouchers. However, those complexes with more than

one taxon (for example Boana xerophylla complex, apparently with at least three or four different putative species)

are not counted as individual species herein. For general purposes, this listing accounts for species included in the

biodiversity of Venezuela, while acknowledging the need to properly work on them. McDiarmid and Paolillo (1988)

provide a very interesting list of species collected during the Neblina expeditions (from 1983 to 1987), which are

mostly in the USNM and AMNH. The importance of checking these collections is acknowledged, as many species

await proper attention. Among them, this catalog does not include species without at least a recognized genus. For

example, the Bufonid sp. nov. from the higher camps of Neblina could be a Metaphryniscus, but without corroborating

data it is not included on this list. In the case of the many “Eleutherodactylus” from Neblina, only those with a decent

number of collected specimens are counted as candidate species. Of these 33 species, some are mentioned for the first

time herein; while some have been mentioned in the literature but are not correctly identified or yet described.

Family Bufonidae

Atelopus sp. |

Distribution: Endemic to Macizo de Guaramacal, between Trujillo and Portuguesa states, Venezuelan Andes.

Remarks: Though this new Ate/opus was discovered in December 1987, it has not been described yet (Garcia-Pérez

and La Marca 2015). Its population status has been mentioned several times (Garcia-Pérez 1997, 2005; Garcia-Pérez

and La Marca 2015). It appears as Ate/opus sp. 2 in Rueda-Almonacid et al. (2005). The proper description of this

species 1s imperative.

Selected references: Garcia-Pérez 1997; La Marca and Lotters 1997; La Marca 2004; Garcia-Pérez 2005; La Marca

et al. 2005; Lotters et al. 2005; Rueda-Almonacid et al. 2005.

Atelopus sp. 2

Distribution: Endemic to near Queniquea, estado Tachira.

Remarks: This species was collected in 1988 and is not yet described (A. Chacon and CBA, in prep.).

Rhinella cf. proboscidea (Spix, 1824)

Holotype: ZSM 1145/0

Type locality: River Solimoes, Brazil.

Distribution: Upper and middle Amazon, from Peru to Manaus. This species is referred in Venezuela through a picture

by Javier Mesa taken in the middle section of La Escalera, Sierra de Lema, Bolivar state (Image 26), that was compared

with similar Rhinella (proboscidea, dapsilis). At this time, we concude that it looks more like proboscidea than any

other species, pending the collection and examination of more specimens.

Family Centrolenidae

“Cochranella” sp.

Distribution: Temiche, base of Mount Marahuaka, Amazonas state.

Remarks: Rivero (1961) reported a “Cochranella sp.” from Temiche, at the base of Mount Marahuaka, a tepui in

Amazonas state, Venezuela. Goin (1968) mentions it could be AHyalinobatrachium taylori, but its poor condition

prevents identification. It also could be “C.”’ duidaeana, known from a higher altitude (2,140 m) in the same general

area, or any other kind of glassfrog (or a Hylid, as Rivero proved to confuse his Centrolenella pulidoi also from Duida

with Boana benitezi — Faivovich et al. 2005; this work). As this area has been not well studied, more attention must

be paid for new species. Unfortunately, old specimens of glassfrogs lose their characteristics quickly if not properly

preserved, such as the specimen MCZ 28569.

References: Goin 1968; Rivero 1961.

Family Dendrobatidae

Allobates aff. marchesianus (Melin, 1941)

Syntype: NHMG 509.

Type locality: “Taracua, Rio Uaupés [Estado do Amazonas, ] Brazil”.

Distribution: Reported from the foothills of Cerros Duida and Marahuaka, two neighboring tepuis in Amazonas state.

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Catalogue of the amphibians of Venezuela

Remarks: Considered herein under one putative species name are all reports of A//obates brunneus and A. marchesianus

from Venezuela, until the proper identity of such populations is resolved. They are considered aff. marchesianus rather

than aff. brunneus as the marchesianus type locality is closer to the Venezuelan southern border and the possibility of

its presence is still likely. A//obates brunneus 1s certainly not present in Venezuela, as its type locality lies south of the

Amazon and, though similar, the Venezuelan material does not fall into the recent redescription of A. brunneus from

its type locality (Lima et al. 2009).

Selected references: Caldwell et al. 2002; La Marca 1996e; Lima et al. 2009; Morales 1994; Rivero 1961,

1964a,b,d, 1988.

Allobates “Neblina species”

Remarks: This is the Co/osthetus sp. A in McDiarmid and Paolillo (1988) and already tested by Grant et al. (2006).

It awaits formal description.

Aromobates sp. |

Remarks: This species was discovered during the Calderas RAP (Barrio-Amoros 2010; Barrio-Amoros and Molina

2010) and demonstrated as a candidate species by Barrio-Amoros and Santos (2012); however, it was not described

since the only materials were tadpoles and recent metamorphs.

References: Barrio-Amoros 2010a; Barrio-Amoros and Molina 2010; Barrio-Amoros and Santos 2012.

Mannophryne sp. 1| (aff. herminae)

Distribution: Santos and Barrio-Amoros (genetic unpub. data) indicate that Manophryne from Bejuma and

surroundings in Carabobo state, is not the same M. herminae that inhabits Rancho Grande. FRR, based on preliminary

morphology and bioacoustics, suggests that 1Z. herminae from RG is apparently conspecific to those in San Esteban

valley, but differs from the population in Bejuma.

References: Barrio-Amor0os 2006; Barrio-Amoros et al. 201 0b.

Mannophryne sp. 2

Distribution: The Mannophryne sympatric with M. oblitteratus at Guatopo National Park is indeed a new species to

be described (J.C. Santos and CBA; unpub. data). Barrio-Amoros and Santos refrained from describing it due to the

lack of permits at the time.

Family Hemiphractidae

Stefania sp. |

Distribution: Serrania de la Neblina, Amazonas state.

References: McDiarmid and Paolillo 1988.

Stefania sp. 2

Distribution: Abakapa-tepui, Bolivar state.

References: Kok et al. 2016.

Stefania sp. 3

Distribution: Angasima-tepui, Bolivar state.

References: Kok et al. 2016.

Stefania sp. 4

Distribution: Upuigma-tepui, Bolivar state.

References: Kok et al. 2016.

Stefania sp. 5

Distribution: Amuri-tepui, Bolivar state.

References: Kok et al. 2016.

Stefania sp. 6

Distribution: Murisipan-tepui, Bolivar state.

References: Kok et al. 2016.

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Family Hylidae

Dendropsophus aff. minutus (Peters, 1872)

Syntypes: ZMB 7456 (five specimens. D. minutus sensu stricto).

Type locality: Nova Friburgo, Rio de Janeiro, Brazil (for D. minutus sensu stricto).

Distribution: Based on the phylogeny of Gehara et al. (2014) the Guianan lineages are widely distributed in Venezuela

(Andes, Cordillera de La Costa, Amazon, and Guiana Shield), Guiana, Suriname, French Guiana, and northern Brazil.

No name is available yet for those populations that could represent more than one species.

Remarks: Gehara et al. (2014) showed this is a composite of species with at least four mitochondrial lineages in

Venezuela that are not part of the clade of D. minutus species complex. Gehara et al. (2014) proposed to remove

Hyla goughi Boulenger, 1911 from the synonymy of D. minutus and allocate it in the “Guianan lineages”. However,

in a recent visit to the BMNH, CBA corroborated the holotype of Hyla goughi from Trinidad as Dendropsophus

microcephalus.

Selected references: Barrio-Amoros 2010a; Barrio-Amoros and Duellman 2009; Barrio-Amoros et al. 2011b; Donnelly

and Myers 1991; Duellman 1997; Gehara et al. 2014; Gorzula and Sefiaris 1998; Heatwole et al. 1965; Hoogmoed

and Gorzula 1979; Magdefrau et al. 1991; Rivero 1961, 1964b, 1971b; Rivero and Esteves 1969; Sefiaris et al. 2014.

Boana sp. (cf. rufitela)

Distribution: A few localities along the Caribbean coastal range and Maracaibo Lake basin, also scattered in Caribbean

Colombia. Those localities are widespread in the literature but there 1s no mention of a viable population.

Remarks: In all previous lists of Venezuelan amphibians this species was confused with H. albomarginatus, a species

found in southeastern Brazil. Barrio-Amoros (2004) identified Venezuelan populations as H. rufitelus, apparently the

closest similar congeneric, known from Nicaragua through southern Central America to northern Colombia. Only one

specimen collected in Maracaibo, Zulia, Venezuela (MCZ 15369; Fig. 185) looks very much like H. albomarginatus; all

other citations are based on misidentifications of juvenile H. crepitans or H. pugnax, both in Venezuela and Colombia.

Here it is concluded that: 1- the specimen MCZ 15369 from a site in Brazil was mislabeled and corresponds with H.

albomarginatus, or 2- there is an unidentified species (not a/bomarginatus or rufitelus) to be collected and described in

Caribbean Venezuela and Colombia.

Selected references: The following references are about either H. rufitelus or H. albomarginatus: Barrio-Amoros

2004, 2013; Fouquette 1968; Ginés 1959; Lutz 1927; Mendoza 2014; Rivero 1961; Spix 1824; Tello 1968.

Osteocephalus aff. planiceps Cope, 1874 [Cal_Neblina411]

Referred specimen: AMNH A-131254.

Distribution: Only known from base of Cerro de la Neblina (Base Camp, Rio Baria, Amazonas state) at southern

border of Venezuela. Probably more extensive through southern Venezuela and northern Brazil.

Remarks: There is only a single report of this species in the country by Jungfer et al. (2013), based on specimen

AMNH A-131254. In the phylogeny of Osteocephalus inferred by these authors, the species is recovered as sister to

a Clade integrated by O. leoniae, O. fuscifacies, and O. deridens, and not clustered with any terminal of O. planiceps.

Based on this phylogenetic evidence, they refer it as a confirmed candidate species [Cal_Neblina411] related to

Osteocephalus planiceps. In the Osteocephalus planiceps species group (Jungfer et al. 2013). Description of this new

species 1s pending. Other specimens collected during the 1983-1987 expedition to Neblina and deposited in AMNH,

USNM and MBUCYV as O. taurinus (McDiarmid and Paolillo 1988) probably correspond to this new taxon.

References: McDiarmid and Paolillo 1988; Jungfer et al. 2013.

Osteocephalus aff. taurinus [Ca2_Neblina410]

Referred specimen: AMNH A-131245.

Distribution: Only known from the base of Cerro de la Neblina (Base Camp, Rio Baria, Amazonas state) at the

southern border of Venezuela. Probably more extensive through southern Venezuela and northern Brazil.

Remarks: Jungfer et al. (2013) discovered this species from a single specimen (AMNH A-131245). In the phylogeny

of Osteocephalus inferred by these authors it is recovered as sister to a clade integrated by O. oophagus, O. taurinus,

and three other undescribed species, and is not clustered with any terminal of O. taurinus sensu stricto. They refer it as

a confirmed candidate species [Ca2_Neblina410] related to O. taurinus. In the Osteocephalus taurinus species group

(Jungfer et al. 2013). Description of this new species is pending. Other specimens collected during the 1983-1987

expedition to Neblina and deposited in AMNH, USNM and MBUCYV as O. taurinus (McDiarmid and Paolillo 1988)

probably correspond to this new taxon.

References: McDiarmid and Paolillo 1988; Jungfer et al. 2013.

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Osteocephalus aff. taurinus [Ca3_AJC2959_ 3181]

Referred specimens: MHNLS 18325, 19907, 20034, and 20083.

Distribution: Only known from northwestern Amazonas state, southern Venezuela.

Remarks: In the phylogeny of Osteocephalus inferred by Jungfer et al. (2013) this taxon is recovered as sister to a

clade integrated by O. oophagus, O. taurinus, and two other undescribed species, and is not clustered with any terminal

of O. taurinus sensu stricto. Jungfer et al. (2013) refer it as a confirmed candidate species [Ca3_AJC2959 3181]

related to O. taurinus. In the Osteocephalus taurinus species group (Jungfer et al. 2013). Description of this new

species 1s pending. Numerous specimens deposited in MHNLS, EBRG, and MBUCYV as O. taurinus (McDiarmid

and Paolillo 1988) probably correspond to this new taxon. The identity of populations in Bolivar and Delta Amacuro

previously referred as O taurinus (Barrio-Amoros 1998; Gorzula and Sefiaris 1999; Sefiaris and Ayarzagtiena 2004)

must be reevaluated.

References: Jungfer et al. 2013.

Family Leptodactylidae

Adenomera sp. |

Distribution: A single sample was analyzed by Fouquet et al. (2014) from the border between Brazil, Colombia, and

Venezuela.

Remarks: This is the Adenomera sp. Q of Fouquet et al. (2014).

References: Fouquet et al 2014.

Adenomera cf. simonstuarti Angulo and Icochea, 2010

Holotype: MHNSM 18218.

Type locality: “Campamento Segakiato, c. 340 m asl, Rio Camisea, District of Echarate, Province of La Convencion,

Region of Cusco, Peru.”

Distribution: Recently described from the eastern versant of the Peruvian Andes. Widely distributed along the eastern

Andean piedmont from Peru to Venezuela. In Venezuela, reported from Calderas, Barinas state by Barrio-Amoros

(2010a) as Adenomera sp.

Remarks: Fouquet et al. (2014) found a sample from Calderas, Barinas, and embedded it into a clade with A.

simonstuart.

Selected references: Angulo and Icochea 2010; Barrio-Amoros et al. 2010a; Fouquet et al. 2014.

Leptodactylus sp. 1

Distribution: Andes of Venezuela, at least in Mérida, Trujillo, and Tachira states, around 1,000 m asl (CBA personal

observation). E. La Marca restricts it to the Meseta de Merida.

Remarks: Currently in description by E. La Marca, who listed it as Endangered in the fourth edition of the

Venezuelan Red List (La Marca 2015).

References: La Marca 2015.

Leptodactylus aff. macrosternum Miranda-Ribeiro, 1926

Type: MZUSP 448.

Type locality: Bahia (Brazil).

Distribution: Venezuela north of the Orinoco river, in open areas.

Remarks: De Sa et al. (2014) restrict the distribution of L. macrosternum to its type locality in Bahia, Brazil, leaving

the Venezuelan populations without a proper name to be applied. Leptodactylus ocellatus Girard, 1853, a previous

name by which many Venezuelan populations were known, is now under synonymy of L. /atrans (Steffen, 1815), a

southern South American species. A thorough review of the /atrans species group is needed. Heyer (2014) indicates the

Venezuelan examples in his work are a different new species, compared with L. /atrans sensu lato. In the meantime,

Leptodactylus aff. macrosternum is used for the Venezuelan populations to be consistent with the classic literature. In

the /atrans species group of De Sa et al. (2014).

Selected references: Barrio-Amoros et al. 2011b; De Sa et al. 2014; Dixon and Staton 1976; Gorzula and Sefiaris

1998; Heyer 2014; Hoogmoed and Gorzula 1979; La Marca 1992; Rada 1981; Ramo and Busto 1989, 1990; Rivero

1967c; Staton and Dixon 1977; Tarano 2010.

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Family Strabomantidae

Pristimantis sp. |

Distribution: Paramo Los Granates, Mérida state.

Remarks: The Andes harbor a surprising diversity of high mountain Pristimantis, all very similar in shape. Some

preliminary molecular work (Barrio-Amoros et al. 2013) determined the validity of several candidate species.

References: Barrio-Amoros et al. 2013.

Pristimantis sp. 2

Distribution: La Motus, subparamo in Mérida Andes.

Remarks: A second new species, allied to P. /ancinii, but different in some morphological and molecular traits,

awaits proper attention.

References: Barrio-Amoros et al. 2013.

Pristimantis sp. 3

Distribution: Pifiango, Mérida state.

Remarks: Barrio-Amoros et al. (2013) show this species as a close ally of P. bricenii.

References: Barrio-Amoros et al. 2013.

Pristimantis sp. 4

Distribution: Murisipan-tepui, 05°53’N, 62°04’ W, 2,350 m asl, Estado Bolivar (MHNLS 11383).

Remarks: This species, collected by S. Gorzula, awaits a proper description (CBA, in prep.).

Pristimantis sp. 5

Distribution: Cerro de la Neblina, Amazonas state; camps I, II, and VII.

Remarks: This species corresponds to Eleutherodactylus “bromeliad” in McDiarmid and Paolillo (1988).

Pristimantis sp. 6

Distribution: Cerro de la Neblina, Amazonas state; camps I, II, X, and XI.

Remarks: This species corresponds to Eleutherodactylus “bromlike” in McDiarmid and Paolillo (1988).

Pristimantis sp. 7

Distribution: Cerro de la Neblina, Amazonas state; camps VII and XI.

Remarks: This species corresponds to Eleutherodactylus “stream” in McDiarmid and Paolillo (1988).

Pristimantis sp. 8

Distribution: Cerro de la Neblina, Amazonas state; camps VII, XI.

Remarks: This species corresponds to Eleutherodactylus “violet” in McDiarmid and Paolillo (1988).

Family Plethodontidae

Bolitoglossa cf. altamazonica (Cope, 1874)

Syntypes: ANSP or USNM, now lost or destroyed.

Terra typica: “Nauta,” Departamento Loreto, Peru.

Distribution: Valle del rio Doradas.

Remarks: Barrio-Amoros et al. (2015), when reporting B. /eandrae for Venezuela, state the presence of B. altamazonica

must be corroborated. The only specimen of this species reported, a juvenile (ULABG 3392), cannot be positively

identified and possibly represents B. Jeandrae. However, until more data is available, the possibility of a widespread

Amazonian species in the Doradas River valley cannot be ruled out, as the area is an important locality for upper

Amazonian herpetofauna (Barrio et al. 1999; Barrio 1999a, 2001; Barrio-Amoros et al. 2002, 2003; Chacon et al. 2002:

Barrio-Amoros and Chacon 2004; Barrio-Amoros and Diaz de Pascual 2008).

References: Barrio-Amoros 2004; Barrio-Amoros et al. 2015; Schargel and Rivas 2003.

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Appendix 2. Taxonomic list of the Amphibians of Venezuela.

In the past 15 years, amphibian taxonomy has undergone many changes. As a living science, many genetic studies

on previously unknown species or groups are still being performed, and the taxonomic panorama is being modified

continuously. While it is not imperative to follow the latest papers on each taxon, here the most recent taxonomic

reviews based on genetic and morphological data are presented, and specific points of disagreement are explained. The

following sources are used for the taxa indicated: Faivovich et al. (2005) and Duellman et al. (2016) for Arboranae;

Faivovich et al. (2010) and Duellman et al. (2016) for Phyllomedusidae; Faivovich et al. (2012) for Pleurodema,

and Faivovich et al. (2014), for Ceratophrydae; Frost et al. (2006) for Amphibia in general (with exceptions); Grant

et al. (2006, 2017) and Santos et al. (2009) for Dendrobatidae; Guayasamin et al. (2009) for Allophrynidae and

Centrolenidae; Castroviejo-Fisher et al. (2015) for Hemiphractidae; Hedges et al. (2008) and Heinicke et al. (2009,

2018) for Terraranae; Jungfer et al. (2013) for Osteocephalus and Tepuihyla; De Sa et al. (2012, 2014) for Microhylidae

and Leptodactylus; and Blackburn and Wake (2011) for higher amphibian taxa. Many of these works are in conflict,

and decisions are presented here on which is the most appropriate for each taxon. This comprehensive list also shows

the species known to exist in Venezuela but not yet described, and under our subjective view taxa that have been

incorrectly identified (see Appendix 1).

* Species with an asterisk are endemic to Venezuela.

CLASS AMPHIBIA Gray, 1825

Order ANURA Fischer von Waldheim, 1813

Family Allophrynidae Goin, Goin and Zug, 1978

Genus Allophryne Gaige, 1926

I. Allophryne ruthveni Gaige, 1926

Family Bufonidae Gray, 1825

Genus Amazophrynella Fouquet, Recoder, Teixeira, Cassimiro, Amaro, Camacho, Damasceno, Carnaval, Moritz

and Rodrigues, 2012

2. Amazophrynella minuta (Melin, 1941)

Genus Afelopus Dumeril and Bribon, 1841

Atelopus carbonerensis Rivero, 1972*

Atelopus chrysocorallus La Marca, 1996*

Atelopus cruciger (Lichtenstein and Martens, 1856)*

Atelopus mucubajiensis Rivero, 1974*

Atelopus oxyrhynchus Boulenger, 1903*

Atelopus pinangoi Rivero, 1980*

Atelopus sorianoi La Marca, 1983*

0. Atelopus tamaense La Marca, Garcia-Perez and Renjifo, 1990

11. Atelopus vogli Miller, 1934*

12. Atelopus sp.1

13. Atelopus sp. 2

mo MN DUA W

Genus Metaphryniscus Sefiaris, Ayarzagutiena and Gorzula, 1994

14. Metaphryniscus sosae Sefiaris, Ayarzaguena and Gorzula, 1994*

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Genus Oreophrynella Boulenger, 1895

15. Oreophrynella cryptica Sefiaris, 1995*

16. Oreophrynella huberi Diego-Aransay and Gorzula, 1988*

17. Oreophrynella macconelli Boulenger, 1900

1&8. Oreophrynella nigra Sefiaris, Ayarzagtiena and Gorzula, 1994*

19. Oreophrynella quelchii (Boulenger, 1895)

20. Oreophrynella vasquezi Sefiaris, Ayarzagtiena and Gorzula, 1994*

Genus Rhaebo Cope, 1862

21. Rhaebo glaberrimus (Gunther, 1868)

22. Rhaebo guttatus Schneider, 1799

23. Rhaebo haematiticus Cope, 1862

24. Rhaebo nasicus (Werner, 1903)

Genus Rhinella Fitzinger, 1826

25. Rhinella ceratophrys (Boulenger, 1882)

26. Rhinella horribilis (Wiegmann, 1833)

27. Rhinella humboldti (Gallardo, 1965)

28. Rhinella margaritifera (Laurenti, 1758)

29. Rhinella marina (Linnaeus, 1758)

30. Rhinella merianae (Gallardo, 1965)

31. Rhinella nattereri (Bokermann, 1967)

32. Rhinella cf. proboscidea (Spix, 1824)

33. Rhinella sclerocephala (Mijares-Urrutia and Arends, 2001)*

34. Rhinella sternosignata (Gunther, 1859)

Family Centrolenidae Taylor, 1951

Subfamily Centroleninae Taylor, 1951

Genus Centrolene Jiménez de la Espada, 1872

35. Centrolene altitudinalis (Rivero, 1968)*

36. Centrolene daidalea (Ruiz-Carranza and Lynch, 1991)

37. Centrolene notosticta Ruiz-Carranza and Lynch, 1991

38. Centrolene venezuelensis (Rivero, 1968)*

Genus Cochranella Taylor, 19511

39. “Cochranella” duidaeana (Ayarzagiiena, 1992)*

40. “Cochranella” riveroi (Ayarzaguena, 1992)*

41. “Cochranella” sp. |

Genus Espadarana Guayasamin, Castroviejo-Fischer, Trueb, Ayarzagtiena, Rada and Vila, 2009

42. Espadarana andina (Rivero, 1968)

Genus Vitreorana Guayasamin, Castroviejo-Fischer, Trueb, Ayarzagutena, Rada and Vila, 2009

43. Vitreorana antisthenesi (Goin, 1963)*

44. Vitreorana castroviejoi (Ayarzagtiena and Sefiaris, 1997)*

45. Vitreorana gorzulae (Ayarzagiiena, 1992)

46. Vitreorana helenae (Ayarzaguena, 1992)

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Subfamily Hyalinobatrachinae Guayasamin, Castroviejo-Fischer, Trueb, Ayarzagiiena, Rada and Vila, 2009

Genus Celsiella Guayasamin, Castroviejo-Fischer, Trueb, Ayarzagtiena, Rada and Vila, 2009

47. Celsiella revocata (Rivero, 1985)*

48. Celsiella vozmedianoi (Ayarzagtiena and Sefiaris, 1997)*

Genus Hyalinobatrachium Ruiz-Carranza and Lynch, 1991

49. Hyalinobatrachium cappellei van Lidth de Jeude, 1904

50. Hyalinobatrachium duranti (Rivero, 1985)*

51. Hyalinobatrachium fragile (Rivero, 1985)*

52. Hyalinobatrachium guairarepanense Sefiaris, 1999*

53. Hyalinobatrachium iaspidiense (Ayarzaguena, 1992)*

54. Hyalinobatrachium mesai Barrio-Amoros and Brewer-Carias, 2008*

55. Hyalinobatrachium mondolfii Ayarzagiena and Sefiaris, 2001*

56. Hyalinobatrachium orientale (Rivero, 1985)*

57. Hyalinobatrachium orocostale (Rivero, 1968)*

58. Hyalinobatrachium pallidum (Rivero, 1985)*

59. Hyalinobatrachium tatayoi (Castroviejo-Fisher, Ayarzagtiena and Vila, 2007)*

60. Hyalinobatrachium taylori (Goin, 1968)

Family Ceratophrydae Tschudi, 1838

Genus Ceratophrys Wied-Neuwied, 1824

61. Ceratophrys calcarata Boulenger, 1890

Family Ceuthomantidae Heinicke, Duellman, Trueb, Means, MacCulloch and Hedges, 2009

Genus Ceuthomantis Heinicke, Duellman, Trueb, Means, MacCulloch and Hedges, 2009

62. Ceuthomantis aracamuni (Barrio-Amoros and Molina, 2006)*

63. Ceuthomantis cavernibardus (Myers et Donnelly, 1997)

64. Ceuthomantis duellmani Barrio-Amoros, 2010*

Family Craugastoridae Hedges, Duellman and Heinicke, 2008

Genus Tachiramantis Heinicke, Barrio-Amoros and Hedges, 2015

65. Tachiramantis lentiginosus (Rivero, 1984)*

66. Tachiramantis prolixodiscus Lynch, 1978

Family Dendrobatidae Cope, 1865

Subfamily Aromobatinae Grant, Frost, Caldwell, Gagliardo, Haddad,

Kok, Means, Noonan, Schargel and Wheeler, 2006

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Genus Prostherapis Cope, 1868

67. Prostherapis dunni Rivero, 1961*

Genus Allobates Zimmermann and Zimmermann, 1988

68. Allobates algorei Barrio-Amoros and Santos, 2009*

69. Allobates bromelicola (Test, 1956)*

70. Allobates caribe (Barrio-Amoros, Rivas and Kaiser, 2006)*

71. Allobates femoralis (Boulenger, 1884)

72. Allobates humilis (Rivero, 1980)*

73. Allobates mandelorum (Schmidt, 1932)*

74. Allobates aff. marchesianus (Melin, 1941)

75. Allobates pittieri (La Marca, Manzanilla and Miyares-Urrutia, 2004)*

76. Allobates sanmartini (Rivero, Langone and Prigioni, 1986)*

77. Allobates undulatus (Myers and Donnelly, 2001)*

Genus Anomaloglossus Grant, Frost, Caldwell, Gagliardo, Haddad, Kok, Means, Noonan, Schargel and Wheeler, 2006

78. Anomaloglossus ayarzaguenai (La Marca, 1997)*

79. Anomaloglossus breweri (Barrio-Amoros, 2006)*

80. Anomaloglossus guanayensis (La Marca, 1997)*

81. Anomaloglossus moffetti Barrio-Amoros and Brewer-Carias, 2008*

8&2. Anomaloglossus murisipanensis (La Marca, 1997)*

83. Anomaloglossus parimae (La Marca, 1997)*

84. Anomaloglossus parkerae (Meinhardt and Parmalee, 1996)*

85. Anomaloglossus praderioi (La Marca, 1997)*

86. Anomaloglossus roraima (La Marca, 1997)*

87. Anomaloglossus rufulus (Gorzula, 1990)*

88. Anomaloglossus shrevei (Rivero, 1961 )*

89. Anomaloglossus tamacuarensis (Myers and Donnelly, 1997)

90. Anomaloglossus tepuyensis (La Marca, 1997)*

91. Anomaloglossus triunfo (Barrio-Amoros, Fuentes and Rivas, 2004)*

92. Anomaloglossus verveeksnyderorum Barrio-Amoros, Santos and Jovanovic 2009*

93. Anomaloglossus wothuja (Barrio-Amoros, Fuentes and Rivas, 2004)*

Genus Aromobates Myers, Paolillo and Daly, 1991

94. Aromobates alboguttatus (Boulenger, 1903)*

95. Aromobates cannatellai Barrio-Amoros and Santos, 2012*

96. Aromobates capurinensis (Péfaur, 1993)*

97. Aromobates duranti (Péfaur, 1985)*

98. Aromobates ericksonae Barrio-Amoros and Santos, 2012*

99. Aromobates haydeeae (Rivero, 1978)*

100.Aromobates inflexus (Rivero, 1978)*

101.Aromobates leopardalis (Rivero, 1978)*

102.Aromobates mayorgai (Rivero, 1980)*

103.Aromobates meridensis (Dole and Durant, 1973)*

104. Aromobates molinarii (La Marca, 1985)*

105. Aromobates nocturnus Myers, Paolillo, and Daly, 1991*

106. Aromobates ornatissimus Barrio-Amoros, Rivero and Santos, 2011*

107.Aromobates orostoma (Rivero, 1978)*

108.Aromobates saltuensis (Rivero, 1980)*

109.Aromobates serranus (Péfaur, 1985)*

110. Aromobates tokuko Rojas-Runjaic, Infante and Barrio-Amoros, 2011*

111. Aromobates walterarpi (La Marca et Otero, 2012)*

112. Aromobates zippeli Barrio-Amoros and Santos, 2012*

113. Aromobates sp. 1

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Genus Mannophryne La Marca, 1992

114. Mannophryne caquetio Mijares-Urrutia and Arends-R., 1999*

115. Mannophryne collaris (Boulenger, 1912)*

116. Mannophryne cordilleriana La Marca, 1995*

117. Mannophryne herminae (Boettger, 1893)*

118. Mannophryne lamarcai Mijares-Urrutia and Arends-R., 1999*

119. Mannophryne larandina (Yustiz, 1991)*

120. Mannophryne leonardoi Manzanilla, La Marca, Jowers, Sanchez and Garcia-Paris, 2007*

121. Mannophryne molinai Rojas-Runjaic, Matta-Pereira and La Marca, 2018*

122. Mannophryne neblina (Test, 1956)*

123. Mannophryne oblitterata (Rivero, 1984)*

124. Mannophryne orellana Barrio-Amoros, Santos and Molina, 2010*

125. Mannophryne riveroi (Donoso-Barros, 1965)*

126. Mannophryne speeri La Marca, 2009*

127. Mannophryne trujillensis Vargas and La Marca, 2007*

128. Mannophryne urticans Barrio-Amoros, Santos and Molina, 2010*

129. Mannophryne venezuelensis Manzanilla, Jowers, La Marca and Garcia-Paris, 2007*

130. Mannophryne vulcano Barrio-Amoros, Santos and Molina, 2010*

131.Mannophryne yustizi (La Marca, 1989)*

132. Mannophryne sp. |

133. Mannophryne sp. 2

Subfamily Dendrobatinae Cope, 1865

Genus Ameerega Bauer, 1986

134. Ameerega picta (Tschudi, 1838)

135.Ameerega trivittata (Spix, 1824)

Genus Dendrobates Wagler, 1830

136. Dendrobates leucomelas Steindachner, 1864

Genus Minyobates Myers, 1997

137. Minyobates steyermarki Rivero, 1971*

Family Eleutherodactylidae Lutz, 1954

Subfamily Eleutherodactylinae Lutz, 1954

Genus Eleutherodactylus Dumeril and Bribon, 1841

138. Eleutherodactylus johnstonei Barbour, 1914

Subfamily Phyzelaphryninae Hedges, Duellman and Heinicke, 2008

Genus Adelophryne Hoogmoed and Lescure, 1984

139.Adelophryne gutturosa Hoogmoed and Lescure, 1984

Family Hemiphractidae Peters, 1862

Genus Cryptobatrachus Ruthven, 1916

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140.Cryptobatrachus remotus Infante, Rojas-Ruijac and Barrio-Amoros, 2009*

Genus Flectonotus Miranda-Ribeiro, 1920

141. Flectonotus fitzgeraldi (Parker, 1933)

142. Flectonotus pygmaeus (Boettger, 1893)

Genus Gastrotheca Fitzinger, 1843

143.Gastrotheca helenae Dunn, 1944

144. Gastrotheca nicefori Gaige, 1933

145.Gastrotheca ovifera Lichtenstein and Weinland, 1854*

146.Gastrotheca walkeri Duellman, 1980*

147.Gastrotheca williamsoni Gaige, 1922*

148. Gastrotheca yacambuensis Yustiz, 1976*

Genus Stefania Rivero, 1968

149. Stefania breweri Barrio-Amoros and Fuentes, 2003*

150.Stefania ginesi Rivero, 1968*

151.Stefania goini Rivero, 1968*

152. Stefania marahuaquensis (Rivero, 1961)*

153.Stefania oculosa Sefiaris, Ayarzaguena and Gorzula, 1997*

154. Stefania percristata Sefiaris, Ayarzaguena and Gorzula, 1997*

155.Stefania riae Duellman and Hoogmoed, 1984*

156. Stefania riveroi Sefiaris, Ayarzaguena and Gorzula, 1997*

157. Stefania satelles Sefiaris, Ayarzagiena and Gorzula, 1997*

158.Stefania scalae Rivero, 1970*

159. Stefania schuberti Sefiaris, Ayarzaguena and Gorzula, 1997*

160. Stefania tamacuarina Myers and Donnelly, 1997

Family Hylidae Rafinesque,1815

Subfamily Cophomantinae Hoffmann, 1878

Genus Ayloscirtus Peters, 1882

161. Hyloscirtus jahni (Rivero, 1961)*

162. Hyloscirtus japreria Rojas-Runjaic, Infante-Rivero, Salerno and Meza-Joya, 2018

163. Hyloscirtus lascinius (Rivero, 1969)*

164. Hyloscirtus platydactylus (Boulenger, 1905)*

Genus Boana Wagler, 1830

165. Boana alemani (Rivero, 1964)*

166. Boana benitezi (Rivero, 1961)*

167. Boana boans (Linnaeus, 1758)

168. Boana calcarata (Troschel, 1848)

169. Boana cinerascens (Spix, 1824)

170.Boana geographica (Spix, 1824)

171.Boana hobbsi (Cochran and Goin, 1970)

172. Boana jimenezi Sefiaris and Ayarzaguiena, 2006*

173.Boana lanciformis (Cope, 1870)

174. Boana lemai (Rivero, 1971)

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175. Boana multifasciata (Gunther, 1859)

176. Boana ornatissima (Noble, 1923)

177. Boana pugnax (Schmidt, 1857)

178. Boana punctata (Schneider, 1799)

179. Boana roraima Duellman and Hoogmoed, 1992

180. Boana rhythmica (Sefiaris and Ayarzaguena, 2002)*

1&1. Boana sibleszi (Rivero, 1971)

182. Boana tepuiana Barrio-Amoros and Brewer-Carias, 2008

183. Boana wavrini (Parker, 1936)

184. Boana xerophylla (Dumeéril and Bibron, 1841)

185. Boana sp. (cf. rufitela)

Genus Myersiohyla Faivovich, Haddad, Garcia, Frost, Campbell and Wheleer, 2005

186. Myersiohyla aromatica (Ayarzagtiena and Sefiaris, 1994)*

187. Myersiohyla chamaeleo Faivovich, McDiarmid and Myers, 2013*

188. Myersiohyla inparquesi (Ayarzagtiena and Sefiaris, 1994)*

189. Myersiohyla loveridgei (Rivero, 1961)*

190. Myersiohyla neblinaria Faivovich, McDiarmid and Myers, 2013*

Subfamily Dendropsophinae Fitzinger, 1843

Genus Dendropsophus Fitzinger, 1843

191. Dendropsophus amicorum (Mijares-Urrutia, 1998)*

192. Dendropsophus battersbyi (Rivero, 1961)*

193. Dendropsophus luteoocellatus (Roux, 1927)*

194. Dendropsophus marmoratus (Laurenti, 1768)

195. Dendropsophus meridensis (Rivero, 1961)*

196. Dendropsophus microcephalus (Cope, 1886)

197. Dendropsophus minusculus (Rivero, 1971)

198. Dendropsophus aff. minutus (Peters, 1872)

199. Dendropsophus parviceps (Boulenger, 1882)

200. Dendropsophus pelidnus (Duellman, 1989)

201.Dendropsophus sarayacuensis (Shreve, 1935)

202. Dendropsophus yaracuyanus (Mijares-Urrutia and Rivero, 2000)*

Subfamily Lophyohylinae Miranda-Ribeiro, 1926

Genus Aparasphenodon Miranda-Ribeiro, 1920

203. Aparasphenodon venezolanus (Mertens, 1950)

Genus Osteocephalus Steindachner, 1862

204. Osteocephalus helenae (Ruthven, 1919)

205. Osteocephalus leprieurii (Dumeéril and Bibron, 1841)

206. Osteocephalus aff. planiceps (Cal)

207. Osteocephalus taurinus Steindachner, 1 862

208. Osteocephalus aff. taurinus (sp 1; Ca2)

209. Osteocephalus aff. taurinus (sp 2; Ca3)

Genus Phytotriades Jowers, Downie and Cohen, 2008

210.Phytotriades auratus (Boulenger, 1917)

Genus Tepuihyla Ayarzagiiena, Sefiaris and Gorzula, 1993

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Barrio-Amoros et al.

211. Tepuihyla aecii (Ayarzagtiena, Sefiaris and Gorzula, 1992)*

212. Tepuihyla edelcae (Ayarzagiiena, Sefiaris and Gorzula, 1992)*

213. Tepuihyla exophthalma (Smith and Noonan, 2001)

214. Tepuihyla luteolabris (Ayarzaguena, Sefiaris and Gorzula, 1992)*

215. Tepuihyla obscura (Kok, Ratz, Tegelaar, Aubret and Means, 2015)*

216. Tepuihyla rodriguezi (Ayarzagtena, Sefiaris and Gorzula, 1992)

Genus Trachycephalus Fitzinger, 1843

217. Trachycephalus resinifictrix (Goeldi, 1907)

218. Trachycephalus typhonius (Laurenti, 1768)

Subfamily Pseudinae Fitzinger, 1843

Genus Pseudis Wagler, 1830

219. Pseudis paradoxa (Linnaeus, 1758)

Genus Scarthyla Duellman et de Sa, 1988

220.Scarthyla vigilans (Solano, 1971)

Subfamily Scinaxinae Duellman, Marion and Hedges, 2016

Genus Scinax Wagler, 1830

221.Scinax baumgardneri (Rivero, 1961)*

222.Scinax boesemani (Goin, 1966)

223.Scinax danae (Duellman, 1986)*

224.Scinax exiguus (Duellman, 1986)*

225.Scinax fuscomarginatus (Lutz, 1925)

226.Scinax garbei (Miranda-Ribeiro, 1926)

227.Scinax kennedyi (Pyburn, 1973)

228.Scinax manriquei Barrio-Amoros, Orellana and Chacon, 2004

229.Scinax nebulosus (Spix, 1824)

230.Scinax rostratus (Peters, 1863)

231.Scinax ruber (Laurenti, 1768)

232.Scinax wandae (Pyburn and Fouquette, 1971)

233.Scinax x-signatus (Spix, 1824)

Genus Sphaenorhynchus Tschudi, 1838

234.Sphaenorhynchus lacteus Daudin, 1802

Family Leptodactylidae Werner, 1896

Subfamily Leiuperinae Bonaparte, 1850

Genus Engystomops Jiménez de la Espada, 1872

235. Engystomops pustulosus (Cope, 1864)

Genus Physalaemus Fitzinger, 1826

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236.Physalaemus cuvieri (Fitzinger, 1826)

237.Physalaemus ephippifer (Steindachner, 1864)

238.Physalaemus fisheri (Boulenger, 1890)

Genus Pleurodema Tschudi, 1838

239. Pleurodema brachyops (Cope, 1869)

Genus Pseudopaludicola Miranda-Ribeiro, 1926

240. Pseudopaludicola boliviana Parker, 1927

241.Pseudopaludicola llanera Lynch, 1989

242. Pseudopaludicola pusilla (Ruthven, 1916)

Subfamily Leptodactylinae Werner, 1896

Genus Adenomera Fitzinger, 1867

243.Adenomera andreae (Miller, 1923)

244. Adenomera hylaedactyla (Cope, 1868)

245. Adenomera cf. simonstuarti (Angulo and Icochea, 2010)

246. Adenomera sp. |

Genus Leptodactylus Fitzinger, 1826

247. Leptodactylus bolivianus Boulenger, 1898

248. Leptodactylus colombiensis Heyer, 1994

249. Leptodactylus diedrus Heyer, 1994

250. Leptodactylus fragilis (Brocchi, 1877)

251. Leptodactylus fuscus (Schneider, 1799)

252.Leptodactylus guianensis Heyer and de Sa, 2011

253.Leptodactylus insularum Barbour, 1906

254. Leptodactylus knudseni Heyer, 1972

255. Leptodactylus leptodactyloides (Andersson, 1945)

256.Leptodactylus lithonaetes Heyer, 1996

257.Leptodactylus longirostris Boulenger, 1882

258.Leptodactylus aff. macrosternum Miranda-Ribeiro, 1926

259. Leptodactylus magistris Mijares-Urrutia, 1997*

260. Leptodactylus mystaceus (Spix, 1824)

261.Leptodactylus pentadactylus (Laurenti, 1768)

262. Leptodactylus petersii (Steindachner, 1864)

263.Leptodactylus poecilochilus (Cope, 1862)

264. Leptodactylus rhodomystax Boulenger, 1884

265. Leptodactylus riveroi Heyer and Pyburn, 1983

266. Leptodactylus rugosus Noble, 1923

267.Leptodactylus sabanensis Heyer, 1994

268. Leptodactylus turimiquensis Heyer, 2005*

269. Leptodactylus validus Garman, 1888

270.Leptodactylus sp. 1

Genus Lithodytes Fitzinger, 1843

271.Lithodytes lineatus (Schneider, 1799)

Family Microhylidae Giinther, 1858

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Genus Adelastes Zweifel, 1986

272. Adelastes hylonomos Zweifel, 1986*

Subfamily Gastrophryninae Fitzinger, 1843

Genus Chiasmocleis Méhely, 1904

273.Chiasmocleis hudsoni Parker, 1940

Genus Ctenophryne Mocquard, 1904

274.Ctenophryne geayi Mocquard, 1904

Genus Elachistocleis Parker, 1927

275.Elachistocleis ovalis (Schneider, 1799)

276. Elachistocleis pearsei (Ruthven, 1914)

277.Elachistocleis surinamensis (Daudin, 1802)

Genus Hamptophryne Carvalho, 1954

278.Hamptophryne boliviana (Parker, 1927)

Subfamily Otophryninae Wasserssug and Pyburn, 1987

Genus Otophryne Boulenger in Lankaster, 1900

279. Otophryne pyburni Campbell and Clarke, 1998

280. Otophryne robusta Boulenger, 1900

281.Otophryne steyermarki Rivero, 1968

Genus Synapturanus Carvalho, 1954

282.Synapturanus mirandaribeiroi Nelson and Lescure, 1975

283.Synapturanus salseri Pyburn, 1975

Family Phyllomedusidae Giinther 1859

Genus Agalychnis Cope, 1864

284. Agalychnis medinae (Funkhouser, 1962)*

Genus Callimedusa Duellman, Marion and Hedges, 2016

285.Callimedusa tomopterna (Cope, 1868)

Genus Pithecopus Cope, 1866

286. Pithecopus hypochondrialis (Daudin, 1802)

Genus Phyllomedusa Wagler, 1830

287. Phyllomedusa bicolor (Boddaert, 1772)

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288.Phyllomedusa neildi Barrio-Amoros, 2006*

289. Phyllomedusa tarsius (Cope, 1868)

290. Phyllomedusa trinitatis Mertens, 1926

291.Phyllomedusa vaillanti Boulenger, 1882

292. Phyllomedusa venusta Duellman and Trueb, 1971

Family Pipidae Gray, 1825

Genus Pipa Laurenti, 1768

293. Pipa arrabali \zecksohn, 1976

294. Pipa parva Ruthven and Gaige, 1923

295.Pipa pipa (Linnaeus, 1758)

Family Ranidae Rafinesque-Schmaltz, 1814

Genus Lithobates Fitzinger, 1843

296. Lithobates catesbeianus (Shaw, 1802)

297.Lithobates palmipes (Spix, 1824)

Family Strabomantidae Hedges, Duellman and Heinicke, 2008

Incertae sedis

Genus Dischidodactylus Lynch, 1979

298. Dischidodactylus colonnelloi Ayarzaguena, 1985*

299. Dischidodactylus duidensis (Rivero, 1968)*

Subfamily Pristimantinae Ohler and Dubois, 2012

Genus Pristimantis Jiménez de la Espada, 1871

300. Pristimantis abakapa Rojas-Runjaic, Salerno, Sefiaris and Pauly, 2013*

301.Pristimantis ameliae Barrio-Amoros, 2011*

302. Pristimantis anolirex (Lynch, 1983)

303.Pristimantis anotis (Walker and Test, 1955)*

304. Pristimantis aureoventris Kok, Means and Bossuyt, 2011

305.Pristimantis auricarens (Myers and Donnelly, 2008)*

306. Pristimantis avius (Myers and Donnelly, 1997)*

307.Pristimantis bicumulus (Peters, 1864)*

308.Pristimantis boconoensis (Rivero and Mayorga, 1973)*

309. Pristimantis briceni (Boulenger, 1903)*

310.Pristimantis cantitans (Myers and Donnelly, 1996)*

311. Pristimantis colostichos (La Marca and Smith, 1982)*

312.Pristimantis conservatio Barrio-Amoros, Heinicke and Hedges, 2013*

313.Pristimantis culatensis (La Marca, 2007)*

314. Pristimantis fasciatus Barrio-Amoros, Rojas-Runjaic and Infante, 2007*

315.Pristimantis flabellidiscus (La Marca, 2007)*

316. Pristrimantis geminus Kaiser, Barrio-Amoros, Rivas-Fuenmayor, Steiletn and Schmidt, 2015*

317.Pristimantis ginesi (Rivero, 1964)*

318.Pristimantis gryllus Barrio-Amoros, Guayasamin and Hedges, 2012

319. Pristimantis guaiquinimensis (Schliter and Rédder, 2007)*

320. Pristimantis hoogmoedi Kaiser, Barrio-Amoros, Rivas-Fuenmayor, Steilein and Schmidt, 2015*

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Barrio-Amoros et al.

321.Pristimantis imthurni Kok, 2013*

322.Pristimantis incertus (Lutz, 1927)*

323.Pristimantis jabonensis (La Marca, 2007)*

324. Pristimantis jamescameroni Kok, 2013*

325.Pristimantis lancinii (Donoso-Barros, 1965)*

326.Pristimantis lassoalcalai Barrio-Amoros, Rojas-Runjaic and Barros, 2010*

327.Pristimantis longicorpus Kaiser, Barrio-Amoros, Rivas-Fuenmayor, Steilein and Schmidt,

2015*

328. Pristimantis marahuaka (Fuentes and Barrio-Amoros, 2004)*

329. Pristimantis marmoratus (Boulenger, 1900)

330.Pristimantis melanoproctus (Rivero, 1984)*

331.Pristimantis memorans (Myers and Donnelly, 1997)*

332.Pristimantis mondolfii (Rivero, 1984)*

333.Pristimantis muchimuk Barrio-Amoros, Mesa, Brewer-Carias and McDiarmid, 2010*

334. Pristimantis nicefori (Cochran and Goin, 1970)

335. Pristimantis nubisilva Kaiser, Barrio-Amoros, Rivas-Fuenmayor, Steilein and Schmidt,

2015*

336.Pristimantis paramerus (Rivero, 1984)*

337. Pristimantis pariagnomus Kaiser, Barrio-Amoros, Rivas-Fuenmayor, Steilein and

Schmidt, 2015*

338.Pristimantis pedimontanus (La Marca, 2004)*

339. Pristimantis pleurostriatus Rivero, 1984*

340. Pristimantis pruinatus Myers and Donnelly, 1996*

341.Pristimantis pulvinatus Rivero, 1984*

342.Pristimantis reticulatus Walker and Test, 1955*

343. Pristimantis rhigophilus (La Marca, 2007)*

344. Pristimantis rivasi Barrio-Amoros, Rojas-Runjaic and Barros, 2010*

345. Pristimantis riveroi Lynch and La Marca, 1993*

346. Pristimantis rozei Rivero, 1961*

347.Pristimantis sarisarinama Barrio-Amoros and Brewer-Carias, 2008*

348.Pristimantis stenodiscus (Walker and Test, 1955)*

349. Pristimantis telefericus La Marca, 2005*

350. Pristimantis thyellus (La Marca, 2007)*

351.Pristimantis tubernasus Rivero, 1984 *

352.Pristimantis turik Barrio-Amoros, Rojas-Runjaic and Infante, 2007*

353.Pristimantis turumiquirensis Rivero, 1961*

354. Pristimantis vanadisae La Marca, 1984*

355.Pristimantis vilarsi Melin, 1941

356. Pristimantis yaviensis Myers and Donnelly, 1996*

357. Pristimantis yukpa Barrio-Amoros, Rojas-Runjaic and Infante, 2007*

358. Pristimantis yuruaniensis Rodder and Jungfer, 2008*

359. Pristimantis yustizi Barrio-Amoros and Chacon, 2004*

360. Pristimantis zeuctotylus Lynch and Hoogmoed, 1977

361.Pristimantis sp. |

362. Pristimantis sp.

363.Pristimantis sp.

364. Pristimantis sp.

365. Pristimantis sp.

366. Pristimantis sp.

367. Pristimantis sp.

368. Pristimantis sp.

CONN FW WN

Subfamily Strabomantinae Hedges, Duellman and Heinicke, 2008

Genus Strabomantis Cope, 1862

Amphib. Reptile Conserv.

369. Strabomantis biporcatus (W. Peters, 1863)*

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Catalogue of the amphibians of Venezuela

Order URODELA Duméril, 1805

Family Plethodontidae Gray, 1850

Subfamily Hemidactylinae Hallowell, 1856

Genus Bolitoglossa Duméril, Bibron and Dumeril, 1854

370. Bolitoglossa cf. altamazonica (Cope, 1884)

371.Bolitoglossa borburata Trapido, 1942*

372. Bolitoglossa guaramacalensis Schargel, Garcia-Pérez and Smith, 2002*

373. Bolitoglossa leandrae Acevedo, Wake, Marquez, Silva, Franco and Amezquita, 2013

374. Bolitoglossa mucuyensis Garcia-Gutiérrez, Escalona, Mora, Diaz de Pascual and Fermin, 2013*

375. Bolitoglossa orestes Brame and Wake, 1962*

376. Bolitoglossa tamaense Acevedo, Wake, Marquez, Silva, Franco and Amézquita, 2013

Order GYMOPHIONA Miller, 1831

Family Caeciliaidae Rafinesque, 1814

Genus Caecilia Linnaeus, 1758

377. Caecilia flavopunctata Roze and Solano, 1963*

378.Caecilia subnigricans Dunn, 1942

379.Caecilia tentaculata Linnaeus, 1758

Family Rhinatrematidae Nussbaum, 1977

Genus Epicrionops Boulenger, 1883

380. Epicrionops niger (Dunn, 1942)

Family Siphonopidae Bonaparte, 1850

Genus Microcaecilia Taylor, 1968

381. Microcaecilia rabei (Roze and Solano, 1963)*

Genus Siphonops Wagler, 1830

382.Siphonops annulatus (Mikan, 1820)

Family Typhlonectidae Taylor, 1968

Genus Nectocaecilia Taylor, 1968

383. Nectocaecilia petersii (Boulenger, 1882)

Genus Potomotyphlus Taylor, 1968

384. Potomotyphlus kaupii (Berthold, 1858)

Genus 7yphlonectes Peters, 1879

385. Typhlonectes natans (Fisher, 1879)

386. Typhlonectes compressicauda Dumeril and Bibron, 1841

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Appendix 3. List of amphibian species of Venezuela by biogeographic areas. 1. Andes (including Cordillera de

Merida, Venezuelan part of Cordillera Oriental de Colombia, and Sierra de Perija); 2. Coastal Range (including eastern

and western sections); 3. Llanos; 4. Amazon lowlands; 5. Guiana shield (including lowlands, uplands, and highlands);

and 6. Maracaibo basin. An asterisk after name indicates species is endemic for Venezuela.

species SCSC—sS 2

[Allopinmeraveni ————SSCS~SSC SE XX

[Amacophrynella inna ————SSSCS~—SCSTS SE

[Atelopus carbonerensis® ———S—S—S~—S XT | dT Sd

[Atelopus chesocoralius® ———SSS—~—SCSCS XE

[Atelopus orwiger*———SSOS—S—SSC KE

[Atelopus mucibaiiensis® ———SSOSC~—SCSCSCSCSCSCSCX YE

[Atelopus axyrhynchus® SSX EET

[Atelopus pinangoi*———SSS~—~—S XP

[Atelopus soranoi® SSCS XP | dT dP

[Atelopustamaense——SSSCS~C—~—SCS XP

[Atlopusvogi® SSCS EE

laelopusp.l®™ SSOSCS—SC EE

[Aelopussp.® SSC XP

[Metaphymiscussosae® ——SSSOSC~SSC Es

[Oreophryneliacrppnca® ————SSSCS~S~—SSCSS Ed i

[Oreophryneliahuberi® ——SSSSCSC~—SCSC—SSSS SE

[Oreophrynetamacconeli SSCS Ei

[Oreophrynellanigra®—SSSCSC~SCSCSSS Es

[Oreophrynellaquech’ SSCS SE i

[Oreophrynetavasques® ———SSCSC~—SCSCSSS SE

[Rhacbo glaberrims ——S—SCS~—S SX TE dT i

[Rhacbo guans ——SSCSC~—SC~S~S~S~S~SCSsSsSsSSSCSE

[Rhacbo haemaniious ————SSSCSCS~—S SX TE

[Rhacbonasiews ———SSOSCSC~—CSCSCSCSCSCSCSSSS SE

[Rhinela bebei SSS EX TX

[Rhinelta margariifera————SOSCS~—S XK Td XY

[Rhineltamerianae———SSCS~—SSSCSESdTS XY

[Rhineta ch. proboscidea—————SSCS~—STC PX

[Rhinetascerocephata® ——SSCS~—SCS—CSCST RP

[Rhineta nernosignata————S—S—S—S~—S XX || dP

[Centrolene ahitudnaie® ———SSSC~C~SCS SX TP

[Centrolene daidalea ——SSCS~—SCS XT Ed

[Centrolene nowostica ———SSSCSC~C~SCS~—~SC~C~S~S~—~Ss~sX YE

[Centrolene venezuclemsis® ———SSSC~—~—S XE ET

[*Cootranelia” dhidaeana® ———SS—~—~SSSS ST

Cochrane rveroi® ———SSOSCS~SSSS SE i

[*Cootranetasp1* STE

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Catalogue of the amphibians of Venezuela

Appendix 3 (continued). List of amphibian species of Venezuela by biogeographic areas. 1. Andes (including

Cordillera de Mérida, Venezuelan part of Cordillera Oriental de Colombia, and Sierra de Perija); 2. Coastal Range

(including eastern and western sections); 3. Llanos; 4. Amazon lowlands; 5. Guiana shield (including lowlands,

uplands, and highlands); and 6. Maracaibo basin. An asterisk after name indicates species is endemic for Venezuela.

[Espadaranaandna ————SSSCS~S~S XY | dd

[Tieorana anisthenes® ————SSCS~SCS—STS KT

[Vireorana casroneioi® ————S—SCSCS~S i X | dT

[Vireoranagorsuiae ——S~—~—STS SE

[Vireorana helenae ———SSSCS~S~S CE dT i x

[Cetsetarevocana® ———SSOS—S—~—S EE

[Cetsictavormediano® —————SSCS~—S XT |

[Hyainobarrachium cappellee ————S~—S Yi i

[Hyalnobarrachium duran ————S—~—S X | | |

[Hyainobarrachiumfragie® ————S—S~—~—ST TE

[Hyalnobarrachium guairarepanensis® __———S~i Sid X | |

[Hyainobatrachiun tspidensis ——~—S~S~S TE Y

[Hyalnobarrachium mesai® ————SCS~S ET i x

[Hyainobatrachiun mondo ———S—S—S~STS TX

[Hyalnobarrachium orienate Si XT dT

[Hyainobatrachium orocostie® ———SSCS~S XT

[Hyalnobarrachium palitum® SCS X || | dd

[Hyainobatrachium tayo” ————SSCS~S XE EP

[Hyalnobarrachium wmiort SCS ET

[Ceranophnyscatcarata—————SCS~—SCSTS

[Ceuthomanisaracamui® —————S~—~S ET

[Ceuthomanis cwernbarus——S~S TE x

[Ceuthomanis duelinani* SCS | | dT i x

[Dischidodacruscolomello®™ ———S—S~S Ei xX

[Dischidodacryus duidensis# Ss | dT dT i x

[Pristimanisabakapa®———SSSCS~ST Ex

[Pristimanis anelae® ————S—S~—SS X | | |

[Pristimanis anoles ———SSOSCS~—SCSCS CX PE

[Pristimanis anots* SSCS dP XT dT

[Prstimanis areovents ————SSSCS~—SCSCS—STS SE

[Pristimanisaurcarens*———S—SCS~—S Yd i x

[Pristimanisavns® ———SSSOSCS—SST Ex

[Pristimanisbicumihus® ——————S—S~S XT dT

[Pristimanis Boconoensis® ———SS—~S~S CX EP

[Pristimanis bricen® ———SS~—S X | | dT i

[Prstimanis canntans®————SSOS—~—~—SCSSTS Ed

[Pristimanis colosichos*® ————S—SCS~S XP TdT

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Barrio-Amoros et al.

Appendix 3 (continued). List of amphibian species of Venezuela by biogeographic areas. 1. Andes (including

Cordillera de Mérida, Venezuelan part of Cordillera Oriental de Colombia, and Sierra de Perija); 2. Coastal Range

(including eastern and western sections); 3. Llanos; 4. Amazon lowlands; 5. Guiana shield (including lowlands,

uplands, and highlands); and 6. Maracaibo basin. An asterisk after name indicates species is endemic for Venezuela.

[Pristimanisgenims® S| dX YT dT dC

[Pristimantisgnesi® ——SSOS—~—S CX EP

[Pristimanis epttus SSS XP | dT

[Prstimanis guaiguininensis® —————SS~S~S TT i

[Pristimanis hoogmoedi®———~—S~S~ST TdT

[Pristimanis miurni® ————SSSCS~—S SE

[Pristimanis ncerus® —————S—SCS~—ST PX TT

[Prstimanisjamescameron® ——S~S ST

[Pristimanisassoateatai® ———S—S~—~S XK | | dT

[Pristimanistansni® ———S—S—S—~—S XP Pd

[Pristimanis tongicorpus® _————SSCS~SCSS ET

[Prstimanis maramaka®———S—S—~S SE

[Pristimantis marmoraus SCS Ei x

[Prstimanis melanoprocus———SSSS~S~—~—SCSC—CSCS SEE

[Pristimanis menorans———SCS~SS ET

[Pristimantis mondolfi———S—S—S—~—S XP | dT

[Pristimanis muchimuk® ST | | dT i

[Pristimanis ceo, ————S—SSCS~—~—S XP EE

[Pristimanis mbisiva® —————S—SCS~—SS XT dT

[Prstimanisparamerws® ——SSCS—~—SCS CX EE

[Pristimanispariagnoms® ————S—S~SS TT

[Prstimanispedimonams®——S—S—S—~S XE

[Pristimanis plewosrias* SCS X | | |

[Pristimanis pruinans*———SS—S~—SS

[Pristimanis puvinanws SCS TdT i x

[Pristimanisreiculams®* ———~—SS—~ST TE

[Pristimanisrhigophius® SS X | | | dP

[Pristimanisrvasi*=————SSCSCS~S XE

[Prisimanis rvero® ———S—S—S—S—S SC XT dT

[Pristimanisroei* ———SSOSC~S~—ST XT

[Pristimanis sarsarnama®_———S—~—S ET i

[Pristimanisstenodwcus® ———S—S~—ST XT

[Pristimanislefecus® ————SS—S—~S || dT Sd

[Pristimanis hyets® ———SS—S~—S XP | |

[Pristimanisbernasis————SSSCSCS~S XP Ed

[Pristimanis writs —————SSOSC~S~—S XP Pd

[Pristimanisturumigurensis® _—————S—SCS~SC TT

Pristimantis vanadisae*

Pristimantis yukpa*

Amphib. Reptile Conserv. 187 July 2019 | Volume 13 | Number 1 | e180

Catalogue of the amphibians of Venezuela

Appendix 3 (continued). List of amphibian species of Venezuela by biogeographic areas. 1. Andes (including

Cordillera de Mérida, Venezuelan part of Cordillera Oriental de Colombia, and Sierra de Perija); 2. Coastal Range

(including eastern and western sections); 3. Llanos; 4. Amazon lowlands; 5. Guiana shield (including lowlands,

uplands, and highlands); and 6. Maracaibo basin. An asterisk after name indicates species is endemic for Venezuela.

[Pristimanisyuruaniensis® SCS dT dT Sd dX

[Pristimanisyusic® ————SSCSCS~—SCS CX EP

[Pristimanisseutoplus SCS Pd XP

[Prisimanissp. 1? —SSSSCS~—S XE

[Pristimanissp.2® SSS X | | dT dd

[Pristimanissp.3@ SSOSOSCS~S XP

[Pristimanisp.4® SSS Ed

[Pristimanissp.5@ SSSCS~STS Ed x

[Pristimanis.6® SSS P| | Td x

[Pristimanissp. 77 SSS x

[Pristimanis sp. ——SSSOSCS—S | | | dx

[Tachiramantis enignosws ————SSCS~—S—~—SCS XE

[tachivamants protwodiscus SCX PE |

[Sirabomanisbiporeams* ———SC~S—STS TE

["Prostherapis” duis ————SSOS~—S XPT

[Allobaresaigorei*———SSSC~—SCS XE

[Allobares brometola® ————SSSCSC~SC XT

[Alloares carte? ———SOSCS~ST XT

[Allobatesfonorais _——S—S~S—S | | | dT i

[Allobares hamiis® ————SS—~—S XE dT

[Allobares mandelonm® ————SSCS—~SC TdT

[Allobates af. marchestamis————SS~ST SP

[Allobarespiiert®™ _————SSSSCS~—SCSCS XK TES

[Allobatessamarin® ———S—S—~—SSS SE

[Allobares undulanus® ————SSCS~SS ST dT i x

[Anomalogiossus quarzaguena® ———SSCS~SS Ex

[Anomalogiosus breveris SCS SY dT i x

[Anomalogiossus guanaensis® ——S—S—~—S SS Ex

[Anomalogiosus mogeni* SCS dT dT i x

[Anomalogossus mursipanensis® ———S—S—S~—S—S Px

[Anomalogiossusparmae®——~—S~S~S ET i x

[Anomalogiossusparkerae® ——S~S Td x

[Anomalogiossus praderoi SS dT dT i xX

[Anomalogiossus roraima ST Tx

[Anomalogiossus rgitus® SSCS | | dT i x

[Anomalogiosus shreve®@ ———SSOSCS~SCS SE x

[Anomalogiossus amacurensis __——S—S~S~S SS ET xX

Anomaloglossus tepuyensis*

Anomaloglossus wothuja*

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Appendix 3 (continued). List of amphibian species of Venezuela by biogeographic areas. 1. Andes (including

Cordillera de Mérida, Venezuelan part of Cordillera Oriental de Colombia, and Sierra de Perija); 2. Coastal Range

(including eastern and western sections); 3. Llanos; 4. Amazon lowlands; 5. Guiana shield (including lowlands,

uplands, and highlands); and 6. Maracaibo basin. An asterisk after name indicates species is endemic for Venezuela.

[Aromobuesalbogmams® SCS XY | i dC

[Aromobaescamaciia® ———S—S~—S XP Ed

[Aromobacescapurnensis® ————S—S~S~—S X | | |

[Aromobaesdurani® ——SSSCS~—SCSCS XE Pd

[Aromobaesericksonae® SSX PY dT

[Aromobaeshaydeeae® ——SSSCS~SCSCSCS CX EP dE

[Aromobaes nfleus* SSX | dT

[Aromobates leopards? —————SSOSCS~SCS CX YP

[Aromobaes mayorgai® SSX PET

[Aromobaes meridenss® ————SS—~S~S XP Ed

[Aromobaes molinari® ————SSC—S~S XE |

[Aromobates nocurms® SSX PP

[Aromobaes oranssims® ————SSOSCS~S XP | dT i

[Aromobates orostoma®——SSOSCS~—SCSCS XE

[Aromobaesserrams® ——SSSOSCS~S XP EE

[Aromobaessaluensis® —————S—~S XE Ed

[Aromobues ohuko® SS X | | | i

[Aromobaeswaterap® —————S—S~S XP Pd

[Aromobaescippeli® ————SSSOSCS~S XP P|

[Aromobaessp.1* SSS XP Ed

[Mamophome capeno® ————S—~—SS XT dT

[Mamophayne coarse ———SS~—S CX EE

[Manopheme corditerana® ———SS~—S X | | dT dd

[Mamophme herninae* ——SSS~S~—STS TE

[Mannophene tamarcai*———S—S~—S i XT dT

[Mamopheyne larandina® ———SSOSC~—~S—S—SCS XE

[Mannopheme leonardo ————SCS~SC TT

[Mamophemne molnai® ST X TT

[Manopheme neblina® _—————S~S i Xx | |

[Mamophye oblieraa® ————SSC~ST XT

[Mamophemne oretana® ————SS~—S CX TdT

[Mamopheymervero™———SSS—S~—~—STC EE

[Mamophemespeer® SSX | | dT Sd

[Mamophememyillewis® ————SS~—~S XP Pd

[Manopheme uticans* SSCS XP | dT

[Mamophynevenezuelonis® —————SS—S—~—SS PP

[Mannophemneviicano® —————SSCS~S SX TT

Ameerega picta

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Appendix 3 (continued). List of amphibian species of Venezuela by biogeographic areas. 1. Andes (including

Cordillera de Mérida, Venezuelan part of Cordillera Oriental de Colombia, and Sierra de Perija); 2. Coastal Range

(including eastern and western sections); 3. Llanos; 4. Amazon lowlands; 5. Guiana shield (including lowlands,

uplands, and highlands); and 6. Maracaibo basin. An asterisk after name indicates species is endemic for Venezuela.

lAneeregarnimaa—————S—S~—~—S| | dT Sd Sd XY

[Dendrobateslewomelas————SSSC~—SCSCSSS PX

[Minyobaressteyermarki# SSCS ET x

[Btewtherodactyusjomnsionsi ————SC~—S~ST TE

[Adelophnyne guturosa—_————S~S~S SE i

[Crprobarrachus remons® ————SS~—~—S PP dT

[Flecononsfizgeraids ———S—SCS~—SC ET

[Fletononspyemacns——SSOSC~—~—SCS XX |

[Gastrotheca helene ————SSOS—S—S X | | dT

[Gastrotheca nico ———SSCSCS~—SCSCSCSCS XP Pd

[Gastrotheca ovfera®———SSCS—~—~—SCSS PE

[Gastrothecawalker® ———SS—~—S XT

[Gastrotecawithanson® ————S—S~—S SX TdT

[Gastrothecaryacambuensi® ———S—~—S XE

[Stam breve® ——SSSSCS~—SS i

[Stmagnes® ———SSSOS—~—ST x

[Stetmiagom® —————SS—SCS—SSSSS | | | | dx

[Stefania marahuaguensis® ———SS~—S Ex

[Stetmiaocuosa® SSCS ET i x

[Stefania percnsaa®™ ————SSOSC~—SSS Ed x

[Stiimiariaes SSP | dT i x

[Stsamarvero® ——SSOSCS—S—ST Ed x

[Stefmiasaetes® SSS | | dT dx

[Stemi scale ————SS—S—S Ed x

[Stetmiaschibor™ ——SSOSCS~—SS TdT i x

[Stefania tamacuarna————SSCS~—SCST Ex

[Aparasphenodon venezolamis——SCS~S Pd

[Dendropsophus amicorm™ SS TT

[Dendropsophus barersbyi® ———S~—~S XPT

[Dendropsophustueoocelas ———S~S—SCSCSC XX ||

[Dendropsophus marmoraus ———S—S—~S SE i XP

[Dendropsophus meridenss® ———S—~—S EP dT

[Dendropsophus microcephahis_———S~S i SX |

[Dendropsophus minwsous——~—S~S EX TX

[Dendropsophus af. minus SS XX | XT

[Dendropsophusparviceps ———SSSC~—C~—~—sSs~SsSsSsSTSSSP dX

[Dendropsophuspeliams ———S—S~S XP | dT

Dendropsophus sarayacuensis

HAyloscirtus japreria

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Appendix 3 (continued). List of amphibian species of Venezuela by biogeographic areas. 1. Andes (including

Cordillera de Mérida, Venezuelan part of Cordillera Oriental de Colombia, and Sierra de Perija); 2. Coastal Range

(including eastern and western sections); 3. Llanos; 4. Amazon lowlands; 5. Guiana shield (including lowlands,

uplands, and highlands); and 6. Maracaibo basin. An asterisk after name indicates species is endemic for Venezuela.

[Hploscimstascniws ——————S—SCS~—SCS Xd dC

[Hifoscirus playdacpius————SSCS~SCS XP Pd

[Boanaatemani® SSCS TdT

[Boaabenei® —SSSCS~S~—ST SS

[Boanaboans SSCS XK EK XX

[Boanacalcaraa————SSCSC~SYS SE XX

[Boanacinerascens SSCS EX x

[Boana geographica——SOSCS~SCSSSS Ed XX

[Boanahobbsi———SSCSCS~S~SS Ed

[Boanajimen® ———SSOSC~—ST SE

[Boana lancifornis SSS PK EX XY

[Boanatemi——SSSOSCS~—SS

[Boana mitifscaa—————SCSCS~SCSCST Ed x

[Boana omatissma——~—S—S—S TE x

[BoanapugnasSSSCSCS~S~CSCS XP

[Boana punctata————SSSCS~—SCSCSCSCSYTS EX

[Boana rorama———S—S—S—SS| | | dT i x

[Boanariythmica® ——SSSCS~S~—ST Ed x

[Boanasibiess: SSCS Ed x

[Boanatepuiana———SSSCS~S Ed x

[Boanawarins SSCS EX

[Boanasp.(ef.rifighsy ————S—SC~S~—ST SX | |

[ijersiohytaaromanca® —————SS~S SE

[Mpersiohytachamacleo*———S~S~S | | dT i x

[iersiohyta pares” ———SSCS~ST Ex

[Mjersiohytaioveridgei® ————SSCS~STSS ST dT i x

[iersiohyla neblinana® ———SSSCS~ST Ex

[Osteovephatus helenae S| | | dT i x

[Oseocephatus eprewh ———SSCSCS~—STS TX

[Osteocephatus aft planieps (Ca) SCS S| dT dX

[Oseocephatustaurims ———SCS~S SS TX

[Oseocephalus at. taurus plc) ——————S~i S| dT dX

[Osteocephatus aft turimus(sp2:C8) ————S~S i PX

[Phytoriadesawans———SSSCSCSC~SCST XE

[Pseudisparadoxa————SSCS~SCSCSCSSS XE

[Seartilavigians ————SSOS—S—S XP XT x | PX

Scinax baumgardneri*

Scinax exiguus*

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Appendix 3 (continued). List of amphibian species of Venezuela by biogeographic areas. 1. Andes (including

Cordillera de Mérida, Venezuelan part of Cordillera Oriental de Colombia, and Sierra de Perija); 2. Coastal Range

(including eastern and western sections); 3. Llanos; 4. Amazon lowlands; 5. Guiana shield (including lowlands,

uplands, and highlands); and 6. Maracaibo basin. An asterisk after name indicates species is endemic for Venezuela.

[Seinax fscomarginanus SSCS YT dT dT Sd XY

[Scmaxgurber ——SSSCSCS—~—ST Ed

[Seinastemedi ——SSOSC—S—SS EX

[Seinax manriguel ———SSCS—~—~—CSCSCSCS XP

[Seinas nebulosus————SSSCSCS—SS Ed

[Seinax romans ———SSSOS—SS EK

[Seinax ruber—SSSCSCSS x EE

[ScmaxwandseSSSCS—~—S XP

[Scinarssignaus——SSCS~—SSC EX XP

[Sphanorlynciustactews———SSCS~—SC—SCSCSTSE

[Tepuitylaaec® SSCS Ed x

[Tepuiyaedetoae® ——SSOS~—SCST Tx

[Tepuiyla exophthaina —————SSCSCS~SCST EE i x

[Tepuiyaleolabrs® ——SSCS~—SCS Ed x

Tepuityaobseurat SSCS Ed x

[Tepuiyta rodriguess———SSCSC~—CSC~—CSCSCSSS Ed x

Trachyeephatus resents ———S—SCS~—S | dT dX xX

lAgalyehnis medinae®_————SSSCS~—SCSSSC EC

[Callinedusa tonoptema———SS—S~—ST EX

[Piyllomedusa bicolor SCS | | dX XT

[Piyllomedusaneitd® ———S—S—~—S TT

[Phyllomedusatarsins ————SCS~—SCS SX EX |

[Piyllomedusatrinnas ——S—S~STS |

[Piyllomedusavensta SSCS | | dT i

[Piyllomedusavailland ———~—SCS~SCSCST TX

[Puhecopus lypochondrias ___——S~S~S i XX PX]

[Physalaemus wien —————S—S—~—SS| TdT x

[Physalaemusephippifr SCS | | dT i x

[Physalaemu fishers ———SSSSCS~S XK TX XX]

[Plewodena brachyops————SSSCS~SSS XX

[Pseudopatudcola olviana———~—S—S~S Ed XT

[Pseudopatudcolaitanera SS EX

[Pseudopatudcolapusitla S| | dT

[Adenomeraandreae———SSSCSCS~—SCS Ed

[Adenomerahylacdacyta———S—SCS~S Ed XX

[Adenomeracf.sinonsnard———S—S~—S XT dT

[Adenomerasp.1 SSS dT dX

[Leptodaciylus bolwvamis—————SSCS~S~SCSS EX

[Leptodacilus colonbiensis _——S—S~S X | | | dC

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Appendix 3 (continued). List of amphibian species of Venezuela by biogeographic areas. 1. Andes (including

Cordillera de Mérida, Venezuelan part of Cordillera Oriental de Colombia, and Sierra de Perija); 2. Coastal Range

(including eastern and western sections); 3. Llanos; 4. Amazon lowlands; 5. Guiana shield (including lowlands,

uplands, and highlands); and 6. Maracaibo basin. An asterisk after name indicates species is endemic for Venezuela.

[Lepiodacphusdedrs ————SSSCS~—S Sd dC Sd XY XT

Leprodaciyusftagiis ——S—S~S~—~—ST ET

[Leptodaciyusfscus SSS EX XX

[Leprodaciyus guanensis ———SSC~—SCSSS EX

[Leptodacilus insular SSS XX

Leprodaciyus kmdseni ———SSC~—SS EX

Leptodaciylus leprodacrfoides——S—S~—SCS Ei x

[Leprodaciyustithonaetes————SSSCS~ST Ed XX

[Leptodaciustongirosiis SCS | | dT x

Leprodacilus aff. macrostemmim ————S—~S TX |

[Leptodaciyus magianis® ———SSSCSCS~SCS XT

Leprodaciyus mystacens———SSSCS~S~SCSCSCSCSYSS SE

[Leptodacilus pentadacnyus SCS Ed XX

Leprodaciyuspetesh_————~—SCS~SCSCSTS EX

[Leptodaciyus poecitochins _——S—S~S~S |

Leprodaciylusrhodomystax ———S—S~S~—SCS—ST SE

[Leptodacryusrveroi_—————SSOSCS—~ST Ed

[Leprodaciyusrugosus———SSSCS~—SCSCSCSTS

[Leptodaciussabanensis SCS | | dT i x

Leprodacilus urimiquensis———S—S~S~—~S TE

[Leptodaciyusvalius—————SOSCS~SS XXX

LLeprodacussp.1* —SSSOS—~—S XP | |

[Litodves ineatus————SSSCS~—S XP | dX

[Adelases yionomos———SSS—~—SCSTS EX

[Chiasmoctes hudson’ —————SSOSCS~SS Ed XP

[Gienophimegea —SSSCSCS~—SCSCSCST

[Blachisocteis vais ————SCS~S i XTX]

[Blachisocteispearsei ———SSCSC~—C—C~—~S~—~S~SsSsSsSTS ST

[Elachistocteis surinamensis_———S—SCS~S~S Ed XX]

[Hamprophryne boliina ————SCS~ST EX

[Orophiyne burs ———SSSCSC~SCST

[Otophiynerobusta————SSCSC~C~—~—SCSCSCSCSsSYSS

[Otophiyne seyermarki—————SSSCS~S Ed i x

[Synapturamus mirandariberoa———SSCS~S~S TE x

[Synapturams salves ———————SSSCSC~S~S Ed XX

[Pipaamaba ———SSCSC~—~SCSCST

[Pipapana SOS Ed

Bolitoglossa cf. altamazonica

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Appendix 3 (continued). List of amphibian species of Venezuela by biogeographic areas. 1. Andes (including

Cordillera de Mérida, Venezuelan part of Cordillera Oriental de Colombia, and Sierra de Perija); 2. Coastal Range

(including eastern and western sections); 3. Llanos; 4. Amazon lowlands; 5. Guiana shield (including lowlands,

uplands, and highlands); and 6. Maracaibo basin. An asterisk after name indicates species is endemic for Venezuela.

Bolitoglossa borburata*

Bolitoglossa guaramacalensis*

Bolitoglossa leandrae

Bolitoglossa mucuyensis*

Bolitoglossa orestes*

Bolitoglossa tamaense

Caecilia flavopunctata*

Caecilia subnigricans

Caecilia tentaculata

Epicrionops niger

[Nectocwecliapetersi———S—SCS~—SSCSdSSS dC

Potomotyphlus kaupii |e ili

iphionecres conpressecauda ————S—S—S~S—SSCSSEC XT

[iyphionectes naans ————SSSCSCS*~—C—C—CSsSsSs~SsSsSESSSSCY SC

Towlsy SS OSOSOSOSOSSSSOSOSCSCSCSCSCS ww GO nS

Appendix 4. Addition of species for Venezuela between 2009 and 2018 (not including those in Appendix 1).

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Appendix 4 (continued). Addition of species for Venezuela between 2009 and 2018 (not including those in

Appendix 1).

Appendix 5. Species deleted from previous list (Barrio-Amor6és 2009c) or reported after and deleted herein.

Rhinella granulosa (taxonomic review reveals that the

former subspecies must be elevated to full species rank Narvaes and Rodrigues 2009

and R. granulosa is not present in Venezuela)

Rhinella humboldti (confused with R. beebei, the latter

recovered from synonymy of R. humboldti), could be Murphy et al. 2017

present in the Maracaibo lake basin

Cochranella oyampiensis (confused with C. helenae) Sefiaris 1997; Kok and Castroviejo 2008

Hyalinobatrachium crurifasciatum (synonym of H. Castroviejo-Fisher et al. 2011

cappellei)

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Appendix 5 (continued). Species deleted from previous list (Barrio-Amorés 2009c) or reported after and deleted

herein.

Hyalinobatrachium eccentricum (synonym of H. the

cappellei) Castroviejo-Fisher et al. 2011

Hyalinobatrachium ibama (confused with and probably ;

synonym of H. pallidum) This work

Allobates aff. brunneus (confused with A. aff. ;

Hyloscirtus; synonym of H. jahni)

macrosternum in Venezuela; synonym of L. /atrans)

Appendix 6. Taxonomic changes at the genus and species levels, along with emendations that occurred since the last

systematic list by Barrio-Amoros (2009c), including changes to new species added in that list.

Dendrophryniscus minutus | Amazophrynella minuta Fouquet et al. 2012a,b

Allobates rufulus Anomaloglossus rufulus Dendrobatidae is ame and Santos

Trachycephalus venulosus | Trachycephalus typhonius Hylidae Lavilla et al. 2010

Centrolene altitudinale Centrolene altitudinalis

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Appendix 6 (continued). Taxonomic changes at the genus and species levels, along with emendations that occurred

since the last systematic list by Barrio-Amoros (2009c), including changes to new species added 1n that list.

Centrolene daidaleum Centrolene daidalea Centrolenidae This work

Centrolene notostictum Centrolene notosticta Centrolenidae This work

Centrolene venezuelense Centrolene venezuelensis Centrolenidae This work

ay RUN OD GRE MEAT: H. cappellei Castroviejo-Fisher et al. 2011

crurifasciatum

guairarepanensis

Pristimantis aracamuni Craugastoridae Sed Sear ee aes

Pristimantis lentiginosus Tachiramantis lentiginosus Craugastoridae Heinicke et al. 2015

Pristimantis prolixodiscus | Tachiramantis prolixodiscus Craugastoridae Heinicke et al. 2015

Owe elant

Hypsiboas boans Dubois 2017

Hypsiboas calcaratus Dubois 2017

Hypsiboas cinerascens Boana cinerascens Hylidae

Orrico et al. 2017; Dubois

Hypsiboas crepitans 017

Hypsiboas hobbsi /Boanahobbsi —SSs«d Hylidae Dubois 2017

Hypsiboas jimenezi |Boanajimenezi Ss Hylidae Dubois 2017

Hypsiboas lemai |Boanalemai ————Ss« Mylidae Dubois 2017

Hypsiboas multifasciatus |Boana multifasciata | Hylidae Dubois 2017

Hypsiboas ornatissimus |Boana ornatissima | Hylidae Dubois 2017

Hypsiboas roraima Boana roraima

Hypsiboas rhythmicus Boana rythmica

Hypsiboas sibleszi Hylidae

Hypsiboas tepuianus Hylidae Dubois 2017

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