Published in the United States of America

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AMPHIBIAN & REPTILE

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Oxyrhopus petolarius, Yotoco, Reserva forestal, 2012, Fdo Castro.

Amphib. Reptile Conserv.

1

December 2014 | Volume 8 | Number 2 | e87

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptiie Conservation

8(2) [Special Section]: 1-18; S1-S24 (e87).

Conservation status of the herpetofauna, protected areas,

and current problems in Valle del Cauca, Colombia

^Alejandro Valencia-Zuleta, Andres Felipe Jaramillo-Martmez, Andrea Echeverry-Bocanegra, Ron-

ald Viafara-Vega, Oscar Hernandez-Cordoba, Victoria E. Cardona-Botero, Jaime Gutierrez-Zuhiga,

and Fernando Castro-Herrera

Universidad del Valle, Grupo Laboratorio de Herpetologia, Departamento de Biologia, Cali, COLOMBIA

Abstract . — In this study, we present an analysis of the conservation status of amphibian and

reptile species by associating the natural protected areas and municipalities with the distribution

of richness in Valle del Cauca. We establish the percentage of species of amphibians and reptiles

in each of the lUCN (International Union for Conservation of Nature) threat categories and assign

local conservation categories to all species, construct distribution maps for the records of species

in relation to their threat status, and analyze the endemic and total number of species in each of

the protected areas. We found that nearly 50% of the species in the Valle del Cauca are under some

degree of risk or threat, that the largest percentage are in the Vulnerable (VU) category, whereas

37% of the fauna is not threatened (Least Concern [LC] and Near Threatened [NT]), and 13% is

categorized as Data Deficient (DD). Although the distribution of species is scattered throughout the

territory, patterns are maintained within the various regions, with areas of greater richness found in

the Pacific region and the cordilleras; the municipalities with the largest number of species under

some level of threat are Buenaventura, Darien, El Cairo, Dagua, Cali, La Cumbre, and Yotoco. The

types of protected areas with the largest number of species are the Reserves Forestales Protectoras

Nacionales (RFPN) 37%, followed by the Parques Nacionales Naturales (PNN) 18%, the Reserves

Forestales Protectoras Regionales (RFPR) 10%, and the Parques Naturales Regionales (PNR) 7.5%;

17% (~ 57 spp.) of the species in the Valle del Cauca have not been recorded in any of the protected

areas, and more than 65% of these are under some type of threat. We consider this study a starting

point for evaluating conservation priorities for the herpetofauna of Valle del Cauca.

Key words. Amphibians, reptiles, distribution, lUCN, population declines, threats

Resumen . — En este trabajo presentamos un analisis del estado de conservacion de las especies de

anfibios y reptiles relacionando las areas naturales protegidas y los municipios con la distribucion

de riqueza en Valle del Cauca. Establecemos los porcentajes de especies de anfibios y reptiles

en cada categoria de amenaza establecida por UlCN (Union Internacional para la Conservacion

de la Naturaleza) y asignamos categorias de conservacion local a todas las especies, se realizo

mapas de distribucion de los registros de las especies en relacion a los estados de amenaza, y

analizamos el numero de especies totales y endemicas en cada area protegida. Encontramos que

cerca del 50% de las especies en el Valle del Cauca presentan algun grade de riesgo o amenaza,

que la mayor proporcion se encuentra en la categoria vulnerable (VU), mientras que el 37% de la

herpetofauna no se encuentra en riesgo (preocupacion menor [LC] y casi amenazado [NT]) y el 13%

esta categorizada en datos deficientes (DD). Aunque la distribucion es diferencial a lo largo del

territorio, se conservan patrones a lo largo de las regiones, con sitios de mayor riqueza en la region

pacifica y las cordilleras; y los municipios con mayor numero de especies con algun grado de

amenaza son Buenaventura, Darien, El Cairo, Dagua, Cali, La Cumbre y Yotoco. Los tipos de areas

protegidas con mayor numero de especies son las Reservas Forestales Protectoras Nacionales

(RFPN) 37%, seguidas por los Parques Nacionales Naturales (PNN) 18%, Reservas Forestales

Protectoras Regionales (RFPR) 10%, y los Parques Naturales Regionales (PNR) 7.5%; el 17% (~

Correspondence. Email: ^alejandwvalencia08@ gmail.com (Corresponding author, Alejandro Valencia-Zuleta).

Amphib. Reptile Conserv.

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December 2014

Volume 8

Number 2 | e87

Valencia-Zuleta et al.

57 spp.) de las especies del Valle del Cauca no se han registrado en ningun tipo de area protegida

y mas del 65% de ellas presents algun tipo de amenaza. Consideramos este trabajo un punto de

partida para evaluar prioridades en la conservacion de la herpetofauna vallecaucana.

Palabras claves. Anfibios, reptiles, distribucion, UICN, declive poblacional, amenazas

Citation: Valencia-Zuleta A, Jaramillo-Martmez AF, Echeverry-Bocanegra A, Viafara-Vega R, Hernandez-Cordoba O, Cardona-Botero VE, Gutierrez-

Zuniga J, Castro-Herrera F. 2014. Conservation status of the herpetofauna, protected areas, and current problems in Valle del Cauca, Colombia.

Amphibian & Reptiie Conservation 8(2) [Special Section]: 1-18; S1-S24 (e87).

mercial-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-reptiie-conservation.org>.

Received: 12 March 2014; Accepted: 24 November 2014; Published: 09 December 2014

Introduction

Many populations of amphibians and reptiles are in de-

cline (Mendelson et al. 2006; Bohm et al. 2013), primar-

ily as a result of habitat loss, climate change, introduced

species, diseases, and illegal trafficking (Young et al.

2001; Stuart et al. 2004; Mendelson et al. 2006; Wake

2007; Rovito et al. 2009; Bohm et al. 2013). Estimates

indicate that 15-36% of the world’s species of reptiles

are threatened (Bohm et al. 2013), and according to Stu-

art et al. (2004) 22.5% of the species evaluated by lUCN

lacked sufficient information to evaluate their status. Al-

though the lUCN standardized the use of categories that

can be applied to any taxon and has attempted to cata-

logue the majority of species (lUCN 2012), many spe-

cies still have not been evaluated or lack the necessary

information for conducting an assessment; in the case of

reptiles, 59% of the species have not been assessed.

In Colombia, in addition to the above mention fac-

tors that threaten populations of amphibians and reptiles

(Rueda 1999; Ruiz and Rueda-A 2008; Velasquez et al.

2008; Isaacs and Urbina 2011; Urbina 2011; Urbina et

al. 2011; Vargas and Amezquita 2013), the social prob-

lem associated with the planting and eradication of illicit

crops threatens the fauna because of the destruction of

primary forests and the use of pesticides such as Glifo-

sato (Arroyo and Lynch 2009; Brain and Solomon 2009).

A mining crisis also has developed in the country, where

mining permits are granted to people for economic pur-

poses while the long-term impact on the environment

caused by these activities is ignored (Mancera and Alva-

res 2006; UPME 2007; Hernandez et al. 2013).

In response to these problems, early in the 1930s

“areas naturales protegidas” (= natural protected areas)

were designated in the country, which led to the fomia-

tion of “zonas forestales protectoras” (= protected forest

areas) in the department (dpto= a territorial division m

Colombia that has autonomy in the administration of re-

gional issues, planning, and the promotion of economic

and social development within its territory under the

terms established by the Constitution) of Valle del Cauca

(decree 1393/40). Regulations for deteimining the exact

management categories that competent authorities at dif-

Amphib. Reptile Conserv. 3

ferent levels can assign to protected areas, however, still

have not been implemented in the country (Vasquez and

SeiTano 2009). Currently, 197 reserves of all types exist

in the Valle del Cauca; three natural national parks are the

most important because of their large size and location in

areas of high herpetofaunal diversity, in the dpto and in

the country — the “Cordillera Occidental” (= the Western

Cordillera) and the “Regibn Pacifica” (= Pacific Region)

(Cardona et al. 2013); environmental problems, however,

are present in these areas, as their biological patrimony

has not been fully elucidated (Patino 2010).

Valle del Cauca is one of the dptos with the greatest

amount of herpetofaunal species richness (333 species),

which represents 24% of the amphibian and 25% of the

reptile species recorded from the country (Cardona et

al. 2013). We are unaware, however, of the number of

threatened species in the dptos, or plans for their con-

servation. In a red book of amphibians, Castro-H and

Bohvar-G (2010) included 68 species under some type of

threat, and along with an action plan for the conservation

of amphibians in Valle de Cauca provided by CoiTedor et

al. (2010); these publications are considered pioneer ef-

forts in conseiwation; in general, research programs usu-

ally are developed separately and independently.

The objective of this paper is to present an analysis

of the conservation status of the species of amphibians

and reptiles by associating the natural protected areas and

municipalities with the distribution of richness in Valle

del Cauca, as a starting point for evaluating conservation

priorities for the herpetofauna of this region.

Materials and Methods

Study area

Valle del Cauca is a dpto in southwestern Colombia that

consists of 42 municipalities (Eig. 1) with a total surface

area of 22,142 km^; it contains a diversity of landscapes,

including very humid tropical forests, premontane plu-

vial forests in warm transition, dry and very dry tropical

forests, and lowland montane to pluvial montane forests

that range in elevation from sea level to 4,000 m. This

December 201 4 I Volume 8 I Number 2 I e87

Conservation status of the herpetofauna in Colombia

dpto has allocated 233,889 ha for parks and natural re-

serves, which because of their ecological importance and

abundance of natural resources have been established in

various strategic ecosystems and protected areas (Gomez

et al. 2007).

Sources of information

In updating their list on the herpetofauna of Valle del

Cauca, Cardona et al. (2013) considered the following:

• Geographic data: Obtained from bibliographic sourc-

es, field notes, and biological collections of amphib-

ians and reptiles at the Universidad del Valle (UV-C),

and online databases from the Institute de Ciencias

Naturales (ICN), and the National Museum of Natural

History at the Smithsonian Institution (USNM).

• Threat category (species recorded from the dpto were

catalogued using the following criteria): trafifieking

in speeies, deaths caused by vehicular traffic or by

humans, distribution within the dpto (eco-regions,

localities, life zones), occurrence in disturbed habi-

tats, frequency of observation, number of citations in

publications, and the presence of species in protected

areas. All these criteria were scored from 0 to 4, where

0 means no risk in the particular criteria, 3 high risk,

and 4 is unknown (see supplemental material at am-

phibian-reptile-eonservation.org for the definition of

the score in each criteria). Based on the data obtained

for each species, it calculated the weighted average

for the different natural groups (amphibians, lizards,

and snakes), and assigned a pereentage in the final

score to each criteria according to the natural group,

because the same criteria does not affect each natu-

ral group in the same way (see supplemental material

for the percentage assigned in each criteria). With the

weighted average of each species, these were assigned

to some of the categories proposed by the lUCN, as

follows: LC 0-1.4, NT 1. 5-2.0, VU 2. 1-2.6, EN 2.7-

3.0, CR 3. 1-3.3, DD 3.4^.0. Each category was jus-

tified aecording to the appendix of the lUCN (2012),

especially considering the threats to each species. Ad-

ditionally, the threat status for each species reported

from the dpto was examined by searching through the

lUCN Red List of Threatened Species (http://www.

iuenredlist.org/), the red books of amphibians and rep-

tiles in Colombia (Castano-M 2002; Rueda-A et al.

2004), and the red book of amphibians from Valle del

Cauca (Castro-H and Bohvar-G 2010).

Fig. 1. Political map of Valle del Cauca (Colombia). North: El Aguila (AGE), El Cairo (CR), Ansermanuevo (ASN), Argelia (ARG),

Cartago (CTG), Ulloa (ULA), Alcala (ACL), Toro (TR), Versalles (VRSL), Obando (OBD), La Union (UN), El Dovio (DV), Rolda-

nillo (RDNL), La Victoria (VTR), Zarzal (ZRZ), Bolivar (BLV); East: Sevilla (SVL), Caicedonia (CDN); Middle: Bugalagrande

(BGG), Tmjillo (TIE), Andalucia (ADL), Rio Erio (RE), Tulua (TL), San Pedro (S/PD), Yotoco (YTC), Darien (DR), Buga (BG),

Guacari (GCR), Ginebra (GNB), Vijes (VJ), Restrepo (RTP), Cumbre (CMB), El Cerrito (CRT); South: Palmira (PMR), Yumbo

(YMB), Cali (CL), Candelaria (CDR), Pradera (PDR), Elorida (ERD), Jamundi (JMD); West: Buenaventura (B/tura), Dagua (DG).

Amphib. Reptile Conserv.

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December 2014 I Volume 8 | Number 2 | e87

Valencia-Zuleta et al.

• Protected areas: Each species was recorded according

to geographic location and the use of bibliographic

resources on protected areas within the dpto, consid-

ering the important areas with a wide extension and

with the ability to hold a great diversity of herpeto-

fauna. The definition of protected areas were defined

based on the Decree-Law 622 of 1977 and 2372 of

2010 of the National Government (in parentheses the

areas that were chosen in this work):

• Parques Nacionales Naturales (PNN) is an area

of great extent permitted ecological autoregula-

tion and whose ecosystems in general have not

been substantially altered by human exploitation

or occupation, where plant and animal species,

geomorphological resorts, historical or cultural

events have scientific, educational, aesthetic

and recreational value and their perpetuation is

subjected to an appropriate management regime

(Farallones de Cali, Las Hermosas, Uramba-

Bahia Malaga, Tatama).

• Santuario de Flora y Fauna (SFF) is dedicated to

preserving wildlife species or plant conununi-

ties to preserve genetic resources of native flora

and fauna (Decreto 622 de 1977), (Isla Mal-

pelo).

• Parque Natural Regional (PNR) is a regional

geographic area where landscapes and strate-

gic ecosystems, maintain their structure, com-

position and function. The natural and cultural

values are associated with human disposition

for preservation, restoration, knowledge, and

enjoyment (La Sierpe and Paramo del Duende).

• Reservas Forestales Protectoras (RFP) is a geo-

graphical area where forest ecosystems main-

tain their function, although their structure and

composition have been modified and associ-

ated natural values are accessible to the human

population to who allocated their preservation,

sustainable use, restoration, knowledge, and en-

joyment. In this type of protected area are the

forests, national (RFPN) (Amaime, Anchicaya,

San Cipriano and Escalerete rivers, Bosque de

Yotoco, Dagua, Cali, Tulua, Sonso-Guabas,

Cerro Dapa-Carisucio) and regional (RFPR)

(Bitaco and Frayle-Desbaratado) protection.

• Reserva Natural (RN) is an area in which undis-

turbed conditions exist or have undergone mini-

mal human disturbanee of flora, fauna, and soil,

and it is intended for conseiwation, research, and

study of its natural wealth (Laguna de Sonso).

• Distrito de Manejo Integrado (DM1) is a geo-

graphical space where landscapes and ecosys-

tems retain their composition and function, al-

though their structure have been modified and

whose natural and cultural associated values are

set to reach the human population who allocated

their sustainable use, preservation, restoration,

knowledge, and enjoyment (La Plata and En-

clave Subxerofitico Atuncela).

• Munieipalities: Each species was recorded based on

its documented geographic location within the mu-

nicipalities of the dpto.

Analysis of the Data

The species distribution model for each threat category

was performed using all the records collected from the

different museums and georeferenced using Google

Earth 7.1.2.2014; these models were constructed in Max-

Ent Version 3.3.3a. The software generated models us-

ing the theory of maximum entropy only when presence

data were available (Phillips et al. 2006). For this work,

we used the 19 climate layers of the WorldClim project

(www.worldclim.org, spatial resolution of 30 arc second

or ~ 1 km^). To evaluate the predictive ability of the mod-

els generated, the Area Under the Curve (AUG) score was

taken into account. The AUG is a ranked approaeh for as-

sessing model fit, which determines the probability that

a presence loeation will be ranked higher than a random

background location (Phillips et al. 2006). The predietion

models generated by MAXENT were mapped in ArcGIS

10.1 (ESRI 2013), with only the detection probabilities

above 0.5 taken into aeeount.

The percentages of amphibian and reptile species for

each threat status was determined, and through histo-

grams indicate the endemic number of species and total

number of species in each of the protected areas.

Results

Status of threats to the herpetofauna

Approximately 51% of the species in the dpto showed

some degree of risk or threat. The majority of amphibians

(60%) are in one of the threat categories, with the Vulner-

able (VU) containing the most species (59), followed by

the Critically Endangered (CR) and Endangered (EN),

each with 29, and 27 in the Near Threatened (NT) (Fig.

2 A) categories. Conversely, more than one-third of the

reptile species show some degree of risk, with those in

the NT and VU containing the largest number of species

(38 in each), followed by the EN (14), and a few (six) in

the CR (Fig. 2B) categories. Of the remaining herpeto-

faunal species in the dpto, 17% show no risk (EC), and

14% are Data Deficient (DD; see supplemental material).

Amphib. Reptile Conserv. 5 December 2014 | Volume 8 | Number 2 | e87

Conservation status of the herpetofauna in Colombia

® LC m NT ■ VU □ EN ■ CR □ DD

Fig. 2. Threat status of the percentage of herpetofaunal species in Valle de Cauca: (A) = Amphibians, (B) = Reptiles.

Eighty percent of the amphibian families contain spe-

cies under some level of threat, with 40% of the species

in the family Craugastoridae in one of the threat catego-

ries. In general, the NT species are represented mostly

in the families Craugastoridae, Centrolenidae, Hylidae,

Dendrobatidae, and Leptodactylidae. In addition, more

than one-half of the VU species are in the family Crau-

gastoridae; in particular, the VU and EN species follow

the same pattern and include the families Craugastoridae,

Centrolenidae, Dendrobatidae, and Hylidae. Significant-

ly, 70% of the CR species are grouped in the Craugas-

toridae, Bufonidae, and Centrolenidae, families with the

greatest risk of losing species, along with representatives

of the family Hemiphractidae, which are restricted to the

EN and CR. As with the amphibians, most families of

reptiles (84%) contain species under some level of threat.

In particular, most of the threatened species are in the

families Colubridae, Dactyloidae, Dipsadidae, and Gym-

nophthalmidae, with most in the NT and VU categories.

Over 30% of the NT species are in the family Colubridae,

followed by the Dipsadidae and Dactyloidae, whereas

the VU species are mostly in the Dactyloidae and Dip-

sadidae. The majority of EN species are in the families

Colubridae, Dipsadidae, and Gymnophthalmidae. Eur-

thermore, the CR species are represented by one species

in each family, except for the Dactyloidae.

The modeling of the maps present an AUC of 0.754-

0.83, indicating a better performance than the random

models (Manel et al. 2001). Herpetofaunal richness is

scattered throughout Valle del Cauca, but the areas (see

Cardona-B. et al. [2013] to define ecoregions in the Valle

del Cauca) with the greatest amount of richness are the

Pacific region and the Cordilleras (Fig. 3 A). The NT spe-

cies show a wide distribution along the western Cordil-

lera (specifically in the northern and central area), and

cover a large area along the Interandean Valley and the

Pacific (Fig. 3B). The VU species are found along the

foothills and northern and central portions of the western

Cordillera, but are less represented in the central Cor-

Amphib. Reptile Conserv. 6

dillera and in the Interandean Valley (Fig. 3C). The EN

species are found in two important areas, the Pacific re-

gion and the western Cordillera in the northern part of the

dpto; in the central Cordillera, a few representatives are

found in the high elevation areas of Sevilla, Tulua, and

Buga, to the north, and Palmira, Pradera, and Florida, to

the south (Fig. 3D). The distribution of the CR species is

important, based on the presence of Atelopus in the cen-

tral and western Cordilleras and groups of Pristimantis

in highland areas of the western and central Cordilleras;

in addition, the centrolenids and dendrobatids are found

in the western Cordillera and the Pacific region (Fig. 3E).

Significantly, the DD species are distributed all along the

dpto, but show similar patterns to species in the threat

categories (Fig. 3F).

In particular, 90% of the municipalities in Valle del

Cauca contain one species in at least one of the threat cat-

egories, whereas 62% of the municipalities contain more

than two species. The municipalities of Buenaventura

(82 species), Darien (61), El Cairo (51), Dagua (45), Cali

(42), La Cumbre (19), and Yotoco (11) contain the great-

est number of species under some level of threat (Fig.

4). A similar pattern was found in these municipalities,

where the majority of species fall into the VU category,

followed by the EN, and last by the CR, with the only

exceptions in the municipalities of Cali and La Cumbre.

The municipalities of Buenaventura (six amphibians,

five reptiles), El Cairo (10, zero), and Darien (eight, one)

contain the largest number of CR species.

The herpetofauna and protected areas

Protected areas in Valle del Cauca with the greatest num-

ber of species are the RFNP (228 species), followed by

the PNN (120), RFPR (60), PNR (45), DMI (34), and

RN (21), and the area with the least number is the SSF

(three species of reptiles). The majority of the species in

the RFPN and the PNN are in the VU, and in the remain-

ing areas most of the species are in EC, except for the

December 2014 I Volume 8 | Number 2 | e87

Valencia-Zuleta et al.

Fig. 3. Distribution maps for the richness of herpetofauna in the most documented areas in Valle del Cauca. (A) richness, (B) NT

species, (C) VU species, (D) EN species, (E) CR species, and (E) DD species.

SEE (Eig. 5). The areas that protect the largest number of

species in a threat category are the REPN (162 species),

the PNN (84), the REPR (37), and the PNR (25), and the

areas that protect the least numbers are the DMI, RN, and

SEE (13, four, and three, respectively). Throughout the

dpto, 17% (~ 57 spp.) of the species are not found in a

protected area, and more than 65% of those fall into one

of the threat categories (NT = four, VU =10, EN = seven,

and CR = 15). In addition, information is not available

for 31% of these species (DD). The protected area with

the largest number of species is the REN de Anchicaya

(183 species), followed by the PNN Earallones de Cali

(90), the REPN of the rivers San Cipriano and Escalerete

(84), and the REPR de Bitaco (49).

Endemic species

Nineteen endemic species are found in the dpto, which

represents only 6% of the species diversity. Amphibians

represent the largest number of species (13), with 75% in

one of the threat categories: CR (six species), EN (three),

and VU (two); the remaining 15% are categorized as

DD. With regard to the threat categories for reptiles,

three species are in the CR, and the other three are DD

because they lacked sufficient information for an assess-

ment (Eig. 6). The endemic species are distributed in four

types of protected areas, the PNN and the REPN, which

contain a high number of species, and it is worth not-

ing that the PNR el Paramo del Duende and the SPP Isla

Amphib. Reptile Conserv. 7 December 2014 | Volume 8 | Number 2 | e87

Conservation status of the herpetofauna in Colombia

BVU DEN bCR

Fig. 4. Municipalities in Valle del Cauca with the greatest number of species in the threat categories: (A) Buenaventura, (B) Dagua,

(C) Cali, (D) Darien, (E) El Cairo, and (E) La Cumbre.

de Malpelo are the types of areas with the greatest num-

ber of endemic species. Only four species (Pristimantis

diaphonus, Anomaloglossus atopoglossus, Nymphargus

armatus, and Geophis betaniensis) are not found in any

of these areas.

Discussion

Conservation status of amphibians and reptiies

The need to recognize the status of a species in a specific

area should be considered baseline information for devel-

oping studies and management plans for its conservation.

The lUCN categorizations are generally applied globally

for each taxon to determine the status of a species at the

local or regional levels, and are considered advanced

studies (Castro-H. and Bolfvar-G. 2010); information

on certain species (e.g., population status, natural his-

tory) is necessary to elicit an approximate categorization.

Records for the DD species (13% of the species in this

study) are not well represented in herpetological collec-

tions (some are only known from their original descrip-

tions), and thus it is not possible to determine their status.

The dpto contains a high proportion of amphibian

species (60%) in one of the threat categories, which rep-

resents nearly one-half of the total herpetofauna of the

dpto, and the majority of these species show a moderate

risk of extinction or population decline over the medium

term (VU). In comparison with the results of Castro-H

and Bolivar-G (2010), we show a significant increase

in the number of species in the CR (11), EN (10), VU

(12), and NT (one) categories, indicating that the risk of

disappearance has increased in certain species, which is

troublesome.

The lack of a threat status among the reptiles results

from insufficient basic ecological information and the

actual distribution of their populations (Urbina-Cardona

2008), for which an evaluation of the threat status has

focused on specific species or groups (e.g., the red book

of reptiles in Colombia), and thus has become a problem

for planning conservation strategies. For this reason, the

status of populations of reptile species in a given area has

been proposed as a mechanism to change attitudes and

generate interest in preserving these organisms (Dodd

2001), the protection and restoration of large areas these

organism inhabit (Roe et al. 2004; Franga and Araujo

2006), species-specific information, field studies, de-

mographics, natural history, and possible threats (Cagle

2008; Fifes et al. 2013). Significantly, this study is a local

proposal that easily addresses the status of reptile species

in Valle del Cauca, so that more effective strategies can

be accomplished. This study is the first to assess many

species of reptiles, and in spite of their low density threats

might make them vulnerable and affect their abundance

in the dpto; in many cases, characteristics of their natural

history allow them to avoid these conditions.

The conservation of snakes remains subjective, be-

cause the current status of many species remains un-

Amphib. Reptile Conserv.

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December 2014 I Volume 8 | Number 2 | e87

Valencia-Zuleta et al.

A

Fig. 5. The herpetofauna of Valle del Cauca according to (A)

in each type of protected area.

Fig. 6. Endemic species listed according to (A) threat category, and (B) by protected areas in Valle del Cauca.

known. According to Lynch (2012), these animals are

some of the most threatened because their deaths are

provoked by people living in rural areas, vehieles trav-

eling on highways, the loss of habitat, climate change,

and illegal traffieking. Vargas et al. (2011) showed that

even in a protected area such as the RFPN Bosque de

Yotoeo (Valle del Cauea), some snakes are vulnerable to

the effects of roads on account of the prolonged amount

of time that deaths by vehicles have been caused, con-

sidering the low density of populations and small size of

the reserve.

Threatened species in the municipaiities and

protected areas

Because of their considerable size and strategic loca-

tion in high diversity life zones, municipalities such

as Buenaventura, El Cairo, and Dagua contain a large

number of threatened species of both groups (Cardona

et al. 2013); however, these areas are the focal point of

anthropogenic pressures, and thus certain species have

^

been affected. Moreover, municipalities such as El Agui-

la, Ulloa, El Dovio, Versalles, Ansermanuevo, Elorida,

Pradera, Palmira, El Cerrito, Buga, Tulua, and Sevilla,

among others, laek adequate sampling and are under-

represented in collections, and the few data available

from these municipalities correspond to widely distrib-

uted generalist species, such as colubrid and dipsadid

snakes that because of their high dispersal abilities ean

easily adapt to anthropogenic environments, and thus are

categorized as EC (Adams 1994). Conservation efforts,

therefore, should be foeused in habitats influeneed by the

western versant of the central Cordillera, important areas

for species in the different threat categories.

The distribution of the threat categories in the dpto

refleets the pressures on the categorized species; for ex-

ample, a large number of CR species are in the genus

Atelopus and most of these are distributed in the two

Cordilleras, and like their eongener speeies possibly have

been seriously affected by chytridiomicosis (Bonaccorso

Amphib. Reptile Conserv.

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December 2014 I Volume 8 | Number 2 | e87

Conservation status of the herpetofauna in Colombia

Hypsiboas picturatus, Buenaventura, San Cipriano, 2003, Fdo Castro.

and Guayasainin 2003; Sanchez et al. 2008; Coloma et

al. 2010). In addition, the fragmentation and loss of habi-

tat due to human activities have affected certain sensitive

species (like the centrolenids and dendrobatids) princi-

pally distributed in the Andean Cordilleras (Hutter et al.

2013) and in the Pacific region (Castro-H and Bolivar

2010). Besides these pressures, several species have ex-

perienced a population decline as a result of illegal traf-

ficking and collection for scientific studies (Castro-H and

Bolivar-G 2010; Corredor et al. 2010).

Additionally, in evaluating natural groups the “Siste-

mas Municipales de Areas Protegidas” (Municipal

System of Protected Areas; SIMAP) and the “Sistemas

Nacionales de Areas Protegidas” (National System of

Protected areas; SINAP) have centered in municipalities

such as Buenaventura, Cali, Dagua, La Cumbre, El Cai-

ro, Darien, and Yotoco in an effort to better understand

the conservation status of species in these areas, and to

promote the monitoring of populations of these organ-

isms. Furthermore, a network of community reserves is

present in the municipality of El Cairo, in the Serrania

de los Paraguas (which were not included in our analy-

sis), and we suggest studying and monitoring the natural

populations of many threatened and endemic species in

this area in order to promote their conservation.

A greater number of species are found in REPN than

in the PNN because of three factors: (1) an extensive area

of the REPN (ca. 154,091 ha) lies in Valle del Cauca,

(CVC 2012); although is not larger than that of the PNN,

compared to other types of areas it represents a substan-

tial part of the territory; (2) several reserves in the dpto

are located in areas of great richness, such as the REPN

of Anchicaya, and of the San Cipriano and Escalerete

rivers in the Pacific Region (Cardona et al. 2013); and

(3) extensive research projects have been conducted in

several of these areas, for which many bibliographic ref-

erences are available and a large number of specimens

are present in collections, such as in the REN del Bosque

de Yotoco, in which the research group from the Labo-

ratorio de Herpetologia de la Universidad del Valle has

been conducting inventories from 1978 until the present

and recorded a large list of species, of which some are no

longer being reported from the area (Castro et al. 2007).

In spite that one of the most effective methods for

preserving natural spaces is the use of specific forms

of protection and legal regulation that limit or prohibit

the development of productive or extractive activities

(Vasquez and Serrano 2009), state policies are necessary

to guarantee the conservation of important ecological

areas (Castro-H and Bohvar-G. 2010) by means of the

environmental authority granted to autonomous corpora-

tions. In Valle del Cauca, several CR species undergo-

ing population pressures were found in the PNN and the

REPN, such as Oophaga lehmani, in which the principal

causes for decline in protected areas are the loss of habi-

tat and illegal trafficking (Avila 2007), a clear example

of not applying the articles of the Codigo de Recursos

Naturales Renovables y Proteccion del Medio Ambiente

Amphib. Reptile Conserv.

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December 2014 I Volume 8 | Number 2 | e87

Valencia-Zuleta et al.

Bolitoglossa medemi, Buenaventura, Bendiciones, 2011.

Strobomantis ruizi, Trujillo, Andinapolis, 2010.

Andinobates bombetes, Darien, Lago Calima, 2005.

Gastrotheca antomia, Dagua, Alto Queremal, 1993, Extinct.

Oophaga histrionica, Buenaventura, Anchicaya, 2000.

Agalychnis spurelli, Buenaventura, san Cipriano, 2003.

Diasporus gularis, Buenaventura, Bazan, 2010.

Pristimantis achatinus, Buenaventura, Bazan, 2010.

Amphib. Reptile Conserv.

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December 2014 I Volume 8 | Number 2 | e87

Conservation status of the herpetofauna in Colombia

{Code of Renewable Natural Resources and Environ-

mental Protection; CRN). A similar situation exists with

other species of amphibians and reptiles that are under

great pressure in protected areas of Valle del Cauca, such

as the ones mentioned previously and including mining,

death caused by humans, and pesticide contamination

from the fumigation of illicit crops, which illustrates a

lack of control in these protected areas. In addition, the

current laws in these areas and the reasons for proposing

them are not clear, such as for preservation, conservation,

and ecotourism, and in some areas they could supersede

their carrying capacity.

Castro-H and Bohvar-G (2010) indicated that with-

in the great variety of habitats found in the dpto those

with specific characteristics became inclusive centers

of speciation, and that these unique areas are of great

importance because they contain endemic species. The

endemic species in these areas could easily disappear

on account of an environmental threat, because of their

specialized requirements and limited distribution. The

size of their distributional range is indispensable for their

conservation, and these species should be included in at

least one state protected area where conditions are stable,

so there is less potential for risks and their populations

can continue to develop (Rueda-A. et al. 2004). These

requuements are necessary for their preservation, but it

is worth noting that four endemic species {Nymphargus

armatus md Anomaloglossus atopoglossus [CR], Pristi-

mantis diaphonus [EN]; and the snake Geophis betani-

ensis [DD]) are not found in any of protected areas des-

ignated by the government and/or autonomous regional

corporations (CAR = Institutions that are responsible for

implementing the policies, plans, programs, and projects

on environment and renewable natural resources. Also,

they give a full and application to current legal provi-

sions, under the regulations, standards, and guidelines is-

sued by the ministry of environment), which makes them

even more susceptible to threats.

Global categorization vs local situations

Many species of continental turtles and crocodilians

are sacrificed for consumption of their meat and eggs,

and commercialization of their skins. In addition, pet

commercialization, global warnung, and developmen-

tal activities such as hydroelectric plants also have had

a negative impact on their populations (Rueda-A. et al.

2007; Paez et al. 2012). For these reasons, these char-

ismatic species are used to promote studies (biological

and economic) and the categorization of these organisms

(Castano-M. 2002; Paez et al. 2012). Various local pres-

sures, however, lead to an analysis of the situation or

threat status of these species; for example, Kinosternon

leucostomum (NT in this study) is a broadly-distributed

species for which we have wide information on its ecol-

ogy and reproductive biology (Giraldo et al. 2012), but

it has been affected by habitat deterioration and is con-

Amphib. Reptile Conserv.

sidered the most trafficked pet trade vertebrate species

in southwestern Colombia (Galvis-R. and Corredor-L.

2005), which threatens the natural populations.

Although the loss of biological diversity in Colombia

has been studied for several years, and plans for the man-

agement of threatened species that include a prioritized

list of amphibians (Castro-H and Bolivar 2010) have

been implemented at the regional and national levels, ad-

ditional actions and research are still required. Some spe-

cies in Valle del Cauca that appear in the lUCN category

of LC, such as Gastrotheca argenteovirens (Rarmrez-P.

et al. 2004) mdAnolisfraseri (Castaneda et al. 2011), are

at risk and others listed as VU, such as Centrolene geck-

oideum (Bolivar et al. 2004) and Gastrotheca antomia

(Castro and Lynch 2004), have not been reported from

the dpto in recent years, which suggests a subjectivity

in analyzing the threat category in these species, espe-

cially on a regional basis. Also, the fossorial habits and

difficulty in locating organisms such as caecilians must

be considered, and thus their threat status is difficult to

determine. According to the lUCN, most species of cae-

cilians are categorized as LC and two species {Caecilia

guntheri and Oscaecilia polizona) as DD; however, on

a local scale and considering the lack of information for

these organisms, not enough data is available to establish

a category in the dpto, as reflected in the family Caeci-

liidae. Similarly, other species might appear stable, but

with additional data and the implementation of manage-

ment plans their threat status might be updated so that

protected areas will be able to comply with their func-

tion and agreements, in addition to the implementation of

management plans for the short, medium, and long terms

that are in place but have not been assumed by the envi-

ronmental authority (CAR del Valle del Cauca, CVC),

where all the stakeholders are included.

Conclusions

One -half of the herpetofauna of Valle del Cauca is under

some degree of threat, which is important for the conser-

vation of this fauna, mainly in two areas in the western

Pacific region (municipality of Buenaventura) and north

on western Cordillera (municipality of Cairo). Tliese

hotspots are locations where extensive sampling of the

herpetofauna has been conducted, and where species un-

der some degree of threat occur differentially along the

dpto.

Additional information on the distribution of amphib-

ians and reptiles, the current status of populations, and

the natural history of species in Valle del Cauca are nec-

essary to develop an initiative for a conseiwation program

with specific short-term objectives, so that decisions can

help mitigate negative effects in the populations. Fur-

thermore, the protected areas and municipahties in the

dpto must develop monitoring plans in their areas that

contain detailed information on the presence or absence

December 201 4 I Volume 8 | Number 2 | e87

12

Valencia-Zuleta et al.

Hypsiboas rubracila, Buenaventura, Bazan, 2014.

Centrolene gekkoideum, La Cumbre, Chicoral, 1988, Extinct.

Pristimantis juanchoi, La Cumbre, Chicoral, 2010.

Anolis lyra, Buenaventura, Bazan, 2010.

Kinosternon Leucostomum, Buenaventura, Zaragoza, 2013.

Basiliscus galeritus, Buenaventura, Zaragoza, 2013.

Thecadactylus rapicaudus, Buenaventura, Zaragoza, 2009.

Diploglossus monotropis, Buenaventura, Bahia Malaga, 2013.

Amphib. Reptile Conserv.

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December 2014 I Volume 8 | Number 2 | e87

Conservation status of the herpetofauna in Colombia

of species, so that along with other organizations they

can negotiate the protection and conservation of ecosys-

tems essential for the herpetofauna. In particular, we ask

the CAR to ensure compliance with the development of

these initiatives.

The conservation of endemic species of amphibians

and reptiles should be clear and we must recognize that

this requires special management, but the current regula-

tions are not clear enough to define the measures that ac-

tually will allow the implementation of specific conser-

vation plans for these species; in many places, the type

of area will not allow the sustainability of these species,

which are an emblem for the dpto.

Acknowledgments. — We would like to start by thank-

ing Reynel Galvis for his help in the early construction

of the manuscript. Thanks goes to the Vicerrectoria de In-

vestigaciones of Universidad del Valle and the program

of Jovenes Investigadores of Colciencias for their effort

and assistance. A special thanks is extended to Azul y

Verde Foundation, Serraniagua Foundation, Nasmille

and family in Chicoral, Amparo Bubu and family, and all

those people at the study sites who have supported and

enabled us to gain valuable information for this paper.

Final gratitude is extended to the Herpetology lab and the

reference collection of amphibians and reptiles of Uni-

versidad del Valle (UV-C) where most of the information

obtained for this manuscript is deposited. We thank Louis

Porras for translating the original version of this paper

into English and copy editing the final version.

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servation in Latin America. Conservation Biology

15(5): 1,213-1,223.

Alejandro Valencia-Zuleta is a biologist from the Universidad del Valle (Cali-Colombia), working in

the Young Researcher program of Colciencias and as a volunteer in the group jSalven las ranas! Cali-

Colombia, under the campaign SAVE THE EROGS ! His main interests include the conservation, ecology,

taxonomy, and natural history of amphibians and reptile of Colombia, with special focus in the Colombian

south-west. At present, he studies the eommunity and trophie eeology of anurans in the Colombia Paeifie.

Photo by Andres F. Jaramillo.

Amphib. Reptile Conserv.

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December 2014 I Volume 8 | Number 2 | e87

Conservation status of the herpetofauna in Colombia

Andres Felipe Jaramillo-Martinez is a biologist from the Universidad del Valle (Cali-Colombia). His

■ interests are the photography of natural biodiversity, the assessment of the natural history of amphibians

reptiles for an ecologic and functional perspective. He is also interested in the taxonomy of salaman-

ders, anurans and snakes. He presently works on the ecology of salamanders and anurans populations in

the Colombian south-west. Photo by Eliana Barona.

Andrea Echeverry-Bocanegra is a biologist from the Universidad del Valle (Cali-Colombia). She did her

bachelor thesis research on the presence of skin alkaloids in Colostethus fraterdanieli (Anura: Dendrobati-

dae). Her areas of interest include the physiology, ecology, taxonomy, and ethology of amphibians, reptiles,

and mammals. Photo by Andrea Echeverry.

a Ronald Andres Viafara-Vega is a biologist from the Universidad del Valle (Cali-Colombia) interested

in the natural history, ethology, ecology, and conservation of the amphibians and reptiles, and studies that

apply molecular biology in these groups. His interest and future pursuit will be the taxonomy of Glass and

leptodactylid frogs. Photo by Ronald A. Vidfara.

Oscar D. Hernandez-Cordoba received his B.S degree in biology from the Universidad del Valle (Cali-

Colombia), in Cali, Colombia in 2013. As a student, he joined the Laboratorio de Herpetologia de la

Universidad del Valle (Cali-Colombia) investigation group, guided by Dr. Fernando Castro. In this group

he developed investigative interests in evolutionary ecology, ecotoxicology, and conservation biology of

amphibians, reptiles and birds. Photo by Ana Goeta.

Victoria E. Cardona-Botero is a biologist from la Universidad del Valle (Cali-Colombia) and young re-

searcher of Colciencias. At present, she works on the community ecology of anurans in the Colombian Pa-

cific. Her main topics of interest are the ecology, natural history, ethology, and systematics of amphibians

and reptiles, with a special focus on the ecology and natural history of birds. Photo by Andres E Jaramillo.

Jaime Gutierrez-Zuniga is a biologist from la Universidad del Valle (Cali-Colombia). His topics of inter-

est are the bioacoustics and ecology of amphibians and reptiles. Photo by Ronald A. Vidfara.

Fernando Castro-Herrera, Biologist, Universidad del Valle (1976) and Ph.D. (Community Ecology)

University of North Texas (1988) — advisor Dr. Lloyd C Fitzpatrick. He is presently a professor at la Uni-

versidad del Valle (Cali-Colombia) and head of the research group, Laboratorio de Herpetologia founded

in 1993 training students in the scientific study and research of amphibians and reptiles in Colombia. This

lab has been supported by the Universidad del Valle in Cali Colombia and research focuses on biodiversity,

natural history, ecology, toxicology, and assessment in conservation and sustainable use of biodiversity.

For more information reference: http://herpetologia.correounivalle.edu.co/. Fernando has been the leader

of a generation of Colombian biologists, since the latter part of the twentieth century, in studies on amphib-

ians and reptiles of Colombia. Three of his latest publications already in circulation are:

• Small changes in vegetation structure changes in amphibian create great ensembles in the Colombian

Pacific rainforest. Tropical Conservation Science 6(6): 749-769 (2013). Available: http://tropicalconser-

vationscience.mongabay.eom/content/v6/TCS-2013-Vol6%286%29_749-769_Cortes-et-al.pdf [Accessed:

30 November 2014].

• Amphibia, Anura, Bufonidae, Atelopus eusebianus (Rivero & Granados-Diaz, 1993): Distribution ex-

tension for Valle del Cauca, Colombia. Check List 10(3): 682-683. Available: http://www.biotaxa.org/cl/

article/view/10.3.682/9327 [Accessed: 30 November 2014].

• Diversidad de la herpetofauna en el Valle del Cauca (Colombia): Un enfoque basado en la distribucidn

por ecorregiones, alturay zonas de vida. Biota Colombiana 14(2): 156-233. (2013). Available: http://www.

redalyc.org/articulo. oa?id=4913 1094008 [Accessed: 30 November 2014].

Animal photo credits: Eernando Castro-Herrera.

Amphib. Reptile Conserv.

18

December 2014 I Volume 8 | Number 2 | e87

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptiie Conservation

8(2) [Special Section]: 1-18; S1-S24 (e87).

Supplemental Material

Conservation status of the herpetofauna, protected areas,

and current problems in Valle del Cauca, Colombia

^Alejandro Valencia-Zuleta, Andres Felipe Jaramillo-Martmez, Andrea Echeverry-Bocanegra, Ron-

ald Viafara-Vega, Oscar Hernandez-Cordoba, Victoria E. Cardona-Botero, Jaime Gutierrez-Zuhiga,

and Fernando Castro-Herrera

Universidad del Valle, Grupo Laboratorio de Herpetologla, Departamento de Biologla, Cali, COLOMBIA

Citation: Valencia-Zuleta A, Jaramillo-Martmez AF, Echeverry-Bocanegra A, Viafara-Vega R, Hernandez-Cordoba O, Cardona-Botero VE, Gutierrez-

Zuhiga J, Castro-Herrera F. 2014. Conservation status of the herpetofauna, protected areas, and current problems in Valle del Cauca, Colombia.

Amphibian & Reptile Conservation 8(2) [Special Section]: 1-18; S1-S24 (e87).

mercial-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: 12 March 2014; Accepted: 24 November 2014; Published: 19 December 2014

Table 1. Taxonomic list of amphibians and reptile of the department of Valle del Cauca (Cardona-B. et al. 2014). Actualization of

threat categories based on: lUCN (red list), Red Book of Amphibians (Rueda et al. ) and Reptile (Castano-M. et al 2002) of Colombia,

Red Book of Amphibians of Valle del Cauca (Castro-H. and Bolivar-G 2010), and current categorization of the amphibians and reptile

for Valle del Cauca. Also is denoted the municipalities and protected areas where the species is distributed; and a (|) is noted in the

category when a species is endemic for the department.

TAXON

lUCN

Red book of

Colombia

Red Book of

Valle del Cauca

Categorization for

the departament

Municipality

Protected area

CLASS AMPHIBIA

ORDEN ANURA

FAMILIA AROMOBATIDAE

Allobates talamancae

(Cope, 1875)

LC

NT

Bu, Dar

RFPN Anchicaya

Anomaloglossus

atopoglossus (Grant,

Humphrey & Myers,

1997)

DD

CR

CR, B1ab(iii)(t)

Cai

Anomaloglossus lacri-

mosus (Myers, 1991)

DD

VU

EN, Bla

Bu

RFPN Anchicaya,

PNR La sierpe

FAMILIA BUFONIDAE

Atelopus cf. famelicus

(Rivero and Morales,

1 995) {sensu latu)

CR

CR, Blab(iii)

Bu, Cal, Yo

RFPN Anchicaya,

RFPN de Cali

Atelopus chocoensis

(Lotters, 1992)

CR

CR, B2ab(iii)

Cai

Atelopus eusebianus

(Rivero and Granados,

1993)

CR

EN

CR, B2ab(iii)

FI

Atelopus cf. ebenoides

(Rivero, 1963)

CR

CR, Blab(iii)

Ce

Correspondence. Email: ^ alejandrovalencia08@gmail.com (Corresponding author, Alejandro Valencia-Zuleta).

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TAXON

lUCN

Red book of

Colombia

Red Book of

Valle del Cauca

Categorization for

the departament

Municipality

Protected area

Atelopus pictiventris

(Kattan, 1986)

CR

CR, B2ab(iii)(t)

Cal

PNN Farallones de

Cali, RFPN de Cali

Atelopus spurrelli

(Boulenger, 1914)

VU

EN,A1a

Bu, Dar

RFPN Anchicaya,

DMI La Plata, PNR La

sierpe

Incilius coniferus

(Cope, 1862)

LC

CR, B2ab(iii)

Dar

Rhaebo andi-

nophrynoides (Mue-

ses-Cisneros, 2009)

NE

DD

Bu

PNN Farallones de

Cali

Rhaebo blombergi

(Myers and Funkhous-

er, 1951)

NT

CR, Blab(iii)

Bu, Dar

PNN Farallones de

Cali, RFPN Anchicaya

Rhaebo haematiticus

(Cope, 1862)

LC

Bu, Da, Dar

RFPN Anchicaya,

RFPN de los rios San

Cipriano y Escalerete,

PNN Farallones de

Cali

Rhaebo hypomelas

(Boulenger, 1913)

NT

VU,A1ace

Bu, Da, Dar

RFPN Anchicaya,

RFPN de los rios San

Cipriano y Escalerete,

PNN Farallones de

Cali

Rhinella cf. mar-

garitifera (Laurenti,

1768)

LC

Bu, Dar

RFPN Anchicaya,

PNR La sierpe,

RFPN de los rios San

Cipriano y Escalerete,

DMI La Plata

Rhinella paraguas (Bo-

livar and Grant, 2014)

EN

VU

EN, B2ab(iii)

Cai

Rhinella marina (Lin-

naeus, 1758)

LC

Al, An, Ans,

Ar, Bo, Bu,

Bug, Bui, Ca,

Cal, Can, Car,

Da, Dar, Do,

Cai, Ce, Ag,

FI, Gi, Gu, Ja,

Cu, Vi, Un,

Ob, Pa, Pr,

Re, Ri, Ro,

Sp, Se, To, Tr,

Tu, Ul, Ve, Vij,

Yo, Yu, Za

RFPR Bitaco, RFN

Cerro Dapa-Carisu-

cio, PNN Tatama,

RFPN Anchicaya,

RFPN de Tulua,

RFPN de los rios San

Cipriano y Escalerete,

PNN Farallones de

Cali, RFPN de Cali,

RNR Laguna de

sonso 0 Cienaga de

chircal, RFPN del

Bosque de Yotoco,

FAMILIA CENTROLENIDAE

Centrolene ballux

(Duellman & Bur-

rowes, 1989)

CR

CR, B2ab(iii)

Da

PNN Farallones de

Cali

Centrolene buckleyi

(Boulenger, 1882)

VU

VU, B2ab(iii)

Cal, Cai, Tu,

Tr

PNN Farallones de

Cali, PNR Paramo El

Duende

Centrolene geckoi-

deum (Jimenez de la

Espada, 1872)

VU

CR, Alace

Cal, Cai, Da,

Cu

PNN Farallones de

Cali, RFPN Anchic-

aya, RFPR Bitaco

Centrolene heloderma

(Duellman, 1981)

CR

EN, B2ab(iii)

Cal, Cai

PNN Farallones de

Cali

Centrolene peristictum

(Lynch & Duellman,

1973)

VU

EN, B2ab(iii)

Cal, Cai, Da

PNN Farallones de

Cali, RFPN Anchicaya

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Conservation status of the herpetofauna in Colombia

TAXON

lUCN

Centrolene robledoi

(Ruiz-Carranza &

Lynch, 1995)

vu

Centrolene savage!

(Ruiz-Carranza &

Lynch, 1991)

vu

Cochranella balionota

(Duellman, 1981)

vu

Cochranella megista

(Rivero, 1985)

NT

Espadarana callis-

tomma (Guayasamin &

Trueb, 2007)

DD

Espadarana prosoble-

pon (Boettger, 1892)

LC

Hyalinobatrachium au-

reoguttatum (Barrera &

Ruiz-Carranza, 1989)

NT

Hyalinobatrachium

colymbiphyllum (Tay-

lor, 1949)

LC

Hyalinobatrachium

fleischmanni (Boettger,

1893)

LC

Hyalinobatrachium

valerioi (Dunn, 1931)

LC

Nymphargus armatus

(Lynch & Ruiz-Carran-

za, 1996)

VU

Nymphargus gran-

disonae (Cochran &

Coin, 1970)

LC

Nymphargus griffithsi

(Goin, 1961)

VU

Nymphargus ignotus

(Lynch, 1990)

NT

Nymphargus prasinus

(Duellman, 1981)

VU

Nymphargus ruizi

(Lynch, 1993)

VU

Rulyrana orejuela (Du-

ellman & Burrowes,

1989)

DD

Sachatamia albomacu-

lata (Taylor, 1949)

LC

Sachatamia ilex (Sav-

age, 1967)

LC

Red book of Red Book of

Colombia Valle del Cauca

Categorization for

the departament

Municipality

EN, B2ab(iii)

Cai, Da, Ri

NT

Bo, Cal, Da,

Cu, Yo

CR, B2ab(iii)

Da

CR, B2ab(iii)

Cai

NT

Bu

NT

Bu, Dar

VU,A3ace

Bu, Da, Dar

CR, Blab(iii)

Bu

VU,A3ace

Bu

VU,A3ace

Bu

CR, B2ab(iii)(t)

Cai

VU, Blab(iii)

Cai, Da

VU,A2ace

Cal, Cai, Da

NT

Cal, Cai, Da,

Cu

CR, B2ab(iii)

Da

VU,A2ace

Bug, Cal, Cai,

Da, Pa

EN, Blab(iii)

Da

EN, B2ab(iii)

Bu

NT

Bu, Da, Dar

Protected area

RFPN Anchicaya

PNN Farallones de

Cali, RFPN Anchic-

aya, RFPN de Cali,

RFPN Bosque de

Yotoco, RFPR Bitaco

RFPN de los rios San

Cipriano y Escale-

rete, RFPN Anchi-

caya

RFPN Anchicaya

PNN Farallones de

Cali, RFPN Anchi-

caya

PNN Farallones de

Cali, RFPN Anchic-

aya, RFPN de Cali,

RFPR Bitaco

PNN Farallones

de Cali, PNN Las

hermosas, RFPN

Anchicaya, RFPN de

Amaime

RFPN Anchicaya

RFPN Anchicaya,

DM I La Plata

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Valencia-Zuleta et al.

TAXON

lUCN

Teratohyla pulverata

(Peters, 1873)

LC

Teratohyla spinosa

(Taylor, 1949)

LC

FAMILIA CRAUGASTORIDAE

Red book of

Colombia

Red Book of

Valle del Cauca

Craugastor fitzingeri

(Schmidt, 1857)

LC

Craugastor longirostris

(Boulenger, 1898)

LC

Craugastor opimus

(Savage & Myers,

2002)

LC

Craugastor raniformis

(Boulenger, 1896)

LC

Hypodactylus babax

(Lynch, 1989)

LC

Hypodactylus manti-

pus (Boulenger, 1908)

LC

Pristimantis acatallelus

(Lynch & Ruiz-Carran-

za, 1983)

LC

Pristimantis achatinus

(Boulenger, 1898)

LC

Pristimantis alalo-

cophus (Roa-Trujillo &

Ruiz-Carranza, 1991)

NT

Pristimantis albericoi

(Lynch & Ruiz-Carran-

za, 1996)

CR

Pristimantis angustilin-

eatus (Lynch, 1998)

EN

Pristimantis aurantigut-

tatus (Ruiz-Carranza,

Lynch &Ardila-Ro-

bayo, 1997)

DD

Pristimantis boulengeri

(Lynch, 1981)

LC

Pristimantis brevifrons

(Lynch, 1981)

LC

Categorization for

the departament

Municipality

Protected area

VU, B2ab(iii)

Bu, Da

RFPN de los rios San

Cipriano y Escale-

rete, RFPN Anchi-

caya

NT

Bu, Da

RFPN de los rios San

Cipriano y Escale-

rete, RFPN Anchi-

caya

LC

Bu, Dar

LC

Bu, Dar

CR, B2ab(iii)

Bu, Dar

LC

Bu, Da, Dar

EN, B2ab(iii)

Da, Cai

NT

Cal, Cu, Da,

Dar, Tr, Yo, Yu

VU, B2ab(iii)

Cal, Cai

LC

Bu, Da, Dar

VU, B2ab(iii)

Bug, FI, Pa,

Tu

CR, B2ab(iii)

Cai

VU, B2ab(iii)

Cai, Tr

DD

Cai

VU, B2ab(iii)

Bug, FI, Pa

LC

Cal, Cai, Cu,

Da, FI, Tr, Yo

PNR La sierpe,

RFPN Anchicaya

PNR La sierpe,

RFPN Anchicaya,

RFPN de los rios San

Cipriano y Escalerete

PNR La sierpe

RFPN Anchicaya,

DMI Enclave subxe-

rofitico de Atuncela,

PNR La sierpe,

RFPN de los rios San

Cipriano y Escalerete

RFPN Anchicaya

PNN Farallones de

Cali, RFPN de Cali,

RFPR Bitaco, RFPN

Anchicaya, RFPN del

Bosque de Yotoco;

RFPN Cerro Dapa -

Carisucio

PNN Farallones de

Cali

RFPN Anchicaya,

DMI La Plata.

RFN Amaime, PNN

Las hermosas, RFPR

Frayle-Desbaratado

RFPR Frayle-Des-

baratado, PNN Las

hermosas

RFPR Bitaco, RFR

Frayle-Desbaratado,

PNN Farallones

de Cali, RFPN del

Bosque de Yotoco,

RFPN Cerro Dapa -

Carisucio, RFPN de

Cali, RFPN Anchi-

caya

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Conservation status of the herpetofauna in Colombia

TAXON

lUCN

Red book of

Colombia

Red Book of

Valle del Cauca

Categorization for

the departament

Municipality

Protected area

Pristimantis buckleyi

(Boulenger 1882)

LC

NT

Cal, Cai, Ce,

FI, Pa, Tu

RFN Amaime, PNN

Farallones de Cali,

RFPR Frayle-Des-

baratado.

Pristimantis calcaratus

(Boulenger, 1908)

EN

NT

Cal, Cai, Cu,

Da, Yo

RFPN de Cali, PNN

Farallones de Cali,

RFPR Bitaco, RFPN

del Bosque de Yotoco

Pristimantis calcarula-

tus (Lynch, 1976)

VU

VU, B2ab(iii)

Cal, Cum

PNN Farallones de

Cali, RFPR Bitaco

Pristimantis capitonis

(Lynch, 1998)

EN

VU

VU, B2ab(iii)

Cal, Cai, Da,

Tr

PNN Farallones de

Cali, RFPNAnchi-

caya

Pristimantis caprifer

(Lynch, 1977)

LC

VU, B2ab(iii)

Bu, Da, Dar

RFPN Anchicaya

Pristimantis chalceus

(Peters, 1873)

LC

NT

Bu, Da, Dar

RFPN Anchicaya

Pristimantis chrysops

(Lynch & Ruiz-Carran-

za, 1996)

EN

EN, B2ab(iii)

Cal, Cai, Da,

Yo

RFPN Anchicaya,

RFPN de Cali, RFPN

del Bosque de Yotoco

Pristimantis deinops

(Lynch, 1996)

EN

EN, B2ab(iii)(t)

Cal, Cai, Da,

Yo

RFPN Anchicaya,

RFPN de Cali, RFPN

del Bosque de Yotoco

Pristimantis diaphonus

(Lynch, 1986)

VU

EN, B2ab(iii)(t)

Da, Re

Pristimantis duende

(Lynch, 2001)

DD

CR, B2ab(iii)(t)

Tr

PNR Paramo el

Duende

Pristimantis erythro-

pieura (Boulenger,

1896)

LC

Cal, Cai, Cu,

Da, Dar, Tr,

Yo

RFPR Bitaco, RFPN

de Cali, RFPN Anchi-

caya, PNN Farallones

de Cali, RFPN del

Bosque de Yotoco

Pristimantis gaigeae

(Dunn, 1931)

LC

NT

Bu

RFPN Anchicaya,

DMI La Plata, RFPN

de los rios San

Cipriano y Escalerete

Pristimantis gracilis

(Lynch, 1986)

VU

CR, B2ab(iii)

Cal, Cai, Cu

PNN Farallones de

Cali, RFPR Bitaco

Pristimantis hybotra-

gus (Lynch, 1992)

VU

VU, B2ab(iii)(t)

Bu, Dar

PNR La sierpe,

RFPN Anchicaya,

RFPN de los rios San

Cipriano y Escalerete

Pristimantis juanchoi

(Lynch, 1996)

VU

NT

Cal, Cai, Cu,

Da, Dar, Yo

RFPR Bitaco, RFPN

Anchicaya, RFPN

de Cali, RFPN del

Bosque de Yotoco.

Pristimantis keiephas

(Lynch, 1998)

VU

CR, B2ab(iii)

Cai

Pristimantis labiosus

(Lynch, Ruiz-Carranza

& Ardila-Robayo,

1994)

LC

NT

Bu, Dar

RFPN de los rios San

Cipriano y Escale-

rete, PNN Farallones

de Cali

Pristimantis latidiscus

(Boulenger, 1898)

LC

Bu, Dar

RFPN de los rios San

Cipriano y Escale-

rete, RFPN Anchic-

aya, DMI La Plata

Pristimantis molybri-

gnus (Lynch, 1986)

NT

VU, B2ab(iii)

Cal, Da, Dar,

Yo

RFPN Anchicaya,

RFPN de Cali, RFPN

del Bosque de Yotoco

Amphib. Reptile Conserv.

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Valencia-Zuleta et al.

TAXON

Pristimantis moro

(Savage, 1965)

Pristimantis myops

(Lynch, 1998)

Pristimantis obmutes-

cens (Lynch, 1980)

Pristimantis orpaco-

bates (Lynch, Ruiz-

Carranza SArdila-

Robayo, 1994)

Pristimantis palmeri

(Boulenger, 1912)

Pristimantis peraticus

(Lynch, 1980)

Pristimantis permixtus

(Lynch, Ruiz-Carranza

& Ardila-Robayo,

1994)

Pristimantis phalarus

(Lynch, 1998)

Pristimantis piceus

(Lynch, Ruiz-Carranza

& Ardila-Robayo,

1996)

Pristimantis platychiius

(Lynch, 1996)

Pristimantis ptochus

(Lynch, 1998)

Pristimantis quantus

(Lynch, 1998)

Pristimantis qui-

cato (Ospina-Sarria,

Mendez-Narvaez,

Burbano-Yandi &

Bolivar-Garcia, 2011)

Pristimantis racemus

(Lynch, 1980)

Pristimantis restrepoi

(Lynch, 1996)

Pristimantis ridens

(Cope, 1866)

Pristimantis roseus

(Boulenger, 1918)

Pristimantis sanguin-

eus (Lynch, 1998)

Pristimantis silver-

stonei (Lynch & Ruiz-

Carranza, 1996)

lUCN

LC

DD

Red book of

Colombia

Red Book of

Valle del Cauca

Categorization for

the departament

Municipality

VU, B2ab(iii)

Bu

VU, B2ab(iii)

Cal, Cai, Da

VU, B2ab(iii)

FI, Gi, Tu

VU, Blab(iii)

Cal, Cu, Dar,

Yo

LC

Cal, Cai, Cu,

Da, Dar, Yo,

Yu

VU, Blab(iii)

Ce, FI, Pa

NT

Ce, Cu, FI, Pa

VU, Blab(iii)

Cai

VU, Blab(iii)

Ce, FI, Pa

VU, Blab(iii)

Cal, Da

DD

Cal, Cai

VU, Blab(iii)

Cai

DD (t)

Pa

VU, Blab(iii)

Ce, Gi, Pa, Tu

VU, Blab(iii)

Cai

NT

Bu, Dar

VU, B2ab(iii)

Bu, Da, Dar

DD

Bu

VU, B2ab(iii)

Cai, Da

Protected area

RFPN Anchicaya,

PNR La sierpe

PNN Farallones de

Cali

RFPR Frayle-Des-

baratado, RFPN de

sonso-guabas

RFPR Bitaco, RFPN

de Cali, RFPN del

Bosque de Yotoco

PNN Farallones de

Cali, RFPR Bitaco,

RFPN de Cali, RFPN

del Bosque de

Yotoco, RFPN Cerro

Dapa - Carisucio

RFPN Amaime,

RFPR Frayle-Des-

baratado

RFPR Bitaco, RFPR

Frayle-Desbaratado

RFPN Amaime,

RFPR Frayle-Des-

baratado

RFPN de Cali, RFPN

Anchicaya, RFPR

Bitaco

PNN Farallones de

Cali

PNN Las hermosas

RFPN de sonso-

guabas

PNR La sierpe,

RFPN Anchicaya

PNR La sierpe,

RFPN Anchicaya,

RFPN de los rios San

Cipriano y Escalerete

PNN Farallones de

Cali, PNR Paramo

del Duende

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December 2014 | Volume 8 | Number 2 | e87

Conservation status of the herpetofauna in Colombia

TAXON

lUCN

Red book of

Colombia

Red Book of

Valle del Cauca

Categorization for

the departament

Municipality

Protected area

Pristimantis simoteris-

cus (Lynch, Ruiz-Car-

ranza & Ardila-Robayo

1996)

EN

EN, B2ab(iii)

Gi, Tu

RFPN de sonso-

guabas

Pristimantis supernatis

(Lynch, 1979)

VU

VU, B2ab(iii)

Ce, Gi, Pa, Tu

RFPN de sonso-

guabas

Pristimantis taeniatus

(Boulenger, 1912)

LC

VU, B2ab(iii)

Bu

DMI La Plata

Pristimantis thectopter-

nus (Lynch, 1975)

LC

VU,A1ace

Cal, Cu, Da,

Dar, FI, Pa, Tr

RFPN de Cali, RFPR

Bitaco, RFPN Anchi-

caya

Pristimantis urano-

bates (Lynch, 1991)

LC

EN, B2ab(iii)

Ce, Pa

RFPN de Amaime

Pristimantis viridicans

(Lynch 1977)

EN

VU

CR, Alace

Cal, Cu

PNN Farallones de

Cali, RFPR Bitaco

Pristimantis w-nigrum

(Boettger, 1892)

LC

NT

Cal, Cu, Tr,

Yo

PNN Farallones de

Cali, RFPR Bitaco,

RFPN de Cali, RFPN

del Bosque de Yotoco

Pristimantis xeniolum

(Lynch, 2001)

DD

DD (t)

Tr

PNR Paramo el

Duende

Pristimantis xylocho-

bates (Lynch & Ruiz-

Carranza,1996)

VU

EN, Alace

Cal, Cai

PNN Farallones de

Cali

Strabomantis anatipes

(Lynch and Myers,

1983)

VU

CR, Alace

Da

RFPN Anchicaya

Strabomantis anoma-

/ws (Boulenger, 1898)

LC

Bu, Da, Dar

RFPN Anchicaya

Strabomantis bufoni-

formis (Boulenger,

1896)

LC

VU, Alace

Bu, Dar

PNN Farallones de

Cali, RFPN Anchi-

caya

Strabomantis cerastes

(Lynch, 1975)

LC

VU, Alace

Cal, Cai, Da,

Yo

RFPN de Cali, RFPN

del Bosque de Yotoco

Strabomantis cheirop-

lethus (Lynch, 1990)

VU

VU, Alace

Bu, Cai, Da

PNN Farallones de

Cali.

Strabomantis ruizi

(Lynch, 1981)

EN

VU, Alace (t)

Cal, Cu, Da,

Tr, Yo

RFPR Bitaco, PNN

Farallones de Cali,

RFPN del Bosque de

Yotoco

Strabomantis zygo-

dactylus (Lynch and

Myers, 1983)

LC

NT

Bu, Dar

RFPN de los rios San

Cipriano y Escalerete

RFPN Anchicaya

FAMILIA DENDROBATIDAE

Andinobates bombetes

(Myers & Daly, 1 980)

EN

VU

EN

VU, Blab(iii)

Cal, Cai, Cu,

Da, Dar, To,

Yo

RFPN Anchicaya,

RFPN de Cali, RFPN

Bosque de Yotoco,

RFPR Bitaco, DMI

Enclave subxerofitico

de Atuncela

Andinobates fulguritus

(Silverstone, 1975)

LC

NT

VU, Blab(iii)

Bu

RFPN de los rios San

Cipriano y Escale-

rete, RFPN Anchic-

aya, PNR La Sierpe

Andinobates minutus

(Shreve, 1935)

LC

Bu, Dar

PNN Uramba-Bahia

Malaga, RFPN de

los rios San Cipriano

y Escalerete, RFPN

Anchicaya, PNR La

Sierpe, DMI La plata

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Valencia-Zuleta et al.

TAXON

lUCN

Red book of

Colombia

Red Book of

Valle del Cauca

Categorization for

the departament

Municipality

Protected area

Andinobates viridis

(Myers & Daly, 1976)

vu

EN

VU

EN,A1acde (t)

Bu

RFPN de los rios San

Cipriano y Escale-

rete, RFPN Anchi-

caya

Colostethus agilis

(Lynch & Ruiz-Carran-

za, 1985)

NT

EN, B2ab(iii)

Cal, Cai, Da

PNN Farallones de

Cali, RFPNAnchi-

caya

Colostethus brachistri-

atus (Rivero & Serna,

1986)

DD

VU,A1ace

FI, Pa, Tu

PNN Las Hermosas,

RFPR Frayle-Des-

baratado

Colostethus fraterdan-

ieli (Silverstone, 1971)

NT

Bug, Cal, Cu,

Da, FI, Ri,

Tu, Yo

PNN Farallones de

Cali, RFPN Anchic-

aya, RFPN Bosque

de Yotoco, RFN de

Tulua, RN Laguna de

Sonso

Epipedobates bou-

/enger/ (Barbour, 1909)

LC

Bu

RFPN de los rios San

Cipriano y Escale-

rete, RFPN Anchic-

aya, DMI La plata

Hyloxalus abditauran-

t/ws (Silverstone, 1975)

LC

VU, B2ab(iii)

Cal, Cai, Da,

Dar

PNN Farallones de

Cali, RFPNAnchi-

caya

Hyloxalus chocoensis

(Boulenger, 1912)

DD

EN, B2ab(iii)

Da, Dar

RFPN Anchicaya

Hyloxalus fascianigrus

(Grant & Castro-Herre-

ra, 1998)

NT

VU, B2ab(iii)

Cal, Cu, Da,

Dar, Yo

RFPN Anchicaya,

RFPN Bosque de

Yotoco, RFPN de

Cali, PNR Paramo

El Duende, RFPR

Bitaco, RFPR Frayle-

Desbaratado

Hyloxalus lehmanni

(Silverstone, 1971)

NT

VU,A1ace

Bug, Cal, Cai,

Cu, Tu

PNN Farallones de

Cali, RFPN Anchic-

aya, RFPR Bitaco,

RFN de Amaime

Oophaga histrionica

(Berthold, 1845)

LC

VU

VU,A1acde (t)

Bu, Da, Dar

RFPN Anchicaya,

RFPN Dagua

Oophaga lehmanni

(Myers & Daly, 1976)

CR

CR, A2acde

Bu, Da

RFPN Anchicaya

Phyllobates bicolor

(Dumeril & Bibron,

1841)

NT

EN, Alacde

Bu, Cai

Phyllobates cf. au-

rotaenia (Boulenger,

1913)

NT

EN, A2acde

Bu

RFPN Anchicaya

Phyllobates terribilis

(Myers, Daly & Malkin,

1978)

EN

CR

CR, Blab(iii)

Bu

Silverstoneia nubicola

(Dunn, 1924)

NT

Bu, Dar

RFPN de los rios San

Cipriano y Escale-

rete, RFPN Anchi-

caya

FAMILIA ELEUTHERODACTYLIDAE

Diasporus tinker

(Lynch, 2001)

LC

Bu, Dar

PNR La sierpe, DMI

La Plata, RFPN

Anchicaya, RFPN de

los rios San Cipriano

y Escalerete

Amphib. Reptile Conserv.

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Conservation status of the herpetofauna in Colombia

TAXON

lUCN

Red book of

Colombia

Red Book of

Valle del Cauca

Categorization for

the departament

Municipality

Protected area

Diasporus gularis

(Boulenger, 1898)

LC

Bu, Da, Dar

DMI La Plata, RFPN

Anchicaya, PNN

Uramba Bahia

Malaga, RFPN de los

rios San Cipriano y

Escalerete

Diasporus quidditus

(Lynch, 2001)

LC

DD

Bu, Dar

Eleutherodactylus

johnstonei (Barbour,

1914)

LC

Cal, Ja, Yu

FAMILIA HEMIPHRACTIDAE

Gastrotheca angus-

tifrons (Boulenger,

1898)

VU

CR, Alace

Dar

Gastrotheca anto-

mia (Ruiz-Carranza,

Ardila-Robayo, Lynch

& Restrepo, 1997)

VU

CR, Alace

Cal, Cai, Cu,

Da

RFPN de Cali, RFPN

Anchicaya, RFPR

Bitaco

Gastrotheca argenteo-

virens (Boettger, 1892)

LC

EN, Alace

Ce, Pa, Tu

PNN Las hermosas,

RFN de Amaime

Gastrotheca cornuta

(Boulenger, 1898)

EN

VU

EN, Alace

Bu

RFPN Anchicaya,

RFPN de los rios San

Cipriano y Escalerete

Gastrotheca dendro-

nastes (Duellman,

1983)

VU

CR, Alace

Cal, Cai, Da,

Dar, Yo

RFPN Anchicaya,

PNN Farallones

de Cali, RFPN del

Bosque de Yotoco

Hemiphractus fascia-

tus (Peters, 1862)

NT

EN

EN, B2ab(iii)

Bu, Dar

FAMILIA HYLIDAE

Agalychnis psilopygion

(Cannatella, 1980)

DD

EN, B2ab(iii)

Bu, Dar

RFPN Anchicaya

Agalychnis spurrelli

(Boulenger, 1913)

LC

NT

Bu, Dar

PNN Farallones de

Cali, RFPN Anchi-

caya

Cruziohyla calcarifer

(Boulenger, 1902)

LC

VU,A2acde

Bu, Dar

RFPN Anchicaya

Dendropsophus

columbianus (Boettger,

1892)

LC

Al, An, Ans,

Ar, Bo, Bug,

Bui, Ca, Cal,

Can, Car, Da,

Dar, Do, Cai,

Ce, Ag, FI, Gi,

Gu, Ja, Cu,

Vi, Un, Ob,

Pa, Pr, Re, Ri,

Ro, Sp, Se,

To, Tr, Tu, Ul,

Ve, Vij, Yo,

Yu,Za

RFPN Cerro Dapa

- Carisucio, PNN

Tatama, PNN Faral-

lones de Cali, PNN

Las Flermosas,

RFPR Bitaco, RFPN

de Amaime, RFPN

de Cali

Dendropsophus ebrac-

catus (Cope, 1874)

LC

VU, B2ab(iii)

Bu

RFPN de los rios San

Cipriano y Escalerete

Dendropsophus micro-

cephalus (Boulenger,

1898)

LC

EN, B2ab(iii)

Bu

RFPN de los rios San

Cipriano y Escalerete

Hyloscirtus alytolylax

(Duellman, 1972)

NT

EN, B2ab(iii)

Cal, Cai, Cu,

Da

RFPR BITACO,

RFPN Anchicaya,

RFPN de Cali

Amphib. Reptile Conserv.

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December 2014 | Volume 8 | Number 2 | e87

Valencia-Zuleta et al.

TAXON

lUCN

Hyloscirtus larinopy-

gion (Duellman, 1973)

NT

Hyloscirtus palmeri

(Boulenger 1908)

LC

Hyloscirtus simmonsi

(Duellman, 1989)

EN

Hypsiboas boans (Lin-

naeus, 1758)

LC

Hypsiboas pellucens

(Werner, 1901)

LC

Hypsiboas picturatus

(Boulenger, 1899)

LC

Hypsiboas pugnax

(Schmidt, 1857)

LC

Hypsiboas rosenbergi

(Boulenger, 1898)

LC

Hypsiboas rubrac-

ylus (Cochran & Coin,

1970)

LC

Scinax elaeochroa

(Cope, 1875)

LC

Scinax ruber (Laurenti,

1768) (sensu latu)

LC

Scinax sugillatus (Du-

ellman, 1973)

LC

Smilisca phaeota

(Cope, 1862)

LC

Red book of

Colombia

Red Book of

Valle del Cauca

FAMILIA LEPTODACTYLIDAE

Leptodactylus colom-

biensis (Fleyer, 1994)

LC

Leptodactylus fragilis

(Brocchi, 1877)

LC

Leptodactylus melano-

notus (Flallowell, 1861)

LC

Leptodactylus rhodo-

merus (Heyer, 2005)

LC

Categorization for

the departament

Municipality

VU,A1ace

Cal, Cai, Ce,

Cu, Da, FI, Ri

NT

Bu, Da, Dar

CR, Blab(iii)

Dar

NT

Bu, Da, Dar

VU, B2ab(iii)

Bu

LC

Bu, Dar

LC

Bu, Car, Ja,

Rol

LC

Bu, Dar

VU, Blab(iii)

Bu

VU, B2ab(iii)

Bu

LC

Bu, Ja

EN, B2ab(iii)

Bu

LC

Bu, Dar

Protected area

RFPR Bitaco, PNN

Farallones de Cali,

RFPN Anchicaya,

RFPN de Cali, RFPN

de Amaime

PNN Farallones de

Cali, RFPN Anchi-

caya

DMI La Plata, RFPN

Anchicaya, RFPN de

los rios San Cipriano

y Escalerete

RFPN Anchicaya

PNR La sierpe, DMI

La Plata, RFPN

Anchicaya, RFPN de

los rios San Cipriano

y Escalerete

RFPN Anchicaya

RFPN Anchicaya,

RFPN de los rios San

Cipriano y Escalerete

PNR La sierpe,

RFPN Anchicaya

RFPN de los rios San

Cipriano y Escalerete

RFPN Anchicaya

PNR La sierpe, PNN

Farallones de Cali,

DMI La Plata, RFPN

Anchicaya, RFPN de

los rios San Cipriano

y Escalerete

LC

An, Ans, Bo,

Bug, Bui, Cal,

Can, Ce, FI,

Gu, Ja, Ob,

Pa, Un, Ri,

Ro, Sp, To, Tr,

Tu, Vi, Vij, Yo,

Yu,Za

RNR Laguna de

Sonso

LC

An, Bug, Cal,

Ja, Yu

RNR Laguna de

Sonso

LC

Bu

RFPN Anchicaya,

PNR La Sierpe

LC

Bu, Dar

RFPN de los rios

Escalerete y San

Cipriano, RFPN

Anchicaya, DMI La

Plata

Amphib. Reptile Conserv.

December 2014 | Volume 8 | Number 2 | e87

Conservation status of the herpetofauna in Colombia

TAXON

lUCN

Leptodactylus ventri-

maculatus (Boulenger,

1902)

LC

Leptodactylus wagneri

(Peters, 1862)

LC

FAMILIA MICROHYLIDAE

Nelsonophryne ater-

rima (Gunther, 1900)

LC

FAMILIA RANIDAE

Lithobates catesbei-

ana (Shaw, 1802)

LC

Lithobates vaillanti

(Brocchi, 1877)

LC

Red book of

Colombia

Red Book of

Valle del Cauca

ORDEN CAUDATA

FAMILIA PLETHODONTIDAE

ORDEN GYMNOPHIONA

FAMILIA CAECILIIDAE

Bolitoglossa biseriata

(Tanner, 1962)

LC

Bolitoglossa walker!

(Brame &Wake, 1972)

NT

Bolitoglossa hiemalis

(Lynch, 2001)

DD

Bolitoglossa vallecula

(Brame &Wake, 1963)

VU

Bolitoglossa silver-

stone! (Brame & Wake,

1972)

LC

Oedipina parvipes

(Peters, 1879)

LC

Caecilia guntheri

(Dunn, 1942)

DD

Caecilia leucocephala

(Taylor, 1968)

LC

Caecilia nigricans

(Boulenger, 1902)

LC

Caecilia occidentalis

(Taylor, 1968)

DD

Caecilia orientalis

(Taylor, 1968)

LC

Caecilia perdita (Tay-

lor, 1968)

LC

Caecilia subdermalis

(Taylor, 1968)

LC

Caecilia subnigricans

(Dunn, 1942)

LC

Caecilia cf. tentaculata

(Linnaeus, 1749)

LC

Oscaecilia polyzona

(Fisher, 1880)

DD

Categorization for

the departament

Municipality

NT

Bu, Dar

NT

Bu

Protected area

EN, B2ab(iii)

LC

Bug, Gi, Pa,

Ri, Ro, Tu, Yo

RN Laguna de Sonso

LC

Bu

RFPN de los rios

Escalerete y San

Cipriano, RFPN

Anchicaya

NT

Bu, Da

VU, B2ab(iii)

Cal, Cai, Cu

CR, B2ab(iii)(t)

Tr

EN, B2ab(iii)

Cai

DD

Bu

DD

Bu

RFPN de los rios

Escalerete y San

Cipriano, RFPN

Anchicaya

RFPN de Cali, RFPR

Bitaco

PNR Paramo El

Duende

RFPN Anchicaya

DD

Cai

DD

Bu

DD

Bu

DD

Cal, Da

DD

Se

DD

Bu

DD

Cu, Da, Ja

DD

Re

DD

Bu, Re

DD

Bu

RFPN Anchicaya

RFPN de los rios San

Cipriano y Escalerete

PNN Farallones de

Cali

RFPN Anchicaya,

RFPR Bitaco

RFPN de los rios San

Cipriano y Escalerete

PNR La sierpe

Amphib. Reptile Conserv.

December 2014 | Volume 8 | Number 2 | e87

Valencia-Zuleta et al.

TAXON lUCN

FAMILIA RHINATREMATIDAE

LC

FAMILIA TYPHLONECTIDAE

Epicrionops bicolor

(Boulenger, 1883)

Typhlonectes natans

(Fisher, 1880)

CLASS REPTILIA

ORDEN CROCODYLIA

FAMILIA ALLIGATORIDAE

Caiman crocodilus

(Linnaeus, 1758)

FAMILIA CROCODYLIDAE

Crocodylus acutus

(Cuvier, 1807)

ORDEN SQUAMATA

SUBORDEN AMPHISBAENIA

FAMILIA AMPHISBAENIDAE

Amphisbaena fuligi-

nosa (Linnaeus, 1758)

SUBORDEN SAURIA

INFRAORDEN IGUANIA

FAMILIA CORYTOPHANIDAE

FAMILIA DACTYLOIDAE

Red book of

Colombia

Red Book of

Valle del Cauca

Basiliscus basiliscus

(Linnaeus, 1768)

NE

Basiliscus galeritus

(Dumeril, 1851)

NE

Anolis agassizi (Stej-

neger, 1900)

NE

Anolis anchicayae

(Poe, Velasco, Miyata

& Williams, 2009)

NE

Anolis antonii (Bou-

lenger, 1908)

NE

Anolis auratus

(Daudin, 1802)

NE

Anolis biporcatus

(Wiegmann, 1834)

NE

Anolis calimae (Ayala,

Harris & Williams,

1983)

NE

Anolis chloris (Bou-

lenger, 1898)

NE

Categorization for

the departament

Municipality

NT

Bo, Cu, Da

NT

An, Ans, Bo,

Bug, Bui, Cal,

Can, Car, FI,

Gu, Ja, Un,

Vi, Ob, Ri,

Ro, Sp, To,

Tr, Tu, Vij, Yo,

Yu,Za

VU,A1acd

Protected area

RFPR Bitaco, PNN

Farrallones de Cali

Bu, Bug RN Laguna de sonso

RFPN Anchicaya

NT

Bu, Bug, Ja,

Pa, Yu

RFPN Anchicaya,

RNR Laguna de

Sonso

NT

Bu, Da

PNN Uramba-Bahia

Malaga, RFPN Anchi-

caya

CR, B2ab(iii)(t)

Bu

VU, B2ab(iii)

Bu, Dar

LC

Cal, Cai, Cu,

Ce, Da, Yo

LC

Bu, Bug, Cal,

Ce, Ja, Pa,

Yu

VU, B2ab(iii)

Bu

VU, B2ab(iii)

Cal, Da, Vi

VU, B2ab(iii)

Bu

SFF Isla Malpelo

RFPN de los rios San

Cipriano y Escale-

rete, RFPN Anchi-

caya

PNN Farallones de

Cali, RFPN Anchic-

aya, RFPN de Cali,

RFPN Bosque de

Yotoco, RFPR Bitaco

PNN Farallones de

Cali

RFPN Anchicaya

Amphib. Reptile Conserv.

December 2014 | Volume 8 | Number 2 | e87

Conservation status of the herpetofauna in Colombia

TAXON

lUCN

Anolis chocorum

(Williams & Duellman,

1967)

NE

Anolis eulaemus (Bou-

lenger, 1908)

LC

Anolis fraseri (Gunther,

1859)

LC

Anolis gracilipes (Bou-

lenger, 1898)

NE

Anolis granuliceps

(Boulenger, 1898)

LC

Anolis heterodermus

(Dumeril, 1851)

NE

Anolis latifrons (Ber-

thold, 1846)

NE

Anolis lyra (Poe,

Velasco, Miyata & Wil-

liams, 2009)

NE

Anolis macrolepis

(Boulenger, 1911)

NE

Anolis maculiventris

(Boulenger, 1898)

NE

Anolis mirus (Williams,

1963)

NE

Anolis notopholis

(Boulenger, 1896)

NE

Anolis propinquus

(Williams, 1984)

NE

Anolis rivalis (Williams,

1984)

NE

Anolis ventrimaculatus

(Boulenger, 1911)

NT

Red book of

Colombia

Red Book of

Valle del Cauca

FAMILIA HOPLOCERCIDAE

Enyalioides heterol-

epis (Bocourt, 1874)

NE

Enyalioides oshaugh-

nessyi (Boulenger,

1881)

NE

FAMILIA IGUANIDAE

Iguana iguana (Lin-

naeus, 1758)

NE

FAMILIA POLYCHROTIDAE

Polychrus gutturosus

(Berthold, 1846)

NE

Categorization for

the departament

Municipality

VU, A2ac

Bu

VU,A2ac

Cal, Cu, Vi

VU, B2ab(iii)

Cal, Da, Yo

EN, B2ab(iii)

Bu

VU, B2ab(iii)

Bu, Da

VU, B2ab(iii)

Cal, Cai, Cu,

Da

NT

Bu

VU, B2ab(iii)

Bu, Dar

NT

Bu

NT

Bu

CR, Blab(iii)

Bu

NT

Bu, Da, Dar

CR, Blab(iii)

Da, Dar

DD

Bu

LC

Cal, Cai, Cu,

Da

EN,A2ac

Bu, Bug, Cal,

Can, Ce, Cu,

Da, Gu, Ja,

Pa, Sp, Vi,

Vij, Yu

Bu, Dar

Protected area

RFPN de los rios San

Cipriano y Escale-

rete, RFPN Anchi-

caya

PNN Farallones de

Cali, RFPR Bitaco

PNN Farallones de

Cali, RFPN Anchic-

aya, RFPN Bosque

de Yotoco

RFPN Anchicaya

RFPN Anchicaya,

RFPR Bitaco

RFPN Anchicaya

RFPN de los rios San

Cipriano y Escale-

rete, RFPN Anchi-

caya

RFPN Anchicaya

NT

Bu, Da

DD

Bu

PNN Farallones de

Cali, RFPN Anchic-

aya, RFPN de Cali,

RFPR Bitaco

RFPN Anchicaya

RFPN de los rios

Escalerete y San

Cipriano, RN Laguna

de Sonso

RFPN de los rios San

Cipriano y Escalerete

Amphib. Reptile Conserv.

December 2014 | Volume 8 | Number 2 | e87

Valencia-Zuleta et al.

TAXON

lUCN

Red book of

Colombia

Red Book of

Valle del Cauca

INFRAORDEN GEKKOTA

FAMILIAGEKKONIDAE

Hemidactylus brookii

(Gray, 1845)

NE

Hemidactylus frenatus

(Schlegel, 1836)

LC

Lepidodactylus lugu-

bris (Dumeril & Bibron,

1836)

NE

FAMILIA PHYLLODACTYLIDAE

Phyllodactylus trans-

versalis {Huey, 1935)

NE

Thecadactylus

rapicauda (Flouttuyn,

1782)

NE

FAMILIA SPHAERODACTYLIDAE

Gonatodes albogularis

(Dumeril & Bibron,

1836)

NE

Lepidoblepharis duol-

epis (Ayala & Castro,

1983)

NE

Lepidoblepharis inter-

medius (Boulenger,

1914)

NE

Lepidoblepharis micro-

/ep/s (Noble, 1923)

NE

Lepidoblepharis perac-

cae (Boulenger, 1908)

NE

INFRAORDEN SCINCOMORPHA

FAMILIA GYMNOPHTHALMIDAE

Alopoglossus festae

(Peracca, 1904)

NE

Alopoglossus lehm-

anni (Ayala & Flarris,

1984)

NE

Anadia vittata (Bou-

lenger, 1913)

NE

Cercosaura argulus

(Peters, 1863)

LC

Cercosaura vertebra-

lis (O’Shaughnessy,

1879)

NE

Echinosaura horrida

(Boulenger, 1890)

NE

Categorization for

the departament

Municipality

Protected area

LC

An, Bu, Bui,

Cal, Can, Gu,

Ja, Ob, Pa,

Yu

LC

An

LC

Bu, Cal, Yu

rete, RFPN Anchi-

caya

RFPN de los rios San

Cipriano y Escale-

rete, RFPN Anchi-

caya

CR, B2ab(iii)(t)

Bu

SFF Isla Malpelo

LC

Bu, Dar

RFPN de los rios San

Cipriano y Escalerete

LC

Bu, Bug, Cal,

Cu, Ja, Pa,

To, Tu, Vi, Vij,

Yo, Yu, Za

NT

Cal, Tr, Yo

VU, B2ab(iii)

Bu

VU, B2ab(iii)

Bu

VU, B2ab(iii)

Bu

RN Laguna de Sonso

RFPN Bosque de

Yotoco, RFPN de Cali

RFPN Anchicaya

EN, B2ab(iii)

Bu, Dar

EN, B2ab(iii)

Bu, Dar

VU, B2ab(iii)

Bu

NT

Cal, Da, FI,

Pa, Yo

LC

Cal, Cai, Cu,

Da, To, Tr

NT

Bu

RFPN de los rios San

Cipriano y Escalerete

PNN Farallones de

Cali, RFPN Anchi-

caya

PNN Farallones de

Cali, PNN Las hermo-

sas, RFPN Anchic-

aya, RFPN Bosque

de Yotoco, RFPR

Frayle-Desbaratado

PNN Farallones de

Cali, RFPN Anchic-

aya, RFPN de Cali,

RFPR Bitaco

RFPN de los rios San

Cipriano y Escale-

rete, RFPN Anchi-

caya

Amphib. Reptile Conserv.

December 2014 | Volume 8 | Number 2 | e87

Conservation status of the herpetofauna in Colombia

TAXON

lUCN

Echinosaura orcesi

(Fritts, Almendariz &

Samec, 2002)

NE

Gymnophthalmus

speciosus (Flallowell,

1861)

NE

Leposoma south!

(Ruthven & Gaige,

1924)

NE

Ptychoglossus stenol-

ep/s (Boulenger, 1908)

LC

Ptychoglossus vallen-

s/s(Flarris, 1994)

NE

Riama Columbiana

(Andersson, 1914)

NE

Riama laevis (Bou-

lenger, 1908)

NE

Riama striata (Peters,

1863)

NE

Red book of

Colombia

Red Book of

Valle del Cauca

FAMILIATEIIDAE

Ameiva ameiva (Lin-

naeus, 1758)

NE

Cnemidophorus

lemniscatus (Linnaeus,

1758)

NE

Holcosus anomalus

(Echternacht, 1977)

NE

FAMILIASCINCIDAE

Mabuya sp. {sensu

latu)

INFRAORDEN DIPLOGLOSSA

FAMILIAANGUIDAE

Diploglossus mil-

lepunctatus

(O'Shaughnessy,

1874)

Diploglossus monotro-

p/s (Kuhl, 1820)

SUBORDEN SERPENTES

FAMILIA BOIDAE

Boa constrictor (Lin-

naeus, 1758)

NE

Corallus annulatus

(Cope, 1876)

NE

FAMILIA COLUBRIDAE

Chironius carinatus

(Linnaeus, 1758)

NE

Chironius grandisqua-

mis (Peters, 1868)

NE

Categorization for

the departament

Municipality

VU, B2ab(iii)

Bu

DD

Bug, Ja, Ro,

Yu

NT

Bu

NT

Cai, Da, Tr,

Yo

DD

Cal

VU,A2acd

Cal, Cai, Tu

EN,A2acd

Cai, Cu

DD

Cal

VU, B2ab(iii)

Protected area

RFPN Anchicaya

RFPN de los rios San

Cipriano y Escale-

rete, RFPN Anchi-

caya

RFPN Anchicaya

RFPN de Cali

PNN Farallones de

Cali, RFPN de Tulua

RFPR Bitaco

PNN Farallones de

Cali

VU, B2ab(iii)

Cal, Ja, Pa

LC

Bug, Cal, Ja,

Pa, Vij, Yo, Yu

NT

Bu, Da, Dar

Bu, Cal

RN Laguna de Sonso

RFPN de los rios San

Cipriano y Escale-

rete, RFPN Anchi-

caya

PNN Farallones de

Cali, RFPN de Cali

CR, B1ab(iii)(t)

Bu

SFF Isla Malpelo

VU, B2ab(iii)

Bu, Dar

RFPN Anchicaya

VU,A2acd

Bu

EN, Blab(iii)

Bu

RFPN Anchicaya

NT

Bu, Cal

PNN Farallones de

Cali, RFPN de Cali

NT

Bu

RFPN Anchicaya,

PNR La Sierpe,

RFPN de los rios

Escalerete y San

Cipriano

Amphib. Reptile Conserv.

December 2014 | Volume 8 | Number 2 | e87

Valencia-Zuleta et al.

TAXON

lUCN

Chironius monticola

(Roze, 1952)

NE

Dendrophidion bivit-

tatus (Dumeril, Bibron

& Dumeril, 1854)

NE

Dendrophidion clarkii

(Dunn, 1933)

NE

Dendrophidion percari-

natus (Cope, 1893)

NE

Dendrophidion pro-

lixum (Cadle, 2012)

NE

Drymarchon melanu-

rus (Bibron & Dumeril

1854)

NE

Drymobius rhombifer

(Gunther, 1860)

NE

Lampropeltis triangu-

lum (Lacepede, 1789)

NE

Leptophis ahaetulla

(Linneaus, 1758)

NE

Leptophis depressiros-

tris (Cope, 1861)

NE

Mastigodryas boddae-

rti (Sentzen, 1796)

NE

Mastigodryas danieli

(Amaral, 1935)

NE

Mastigodryas pleei

(Dumeril, Bibron &

Dumeril, 1854)

NE

Mastigodryas pulchri-

ceps (Cope, 1868)

NE

Red book of Red Book of

Colombia Valle del Cauca

Categorization for

the departament

Municipality

Protected area

NT

Ar, Bo, Cal,

Cai, Cu, Da,

Do, Tr, Ve, Yo

RFPN del Bosque de

Yotoco, PNN Faral-

lones de Cali, RFPN

de Cali

NT

Bu, Bug, Cal,

Cu, Da, Yo

PNN Farallones de

Cali, RFPN de Cali,

RFPN del Bosque

de Yotoco, RFPN

Anchicaya

EN, B2ab(iii)

Bu, Dar

PNN Farallones de

Cali, RFPN de los

rios Escalerete y San

Cipriano

EN, Blab(iii)

Ar, Bu, Cai,

Dar, Do, Ve

RFPN de los rios

Escalerete y San

Cipriano

DD

Bu

RFPN Anchicaya

NT

An, Bu, Cal,

Can, Ce, Yo

RFPN Anchicaya,

RFPN del Bosque de

Yotoco

EN, Blab(iii)

Bu

RFPN Anchicaya;

RFPN de los rios

Escalerete y San

Cipriano

LC

Bu, Cal, Cai,

Da, Ja, Pa,

Tu, Vij, Yu

RFPN de los rios

Escalerete y San

Cipriano, RFPN

Anchicaya, RFPN de

Cali, PNN Farallones

de Cali, RFPR Bitaco,

RFPN Cerro Dapa -

Carisucio

NT

Bu, Bui, Cal,

Pa, Yo

RFPN del Bosque de

Yotoco

NT

Bu

RFPN de los rios

Escalerete y San

Cipriano, RFPN

Anchicaya

LC

Bu, Cal

RFPN Anchicaya,

PNN Farallones de

Cali, RFPN de los

rios Escalerete y San

Cipriano, DMI La

Plata, PNN Uramba

Bahia Malaga, PNR

La sierpe, RFPN de

Cali

NT

Cal, Cu, Pa

RFPN de Cali, PNN

Farallones de Cali,

PNN Las hermosas

NT

Bug, Cal, Cu,

Da, Gu, Ja,

Pa, Re, Ro,

Se, Tu, Vi, Vij,

Yo, Yu

PNN Farallones de

Cali

NT

Bu, Cal

RFPN Anchicaya,

RFPN de Cali, RFPN

de los rios Escalerete

y San Cipriano

Amphib. Reptile Conserv.

December 2014 | Volume 8 | Number 2 | e87

Conservation status of the herpetofauna in Colombia

TAXON

lUCN

Red book of

Colombia

Red Book of

Valle del Cauca

Categorization for

the departament

Municipality

Protected area

Oxybelis aeneus (Wa-

gler, 1824)

NE

LC

Al, An, Ans,

Ar, Bo, Bu,

Bug, Bui, Ca,

Cal, Can, Car,

Da, Dar, Do,

Cai, Ce, Ag,

FI, Gi, Gu, Ja,

Cu, Vi, Un,

Ob, Pa, Pr,

Re, Ri, Ro,

Sp, Se, To, Tr,

Tu, Ul, Ve, Vij,

Yo, Yu, Za

RFPN Anchicaya,

PNN Farallones

de Cali, PNN Las

hermosas, RFPN de

los rios Escalerete y

San Cipriano, DMI La

Plata, PNN Uramba

Bahia Malaga, PNR

La sierpe, RN Laguna

de sonso, RFPN de

Cali, RFPN Dagua,

DMI Enclave subxe-

rofitico de Atuncela,

PNR Paramo el Du-

ende, PNN Tatama,

RFPN Amaime,

RFPR Bitaco, RFPN

Cerro Dapa - Ca-

risucio

Oxybelis brevirostris

(Cope, 1861)

NE

LC

Bu, Da

DMI La Plata, RFPN

Anchicaya, PNN

Farallones de Cali

PUocercus euryzonus

Cope, 1862

LC

VU, B2ab(iii)

Bu, Da, Dar

RFPN Anchicaya,

RFPN de los rios

Escalerete y San

Cipriano, PNN Faral-

lones de Cali

Pseustes poecilonotus

(Gunther, 1858)

LC

VU, B2ab(iii)

Bu

RFPN Anchicaya,

PNN Farallones de

Cali

Pseustes Shropshire!

(Barbour & Amaral,

1924)

NE

VU, B2ab(iii)

Bu

RFPN Anchicaya,

PNR La sierpe

Rhinobothryum bovallii

(Anderson, 1916)

NE

EN, Blab(iii)

Bu

RFPN Anchicaya,

PNR La sierpe

Spilotes pullatus (Lin-

naeus, 1758)

NE

LC

Al, An, Ans,

Ar, Bo, Bu,

Bug, Bui, Ca,

Cal, Can, Car,

Da, Dar, Do,

Cai, Ce, Ag,

FI, Gi, Gu, Ja,

Cu, Vi, Un,

Ob, Pa, Pr,

Re, Ri, Ro,

Sp, Se, To, Tr,

Tu, Ul, Ve, Vij,

Yo, Yu, Za

RFPN Anchicaya,

RFPN de los rios

Escalerete y San

Cipriano, RFPN

de Cali, RFPN del

Bosque de Yotoco,

RFPN Dagua, RFPN

de Amaime, RFPN de

sonso-guabas, RFPR

Bitaco, RFPN Cerro

Dapa - Carisucio, RN

Laguna de sonso,

PNR La sierpe,

DMI La Plata, DMI

Enclave subxerofitico

de Atuncela, PNR

Paramo el Duende,

PNN Tatama, PNN

Farallones de Cali,

PNN las hermosas,

PNN Uramba-Bahia

Malaga

Stenorrina degen-

hardtii {Bedho\6, 1845)

NE

NT

Bu, Dar

RFPN Anchicaya,

RFPN de los rios

Escalerete y San

Cipriano, DMI La

Plata

Amphib. Reptile Conserv.

S-17

December 2014 | Volume 8 | Number 2 | e87

Valencia-Zuleta et al.

TAXON

lUCN

Red book of Red Book of

Colombia Valle del Cauca

Categorization for

the departament

Tantilla melanoceph-

ala (Linnaeus, 1758)

{sensu latu)

FAMILIA DIPSADIDAE

Atractus boulengerii

(Peracca, 1896)

NE

Atractus clarki (Dunn &

Bailey, 1939)

NE

Atractus lehmanni

(Boettger, 1898)

NE

Atractus melas (Bou-

lenger, 1908)

NE

Atractus multicinctus

(Jan, 1865)

NE

Atractus obesus

(Marx, 1960)

NE

Clelia clelia (Daudin,

1803)

NE

Clelia equatoriana

(Amaral, 1924)

NE

Diaphorolepis wagneri

(Jan, 1863)

NE

Dipsas sanctijoannis

(Boulenger, 1911)

DD

Dipsas temporalis

(Werner, 1909)

NE

Erythrolamprus bizo-

nus (Jan, 1863)

LC

Erythrolamprus epi-

nephelus {Cope, 1862)

NE

Municipality

Al, An, Ans,

Ar, Bo, Bu,

Bug, Bui, Ca,

Cal, Can, Car,

Da, Dar, Do,

Cai, Ce, Ag,

FI, Gi, Gu, Ja,

Cu, Vi, Un,

Ob, Pa, Pr,

Re, Ri, Ro,

Sp, Se, To, Tr,

Tu, Ul, Ve, Vij,

Yo, Yu, Za

DD (t)

Bu

DD

Re

DD

Cal, Cu, Da,

Se

DD

Bu, Cal, Da

VU, B2ab(iii)

Bu, Cal, Cu,

Da

DD

Cal, Ce, FI

LC

Bu, Cal, Ce,

Cu, Da, Yo,

Yu

EN, B2ab(iii)

Ans, Ar, Bo,

Cai, Do, Un,

Pr, To, Ve

VU, B2ab(iii)

Bu, Cal, Cai,

Cu, Da

LC

Bu, Cal, Cu,

Da, Yo

DD

Bu

LC

Bu, Bug, Cal,

Car, Cu, Da,

Dar, FI, Ja,

Pa, Un, Yu

NT

Bo, Bu, Bug,

Cal, Cai, Da,

Tr

Protected area

RFPN del Bosque

de Yotoco, RFPN de

Cali, RFPNAnchi-

caya, RFPN de los

rios Escalerete y

San Cipriano, RFPR

Bitaco, RFPN Cerro

Dapa - Carisucio,

RFPN Dagua, RFPN

de Amaime, DMI

Enclave subxerofitico

deAtuncela, RN

Laguna de sonso,

PNN Tatama, PNN

Farallones de Cali

RFPN Anchicaya

RFPN Anchicaya,

RFPR Bitaco

PNN Farallones de

Cali, RFPN de Cali

RFPN de Cali, RFPN

Anchicaya

PNN Farallones de

Cali, PNN las hermo-

sas, RFPR Frayle-

Desbaratado

RFPN Anchicaya,

RFPN de Cali,

RFPN del Bosque

de Yotoco, PNN las

hermosas

PNN Las hermosas

PNN Farallones de

Cali

RFPN de los rios

Escalerete y San

Cipriano, PNN Faral-

lones de Cali, RFPN

deCali, RFPN del

Bosque de Yotoco

RFPN Anchicaya

RFPN Anchicaya,

PNN Farallones de

Cali, PNN las hermo-

sas, RFPN de Cali,

RFPR Bitaco, RFPN

Cerro Dapa - Carisu-

cio, RFPN Dagua

RFPN de Cali, RFPN

Anchicaya, PNN

Farallones de Cali

Amphib. Reptile Conserv.

December 2014 | Volume 8 | Number 2 | e87

Conservation status of the herpetofauna in Colombia

TAXON

lUCN

Red book of

Colombia

Red Book of

Valle del Cauca

Erythrolamprus mimus

(Cope, 1868)

Geophis betaniensis

(Restrepo & Wright,

1987)

Geophis nigroalbus

(Boulenger, 1908)

Categorization for

the departament

VU, B2ab(iii)

Imantodes cenchoa

(Linnaeus, 1758)

Imantodes inornatus

(Boulenger, 1896)

Imantodes chocoen-

sis (Torres-Carvajal,

Yanez-Munoz, Quirola,

Smith & Almendariz,

2012)

Leptodeira annulata

(Linnaeus, 1758)

Leptodeira septentrio-

na//s (Kennicott, 1859)

Ninia atrata (Hallowell,

1845)

Nothopsis rugosus

(Cope, 1871)

Municipality

Bu, Da

Protected area

RFPN Anchicaya,

RFPN de los rios

Escalerete y San

Cipriano, PNN Faral-

lones de Cali

Bo, Da, Ri

Bo,Cu, Tr

Al, An, Ans,

Ar, Bo, Bu,

Bug, Bui, Ca,

Cal, Can, Car,

Da, Dar, Do,

Cai, Ce, Ag,

FI, Gi, Gu, Ja,

Cu, Vi, Un,

Ob, Pa, Pr,

Re, Ri, Ro,

Sp, Se, To, Tr,

Tu, Ul, Ve, Vij,

Yo, Yu, Za

RFPN de los rios Es-

calerete y San Cipria-

no, RFPN Anchicaya,

PNN Farallones de

Cali, DMI La Plata,

PNN Uramba Bahia

Malaga, PNR La

sierpe, RN Laguna de

sonso, RFPN de Cali,

RFPN Dagua, DMI

Enclave subxerofitico

deAtuncela, PNR

Paramo el Duende,

RFPN de Amaime,

PNN Tatama, RFPN

del Bosque de

Yotoco, RFPR Bitaco,

RFPN Cerro Dapa -

Carisucio

Al, An, Ans,

Ar, Bo, Bu,

Bug, Bui, Ca,

Cal, Can, Car,

Da, Dar, Do,

Cai, Ce, Ag,

FI, Gi, Gu, Ja,

Cu, Vi, Un,

Ob, Pa, Pr,

Re, Ri, Ro,

Sp, Se, To, Tr,

Tu, Ul, Ve, Vij,

Yo, Yu, Za

RFPN de los rios

Escalerete y San

Cipriano

RFPN Anchicaya,

PNN Farallones de

Cali, RFPN de los

rios Escalerete y San

Cipriano, DMI La

Plata, PNN Uramba

Bahia Malaga, PNR

La sierpe, RN Laguna

de sonso, RFPN de

Cali, RFPN Dagua,

DMI Enclave subxe-

rofitico deAtuncela,

PNN Tatama, RFPN

del Bosque de

Yotoco, RFPR Bitaco;

RFPN Cerro Dapa -

Carisucio, RFPN de

Amaime

EN, Blab(iii)

RFPN Anchicaya

RFPN Anchicaya;

RFPN de los rios

Escalerete y San

Cipriano

Amphib. Reptile Conserv.

December 2014 | Volume 8 | Number 2 | e87

Valencia-Zuleta et al.

TAXON

lUCN

Red book of

Colombia

Red Book of

Valle del Cauca

Oxyrhopus petolarius

(Linnaeus, 1758)

Saphenophis boursieri

(Jan, 1867)

Saphenophis sneiderni

Myers, 1973

Saphenophis tristiatus

(Rendahl & Vester-

gren, 1941)

Sibon nebuiata (Lin-

naeus, 1758)

Siphiophis compres-

sus (Daudin, 1803)

Synophis piectoverte-

braiis (Sheil & Grant,

2001)

Tretanorhinus taenia-

tus (Boulenger, 1903)

Urotheca decipiens

(Gunter 1893)

Urotheca fuiviceps

(Cope, 1886)

Urotheca iateristriga

(Berthold, 1859)

Xenodon rabdocepha-

/us(Wied, 1824)

Protected area

RFPN Anchicaya,

PNN Farallones de

Cali, RFPN de los

rios Escalerete y San

Cipriano, DMI La

Plata, PNN Uramba

Bahia Malaga, PNR

La sierpe, RN Laguna

de sonso, RFPN de

Cali, RFPN Dagua,

DMI Enclave subxe-

rofitico de Atuncela,

PNN Tatama, RFPN

del Bosque de

Yotoco, RFPR Bitaco,

RFN Cerro Dapa -

Carisucio

PNN Farallones

de Cali, PNN Las

hermosas

RFPN Anchicaya,

RFPN de los rios

Escalerete y San

Cipriano, RFPN del

Bosque de Yotoco,

RFPN de Cali, RFPR

Bitaco, RFPN Cerro

Dapa - Carisucio,

PNR La sierpe, RN

Laguna de sonso,

RFPN Dagua, DMI La

Plata, DMI Enclave

subxerofitico de

Atuncela, PNN Tata-

ma, PNN Farallones

de Cali, PNN Uramba

- BahiaMalaga

RFPN Anchicaya,

RFPN de los rios

Escalerete y San

Cipriano

RFPN Anchicaya

PNN Farallones de

Cali, RFPR Bitaco,

RFPN Cerro Dapa -

Carisucio, RFPN del

Bosque de Yotoco

RFPN Anchicaya

RFPN de Cali, RFPN

Anchicaya, RFPR

Bitaco

DMI La Plata, RFPN

Anchicaya, RFPN de

los rios Escalerete y

San Cipriano

Amphib. Reptile Conserv.

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December 2014 | Volume 8 | Number 2 | e87

Conservation status of the herpetofauna in Colombia

TAXON

lUCN

FAMILIA ELAPIDAE

Micrurus ancoralis

(Jan, 1872)

NE

Micrurus clarki

(Schmidt, 1936)

NE

Micrurus dumerilii

(Jan, 1858)

NE

Micrurus mipartitus

(Dumeril, Bibron &

Dumeril, 1854)

NE

Micrurus multifasciatus

(Jan, 1858)

NE

Micrurus multiscutatus

(Rendahl & Vester-

gren, 1940)

DD

Red book of Red Book of

Colombia Valle del Cauca

Categorization for

the departament

Municipality

FAMILIA LEPTOTYPHLOPIDAE

Trilepida joshuai

(Dunn, 1944)

LC

Trilepida macrolepis

(Peters, 1857)

NE

FAMILIA TROPIDOPHIIDAE

Trachyboa boulengeri

(Peracca, 1910)

NE

FAMILIA VIPERIDAE

Bothriechis schlegelii

(Berthold, 1846)

NE

Bothrocophias myersi

(Gutberlet & Campbell,

2001)

LC

NT

Bu

DD

Da

NT

Bu, Da

LC

Bo, Bug, Bu,

Cal, Cai, Cu,

Da, Dar, FI,

Gu, Ja, Pa,

Pr, Ri, Tu, Yo

DD

Dar

DD

Bu, Dar

Bothrops asper (Gar-

man, 1884)

Protected area

DMI La Plata, RFPN

Anchicaya, RFPN de

los rios Escalerete y

San Cipriano

RFPN Anchicaya

RFPN Anchicaya,

RFPN de los rios

Escalerete y San

Cipriano, RFPN de

Cali; PNR La sierpe,

PNN Farallones de

Cali

RFPN del Bosque

de Yotoco, RFPN

Anchicaya, RFPN de

los rios Escalerete y

San Cipriano, RFPN

de Cali, RFPR Bitaco,

PNR La sierpe, RN

Laguna de sonso,

PNN Farallones

de Cali, PNN Las

hermosas

DD

Bo, Cal, Cai,

Da

RFPN de Cali, PNN

Farallones de Cali

DD

Cal, Dar

RFPN de Cali

NT

Bu

RFPN de los rios

Escalerete y San

Cipriano, DMI La

Plata

NT

Bu, Cal, Cai,

Cu, Da, Dar,

FI, Pa, Se,

Tr, Yo

RFPN Anchicaya,

RFPN del Bosque de

Yotoco, RFPR Bitaco,

PNN Las hermosas

VU,A2ac

Bu, Dar

RFPN Anchicaya,

PNR La sierpe

Bu, Cal, Da,

Dar, Pa, Yo

RFPN Anchicaya,

RFPN de los rios

Escalerete y San

Cipriano, RFPN del

Bosque de Yotoco,

DMI Enclave subxe-

rofitico de Atuncela,

DMI La Plata, RFPN

Dagua, RFPN de

Cali, PNR La sierpe,

PNN Farallones de

Cali, PNN Uramba-

Bahia Malaga

Amphib. Reptile Conserv.

December 2014 | Volume 8 | Number 2 | e87

Valencia-Zuleta et al.

TAXON

lUCN

Red book of

Colombia

Red Book of

Valle del Cauca

Categorization for

the departament

Municipality

Protected area

Bothrops punctatus

(Garcia, 1896)

NE

VU, A2ac

Bu

RFPN de los rios

Escalerete y San

Cipriano, PNR La

sierpe

Bothrops rhombeatus

(Garcia, 1896)

NE

VU, B2ab(iii)

Cal, Pa

RFPN de Cali

Lachesis acrochorda

(Garcia 1896)

NE

VU,A2ac

Bu

RFPN Anchicaya

Porthidium nasutum

(Bocourt, 1868)

LC

VU,A2ac

Bu

RFPN Anchicaya,

RFPN de los rios

Escalerete y San

Cipriano

ORDEN TESTUDINATA

SUBORDEN CRYTODIRA

FAMILIA CHELYDRIDAE

Chelydra acutirostris

(Peters, 1862)

NE

DD

NT

An, Bu, Bug,

Cal, Car, Pa,

Yo, Za

RFPN de los rios

Escalerete y San

Cipriano, RN Laguna

de sonso

FAMILIA EMYDIDAE

Trachemys scripta

(Thunberg in Schoepff,

1792)

NE

LC

Bug, Cal

RN Laguna de sonso

FAMILIA GEOEMYDIDAE

Rhinoclemmys annu-

lata (Gray, 1860)

NT

DD

VU,A2acd

Bu

RFPN de los rios

Escalerete y San

Cipriano, PNR La

sierpe

Rhinoclemmys mela-

nosterna (Gray, 1861)

NE

NT

VU,A2acd

Bu

RFPN Anchicaya,

PNR La sierpe

Rhinoclemmys nasuta

(Boulenger, 1902)

NT

DD

NT

Bu

RFPN Anchicaya,

RFPN de los rios

Escalerete y San

Cipriano, PNR La

sierpe

FAMILIA KINOSTERNIDAE

Cryptochelys dunni

(Schmidt, 1947)

VU

VU, B2ab(iii)

Bu

PNR La sierpe

Cryptochelys leucosto-

mum (Dumeril, Bibron

& Dumeril, 1851)

NE

NT

Bu

RFPN Anchicaya,

RFPN de los rios

Escalerete y San

Cipriano, PNR La

sierpe

Considering that recently some taxonomic problems have been resolved, we provide to some species or groups of species, the

taxonomic category of Sensu latu or cf., understanding that these species could eventually change and give new information for the

department.

Amphib. Reptile Conserv.

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December 2014 | Volume 8 | Number 2 | e87

Conservation status of the herpetofauna in Colombia

Table 2. Criteria

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Amphib. Reptile Conserv.

S-23

December 2014 | Volume 8 | Number 2 | e87

Valencia-Zuleta et al.

1 '

>i- ■ -* N ■ -

. ' .a

V. . ■

V 'A V*

« . "if*-

■^v. it.

v/'.

Pristimantis achatinus

Pristimantis palmeri

Anolis maculiventris

Anolis chocorum

Bothrops asper

Clelia clelia

Corallus annulata

Dendrophidion bivittatus

Amphib. Reptile Conserv.

S-24

December 2014 I Volume 8 I Number 2 I e87

Comparative dorsal view of the head of Trimeresurus gunaleni spec. nov. (left) and T. sumatranus (right). Left from above: male,

female (holotype), male, all alive, from Sumatra Utara Province, Sumatra. Right: adult female alive from Bengkulu Province, Su-

matra, adult male alive from Bengkulu Province, Sumatra, preserved female from Borneo. Photos: N. Maury.

( 1 )

September 201 4 | Volume 8 | Number 2 | e80

Amphib. Reptile Conserv. | amphibian-reptile-conservation.org

Amphibian & Reptiie Conservation

8(2) [General Section]: 1-29.

of the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License,

which permits unrestricted use for non-commercial and education purposes only provided

the original author and source are credited. The official publication credit source: Amphib-

ian & Reptile Conservation at: amphibian-reptile-conservation.org

On Trimeresurus sumatranus (Raffles, 1822 ), with the desig-

nation of a neotype and the description of a new species of

pitviper from Sumatra (Squamata: Viperidae: Crotalinae)

^Gernot Vogel, ^Patrick David, and ^Irvan Sidik

^Society for Southeast Asian Herpetology, Im Sand 3, D-69115 Heidelberg, GERMANY ^Reptiles & Amphibiens, UMR 7205 OSEB, Departement

Systematique et Evolution, CP 30, Museum National d’Histoire Naturelle, 57 rue Cuvier, 75231 Paris Cedex 05, PRANCE ^Laboratory of

Herpetology, Museum Zoologicum Bogoriense, Zoology Division, Research Center for Biology, Indonesian Institute of Sciences, Widyasatwa Loka

Jl. Raya Jakarta-Bogor Km 46, Cibinong 16911, INDONESIA

Abstract. — Variation in morphological characters were investigated among 126 specimens

from at least 67 populations covering the whole range of the large pitviper currently known as

Trimeresurus sumatranus (Raffles, 1822). The results showed that two distinct taxa are involved.

Herein Trimeresurus sumatranus is redefined. In order to fix the status of this species, a neotype is

selected and described. Its type locality is restricted to the vicinity of Bengkulu, Bengkulu Province,

Sumatra. The second taxon represents a distinct, previously unnamed species, which is described.

The new species differs from Trimeresurus sumatranus by a lower number of ventrals in males

(162-179 against 178-185) and females (164-171 vs. 175-191); a distinctly longer tail in males

(value of the ratio tail length/total length: 0.201-0.210 vs. 0.150-0.168), the color of the tail (see the

description), the color of the eyes: green in the new species, vs. dark grey in T. sumatranus, the color

of the ventrals, which are green with a pale posterior suture in the new species and pale with dark

posterior suture in T. sumatranus. The new species lives in higher elevations than T. sumatranus

and seems to be endemic to the higher mountain ranges of western Sumatra.

Key words. Sumatra, West Malaysia, Borneo, Trimeresurus gunaleni spec, nov., Trimeresurus malcolmi, Trimeresu-

rus sumatranus

Citation: Vogel G, David P, Sidik I. 2014. On Trimeresurus sumatranus (Raffles, 1822), with the designation of a neotype and the description of a new

species of pitviper from Sumatra (Squamata: Viperidae: Crotalinae). Amphibian & Reptiie Conservation 8(2) [General Issue]: 1-29 (e80).

Introduction

The genus Trimeresurus Lacepede, 1804 is currently

composed of 46 species, of which 12 were described

since the year 2000 (David et al. 2011). This genus and

most of its species have complex systematic and nomen-

clatural histories. For example, the generic nomen P arias

Gray, 1849 was regarded as a valid subgenus only re-

cently (David et al. 2011). This extensive series of confu-

sions arose from the fact that many species of this genus

are arboreal; green snakes that are superficially similar

in color and scalation. This factor is exemplified by the

case of Trimeresurus sumatranus (Raffles, 1 822) and its

relatives. Trimeresurus sumatranus (Raffles, 1822) was

one of the first species of this genus of Asian pitvipers

to have been described (as Coluber sumatranus), a point

that is quite surprising as it is a rather uncommon spe-

cies with a limited distribution in the Sunda Islands. Sir

Thomas Stamford Raffles (1781-1826), best known as

the founder of Singapore, made a long stay in Bencoolen,

now Bengkulu, a city of south-western Sumatra, between

March 1818 and August 1824, although this stay was in-

Correspondence. Email: ^ Gernot.Vogel@t-online.de

Amphib. Reptile Conserv. | amphibian-reptile-conservation.org (2)

terrupted several times for trips to Nias and, especially

Singapore. He held the position of Governor-General of

Bencoolen from 1818 to 1822. An account on the activi-

ties of Sir T. Raffles in Sumatra can be found in Raffles

(1835). Sir Raffles was definitely not a scholarly natural-

ist but had a strong interest in local plants and animals

which he described himself (Raffles 1821, 1822).

Trimeresurus sumatranus has been extensively con-

fused in the literature with Trimeresurus hageni (Van

Lidth de Jeude, 1886), a related but distinct species that

inhabits lower elevations in Sumatra, Borneo, and the

Malay Peninsula. Both species share similar scalation,

and juvenile specimens of both species are uniformly

green in color. However, although the adults are very

different in coloration, the similarities in scalation led

Boulenger, the leading herpetologist of his time, to syn-

onymize these two species (Boulenger 1896). A reason

for this misinterpretation was the fact that at that time,

the British Museum of Natural History, the place where

Boulenger was working, had only a single specimen of

the species T. sumatranus which unfortunately was a ju-

venile and thus shared the green color with T hageni.

September 201 4 | Volume 8 | Number 2 | e80

Vogel et al.

Boulenger’s appraisal was followed by subsequent work-

ers, until the works of van Lidth de Jeude (1922) and

especially Brongersma (1933). Tliis latter author was the

first to show that T. sumatranus and T. hageni were valid,

distinct species, and he provided new characters separat-

ing both species from one another.

Furthermore, it is worth noting that Schlegel (1826)

described the taxon Cophias wagleri (nee Cophias wa-

gleri Boie, 1827) as a replacement name for Coluber

sumatranus (Raffles, 1822). We refer to Savage et al.

(2012) for a discussion on the early confusion between

these species. Nevertheless, other authors such as Schle-

gel (1837), Gray (1842: 48; 1849: 10), Cantor (1847:

1042, PI. 40: Fig. 9), Gunther (1858: 266), Peters (1862:

671) and later as Ouwens (1916: PI. 15: Fig. 22 and 22a)

also confused in part or totally Coluber sumatranus with

Cophias wagleri Boie, 1827, a totally different species

now known as Tropidolaemus wagleri.

Loveridge (1938: 45) described Trimeresurus su-

matranus malcolmi (Type locahty: “Sungii River, near

Bunduntuan, Mount Kinabalu, British Nord Borneo,”

a river in the vicinity of Bundu Tuhan, on the southern

slopes of Mt. Kinabalu, state of Sabah, Borneo, Federa-

tion of Malaysia). This subspecies was regarded as valid

by all subsequent authors for some populations of north-

ern Borneo whereas other populations of this island were

referred to the nominative subspecies (David and Ineich

1999; McDiamiid et al. 1999; Malkmus et al. 2002).

Subsequently, Trimeresurus sumatranus malcolmi was

raised to full species status by Stuebing and Inger (1998).

As a consequence, Trimeresurus sumatranus was subse-

quently considered monotypic.

More recently, two revisions of the systematics of

these two species were published by Sanders et al. (2002,

2004). Results of the first publication, which was based

on scalation characters, pattern and coloration, can be

summarized as follows: (1) T sumatranus and T hageni

are clearly separate as shown by canonical multivari-

ate analysis; (2) T sumatranus inhabits South Thailand,

Borneo, and central western Sumatra, whereas T. hageni

is living in North and South Sumatra, Thailand, Malay-

sia, Singapore, Nias, and Siberut; (3) the authors showed

clear differences between populations of T. sumatranus

inhabiting the central part of western Sumatra and that

one living on Borneo; and (4) morphological differences,

especially in males, were pointed out between popula-

tions of the islands of Nias and Siberut on the one hand,

and all other populations on the other hand. In these is-

lands, specimens referable to Trimeresurus hageni show

some characters of the pattern typical to T. sumatranus,

such as black dorsal crossbars and the presence of dark

edges on head scales. This partial similarity has led to

erroneous records of T. sumatranus from these islands.

In contrast, Sanders et al. (2004) included molecu-

lar analyses and considered all species of the subgenus

Parias Gray, 1849 as defined by Malhotra and Thorpe

(2004) (as a genus). For the species treated here, the

Amphib. Reptile Conserv. | amphibian-reptile-conservation.org (3)

results of Sanders et al. (2004) can be summarized

as follows: (1) T hageni has an expanded distribution

compared with Sanders et al. (2002), i.e. populations

of southern Thailand, West Malaysia, and the islands of

Bangka, Sibemt, Nias, and all islands of the Mentawai

Archipelago are refen-ed to this species; (2) little mor-

phological variation was found between populations of

T hageni, in contrast to results presented by Sanders et

al. (2002); (3) without justification, populations of South

Thailand and West Malaysia were no longer referred to T.

sumatranus but to T hageni', as a consequence, the range

of T. sumatranus was restricted to a naiTow area covering

mid elevations between 650 and 800 m — the central and

southern parts of the Barisan Range, western Sumatra,

and a wide range throughout Borneo mainly below 300

m; (4) populations of Sumatra and Borneo are separated

by a genetic distance of 3.3%; (5) differences in ecol-

ogy were pointed out between the Bornean and Suma-

tran populations; and (6) ecological adaptation has led

to a convergence in the pattern between T hageni and

T. sumatranus in Sumatra. Sanders et al. (2004) consid-

ered that this convergence made useless some characters

which used to be considered diagnostic; for example the

separation of the fourth and fifth supralabial with the

suboculars (see Brongersma 1933). Sanders et al. (2004)

provided quite a different definition of T sumatranus

compared with Brongersma (1933). However, it must be

emphasized that only two populations of T. hageni and

two populations of T. sumatranus, both from Bengkulu

Province for this latter species, were compared in mo-

lecular analyses.

In the frame of a revision of the subgenus Parias in

the Sunda Islands, namely of populations referred in the

literature to T. sumatranus, T hageni, and T. malcolmi,

we examined specimens refeiTed to as Trimeresurus su-

matranus originating from throughout the range of this

taxon, including material not investigated previously.

Our data suggest that the combination of several charac-

ters, both of pattern and scalation, allow a clear distinc-

tion between T sumatranus and T hageni. Definitions

of these two species will be presented in a subsequent

paper. In the first step of this revision, presented here,

we put emphasis on the morphology of the various popu-

lations refen'ed to as T. sumatranus auctorum. Our data

also show that two distinct forms of T. sumatranus auc-

torum can be defined in Sumatra, both deserving to be

recognized as full species.

Material and Methods

The present paper is based on 126 preserved specimens

examined by us from 67 localities covering the whole

range of T. sumatranus auctorum and T. hageni, and sev-

eral live specimens of both species. Preserved examined

specimens of the two forms of T. sumatranus are listed

under their respective account; specimens of T. hageni

are listed in Appendix I.

September 2014 | Volume 8 | Number 2 | e80

A new species of Trimeresurus from Sumatra

Selection of morphological characters

We retained standard morphological characters used in

the genus Trimeresurus by Brongersma (1933), Pope and

Pope (1933), and Regenass and Kramer (1981), along

with other morphometric characters adapted from Vogel

et al. (2004). We made a pre-selection of characters with

a limited number of specimens. Characters not suitable,

due to variability or uniformity were deleted and a set of

30 characters was retained (Table 1).

Measurements, except body and tail lengths, were

taken with a slide-calliper to the nearest 0.1 millimeter

(nun); all measures on body were taken to the nearest

mm. In order to minimize inter-observer error, all mea-

surements considered here were made by Gemot Vogel

(GV). Ventral scales were counted according to Dowl-

ing (1951). The first subcaudal was defined as the first

scale posterior to the vent that touched the opposite scale.

The terminal scute is excluded from the number of sub-

caudals. The numbers of dorsal scale rows are given at

one head length behind the head, at midbody (i.e., at the

level of the ventral plate corresponding to half of the total

number of ventrals), and at one head length anterior to

the vent respectively. Values for symmetric head char-

acters are given in left/right order. The real coloration of

body and eyes were observed only on living animals or

freshly preserved specimens.

Morphometric, meristic, and coloration characters re-

tained for this study are listed in Table 1. Altogether, 30

variables were considered, either standing on their own

or derived from the raw characters listed above. Not all

variables listed in this table proved to be useful to sepa-

rate at least one taxon of the Trimeresurus sumatranus

group from the others, but all were investigated and used

in combinations of characters and/or were used in uni-

variate analyses.

Table 1. List of morphological characters and variables used in this study and their abbreviations.

Number

Abbreviation

Character

Morphometry

1

SVL

Snout-vent length

2

TaL

Tail length

3

TL

Total length

4

TaL/TL

Ratio tail length/Total length

Scalation

5

Dorsal scale rows

6

Do

Dorsal scale rows at midbody

7

Dorsal scale rows at midbody

8

Ven

Ventral plates

9

Sc

Subcaudal plates

10

Cep

Cephalic scales (scales on a line between the middle of supraoculars)

11

InN

Intemasal scale(s)

12

InN sep

Intemasal scales touching each other

13

Keeling of the occipital scales

14

Supralabial scales

15

Number of scales between third supralabial and subocular

16

Number of scales between fourth supralabial and subocular

17

Number of scales between fifth supralabial and subocular

18

CtotSL

Total number of supralabials touching subocular

19

IL

Infralabials

Pattern

20

Presence of black margins on dorsal scales of the head

21

Upper labials being lighter than other parts of the head

22

Ventrals with dark margins

23

Subcaudals with dark margins

24

Presence of bands on the body

25

Presence of dorsolateral light spots on the body

26

Coloration and presence of a ventrolateral stripe

27

Coloration and presence of a temporal streak

28

Color of eyes

29

Posterior part of the tail reddish

30

Pattern of the tail

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Vogel et al.

The analyses of external morphological data were

based on comparisons of statistical values (mean value

and standard deviation). A test of Mann- Whitney {U

test; see Siegel 1956) was applied as necessary. Calcula-

tions were run online on the website: http://elegans.som.

vcu.edu/~leon/stats/utest.html (last accessed on 14 July

2014). Abbreviations are: n\ number of specimens; x\

mean value; standard deviation; P: probability of oc-

currence of a value as extreme as or more extreme than

the observed value; U: the statistic in the Mann- Whitney

test.

The color of the eyes is shown here to be a taxonomic

character. However, it is problematic as it cannot be ob-

served in preserved specimens. According to our obser-

vations, the eye color in adult animals is stable for each

species and sex (Vogel et al. 2004). In the species treated

here, there was no sexual dimorphism in eye coloration.

The color of the tail is diagnostic and we recognize two

patterns: “uniform reddish-brown with dark margins,” or

“mottled,” for specimens with a mixture of brown and

green colors on the tail.

Museum abbreviations

BMNH: The Natural History Museum, London, UK;

CAS: Cahfornia Academy of Sciences, San Francisco,

USA; FMNH: Field Museum of Natural History, Chi-

cago, USA; IRSNB: Institut Royal des Sciences Naturel-

les de Belgique, Bmssels, Belgium; MNHN: Museum

National d’Histoire Naturelle, Paris, France; NHMB:

Naturhistorisches Museum, Basel, Switzerland; NHMW:

Naturhistorisches Museum Wien, Austria; MZB: Mu-

seum Zoologicum Bogoriense, Bogor a Cibinong, Java,

Indonesia; OMNH: Osaka Museum of Natural History,

Osaka, Japan; PSGV: Gemot Vogel’s private collection,

Heidelberg, Germany; RMNH: Nationaal Natuurhisto-

risch Museum (Naturalis), Leiden, Netherlands; SMF:

Natur-Museum und Forschungs-Institut Senckenberg,

Frankfurt-am-Main, Germany; ZFMK: Zoologisches

Forschungsinstitut und Museum Alexander Koenig,

Bonn, Germany; ZMB: Zoologisches Museum fur

Naturkunde der Humboldt-Universitat zu Berlin, Ber-

lin, Germany; ZMH: Zoologisches Institut und Museum,

Universitat Hamburg, Hamburg, Germany; ZRC: Zoo-

logical Reference Collection, National University of Sin-

gapore, Singapore; ZSM: Zoologische Staatssannnlung,

Munchen, Gennany.

Other abbreviations

Measures and ratios: ED: vertical diameter of the eye.

HL: Head length, SVL: Snout- vent length, TaL: Tail

length, TL: Total length, TaL/TL: Ratio tail length/total

length.

Meristic characters: DSR: Formula of dorsal scale rows,

IL: Infralabials, SC: Subcaudals, SL: supralabials, VEN:

Ventrals.

Results

In our sample of 53 specimens referred to Trimeresu-

rus sumatranus, as currently defined, we noticed that

nine specimens from western Sumatra differed in sev-

eral morphological characters from other populations.

Trimeresurus sumatranus (Raffles, 1 822) was briefly de-

scribed (as Coluber sumatranus) without any designation

of a name-bearing type. Therefore, we first redefine this

species and note intraspecific variation of the characters

examined. We then designate a neotype for this species

in agreement with Art. 75.3.1 to 75.3.7 of the Interna-

tional Code of Zoological Nomenclature (I.C.Z.N. 1999;

merely designated below as the Code).

Trimeresurus sumatranus (Raffles, 1822)

Fig. 1-7

Coluber sumatranus Raffles, 1822: 334.

Type locality. By virtue of neotype designation: “SW

Sumatra” (original type locality: implicitly “Suma-

tra;” restricted to vicinity of Bengkulu city, Bengkulu

Province, Sumatra fide Wallach et al. [2014: 527]; see

also the discussion given below).

Neotype. ZFMK 76340, adult female; deposited by An-

dreas Gumprecht (holotype not traced according to

McDiarmid et al. 1999: 345, considered to be lost).

Trigonocephalus formosus Muller and Schlegel, 1842

(in 1842-1845): PI. 7 [dated 1842]; text [dated 1845]:

52 and 55.

Type locality. “Aan de westkust van het eiland Sumatra,

in de omstreken van het dorp Limomanis, eenige uren

beoosten Padang. . .,” i.e.: on the west coast of Suma-

tra Island, in the vicinity of Limomanis, a few hours

east of Padang, now near Limau Manis, Province of

Sumatera Barat, Sumatra, Indonesia.

Holotype. RMNH 1583, adult male; deposited by S.

Muller, 1835.

Status. Junior subjective synonym of Coluber sumatra-

nus Raffles, 1822. Synonymized by Lidth de Jeude

(1886: 51).

Material Examined {n = 44)

Indonesia

Sumatra. Bengkulu Province. MZB 1035, Gunung Ge-

dang; MZB 2180, “Muara Aman, North Bengkulu;”

MZB 3718, ZMB 66177-8; ZMB 76340, ZMB 70490,

“Bengkulu;” ZMH R06936, Lebong-Tandai (3°01’S-

101°5’E). Jambi Province. MZB 457, “Jambi.” Lampung

Province. MZB 2166, Rimba; MZB 2219, “Propinsi

Lampung.” Sumatera Barat Province. OMNH R2135-6,

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A new species of Trimeresurus from Sumatra

Kambot, Ulu Gadut, Mt. Gadut, ca. 800 m; MZB 2443,

MZB 2445, Anai River. No locality. ZFMK 76340 (neo-

type), South-western Sumatra.

Borneo (Kalimantan). Kalimantan Barat Province.

MZB 1052, Sungai Mentawit Balik; ZSM 283/1977-

1-2, Landak River; MZB 2138a-b, Tangung Lokang,

Kapuas Hulu Regency. Kalimantan Tengah Province.

MZB 2647, Mamwai. Kalimantan Timur Province. MZB

1340, Mapa Kelai River. Unidentified locality. MZB

2424, Sungai Auge.

Federation of Malaysia

Borneo (East Malaysia). State of Sabah. FMNH

239949-52, FMNH 239957-8, Tenom District; FMNH

239959, Sipitang District. State of Sarawak. BMNH

91.8.29.33, Mt. Dulit, Miri District, Miri Division;

BMNH 1978.1879, Gunung Mulu National Park, Miri

District, Miri Division; FMNH 138687-8, FMNH

138690, FMNH 148829, Kapit District, Kapit Division;

FMNH 158671, Bintulu, Bintulu Division.

West Malaysia. State of Johore. BMNH 1971.1532, Pan-

ti Forest Reserve, South Johore. State of Pahang. ZRC

2.2929, Kuala Tahan; ZMB 69982, “Pahang.” State of

Trengganu. BMNH 1974.5001-3, Gunong Lawit.

Thailand

Yala Province. BMNH 1936.9.12.3, “Betong, Pattani.”

Taxonomic and nomenclatural comments

There is no doubt about the distinct specific status of T.

sumatranus and T. hageni. Our material shows that, in

contrast to the conclusions of Sanders et al. (2004), Tri-

meresurus sumatranus is a rather wide-ranging species.

We examined several specimens from West Malaysia and

one from extreme southern Thailand, and they are mor-

phologically identical with specimens originating from

most populations of Sumatra and Borneo. There is no

reason for assigning them to any other species of the sub-

genus Parias, and definitely not to T. hageni.

Furthermore, T. sumatranus auctorum is here shown

to be composed of two species in Sumatra. Some popula-

tions of Sumatera Barat Province, in the northern part of

the range of the species, are here referred to a new species

that is described below. We examined the holotype of T.

formosus Muller and Schlegel (1842: Fig. 2). We confirm

that this specimen is definitely referred to Trimeresurus

sumatranus and not to the new species described below

that inhabits the same region. Trimeresurus sumatranus,

as here redefined, is monotypic.

Raffles (1822) deseribed this speeies on the basis of

a single specimen. As he was posted in “Bencoolen,”

and aeeordmg to S. Raffles (1835: 102-104), his widow,

T. Raffles was interested in local “wonders in natural

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history.” Furthermore, T. Raffles announced in a letter

dated on 14 March 1820 that he planned to ship home,

(England) the whole of his zoological collection “in a

few days.” This collection was shipped before 29 March.

So, by all evidence, the holotype of Coluber sumatranus

originated from Bengkulu or its vicinity. On the basis of

these historical considerations, Wallach et al. (2014) re-

stricted the type locality of Coluber sumatranus to this

city on the south-western coast of Sumatra. Considering

that the description of the new species was published in

1822, we may ascertain that the holotype was included in

this shipment and originated from the vicinity of Beng-

kulu. However, the fate of the specimen is unknown and,

by all evidence, it should be considered lost. As a conse-

quence, we here designate a neotype for Coluber suma-

tranus (Raffles, 1 822).

Designation and Description of the Neotype

of Coluber sumatranus Raffles, 1 822

The designation is made on the following basis and in

agreement with the following articles of the Code: (1) the

neotype is designated in order to fix the status of Colu-

ber sumatranus (Raffles, 1 822) aceording to its eurrent

definition in the literature, especially in regards to popu-

lations described below as a new species and of other

species of the subgenus Parias (Art. 75.3.1 of the Code);

(2) diagnostic characters of Coluber sumatranus for

which we designate this neotype, are given below (Art.

75.3.2); (3) the neotype is designated in details below

(Art. 75.3.3); (4) a holotype has never been mentioned

in the literature, for example by Boulenger (1896). It

could not be traced in the collections of the Natural His-

tory Museum (London), in contrast to Cox et al.’s (2012)

statement, or of the Zoological Reference Collection of

the National University of Singapore. For these reasons,

we consider the holotype to be lost (Art. 75.3.4); (5) we

select a specimen, the morphology of which i.e., scala-

tion, pattern, and coloration, that agrees with characters

provided in the original description (Art. 75.3.5); (6) as

shown above, the holotype most probably originated

from Bengkulu Province. We select a neotype from an

area of Sumatra that encompasses Bengkulu Province

(Art. 75.3.6). For these reasons, and in agreement with

Art. 75.3.7 of the Code, we here designate as the neotype

of Coluber sumatranus as the following specimen:

ZFMK 76340, an adult female, from “Southwestern

Sumatra” (Fig. 1)

Morphology and measurements

Body elongate, compressed; head elongate, relatively

naiTow seen from above, massive seen from the side, dis-

tinctly triangular, wide at its base, clearly distinct from

the neck, flattened but thick, 1.8 times as long as wide;

snout quite long, round when seen from above, strongly

obliquely truncated when seen from the side, with a mod-

September 201 4 | Volume 8 | Number 2 | e80

Vogel et al.

A

Fig. 1 A-C. Trimeresurus sumatranus, ZFMK 76340, neotype of Coluber sumatranus Raffles, 1822, southwest Sumatra, Bengkulu

Province, Sumatra. A. dorsal view of the body, B. ventral view of the body, C. lateral view oft the head. Photo: G. Vogel.

September 201 4 | Volume 8 | Number 2 | e80

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A new species of Trimeresurus from Sumatra

Fig.2A-C. Trimeresurus su-

matranus, RMNH 1583, Ho-

lotype of Trigonocephalus

formosus Muller and Schle-

gel, 1842, from Padang, Su-

matera Barat Province, Su-

matra A. dorsal view of the

body, B. ventral view of the

body, C. lateral view of the

head. Photo: G. Vogel

September 201 4 | Volume 8 | Number 2 | e80

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Vogel et al.

Fig. 3A. MZB.Ophi.5452 live holotype of Trimeresurus gunaleni spec. nov. from Mt. Sibayak, ca. 1,800 m a.s.l., west of Brastagi,

Sumatera Utara Province, Sumatra, adult female. Photo: G. Vogel.

Fig. 3B. MZB.Ophi.5452 live holotype of Trimeresurus gunaleni spec. nov. from Mt. Sibayak, ca. 1,800 m a.s.l., west of Brastagi,

Sumatera Utara Province, Sumatra, adult female. Photo: G. Vogel.

September 201 4 | Volume 8 | Number 2 | e80

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A new species of Trimeresurus from Sumatra

Fig. 4A. Live male of Trimeresurus gunaleni spec. nov. from Mt. Singkut, 1,600 m a.s.L, Sumatra Utara Province, Sumatra. Photo:

G. Vogel.

Fig. 4B. Live male of Trimeresurus gunaleni spec. nov. from Mt. Singkut, 1,600 m a.s.L, Sumatra Utara Province, Sumatra. Photo:

G. Vogel.

September 201 4 | Volume 8 | Number 2 | e80

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Vogel et al.

Fig. 5. Comparative dorsal view of the head of Trimeresurus gunaleni spec. nov. (left) and T. sumatranus (right). Left from above:

Male, female (holotype), male, all from Sumatra Utara Province, Sumatra alive, right adult female alive from Bengkulu Province,

Sumatra, adult male alive from Bengkulu Province, Sumatra, preserved female from Borneo. Photos: N. Maury.

September 201 4 | Volume 8 | Number 2 | e80

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A new species of Trimeresurus from Sumatra

Fig. 6. Comparative lateral view of the head of Trimeresurus gunaleni spec. nov. (left) and T. sumatranus (right). Left from above:

Male, female (holotype), male, all from Sumatra Utara Province, Sumatra alive, right adult female alive from Bengkulu Province,

Sumatra, adult male alive from Bengkulu Province, Sumatra, preserved female from Borneo. Photos: N. Maury.

September 201 4 | Volume 8 | Number 2 | e80

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Vogel et al.

Fig. 7. Comparative ventral view of the head of Trimeresurus gunaleni spec. nov. (left) and T. sumatranus (right). Left from above:

Male, female (holotype), male, all from Sumatra Utara Province, Sumatra alive, right adult female alive from Bengkulu Province,

Sumatra, adult male alive from Bengkulu Province, Sumatra, preserved female from Borneo. Photos: N. Maury.

September 201 4 | Volume 8 | Number 2 | e80

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A new species of Trimeresurus from Sumatra

erate canthus rostralis, totalling 28% of head length and

1.9 times as long as diameter of eye; a large, oval nostril

piercing in the middle of nasal scale; nostril-loreal pit

distance about 0.4 tunes the distance between the nostril

and the eye; eye average, amounting for 0.6 times the

distance between the lower margin of eye and upper lip

border; tail, tapering and prehensile. SVL 895 mm, TaL

155 mm, TL 1,050 mm; ratio TaL / TL 0. 148.

Body scalation

DSR: 21-21-15 scales, rhomboid, very weakly keeled

with the exception of scales of first dorsal scale row

which is smooth and not enlarged; 186 ventrals (+ two

pre-ventrals); 61 subcaudals, all paired; anal entke.

Head scalation

Rostral barely visible from above, triangular, about as

high as broad; nasals subrectangular, large, elongate, en-

tirely divided by a funow; two subrectangular, laterally

elongate internasals, about 1.8 times wider than long,

separated each from the other by one small scale; each

internasal followed on each side by one very large scale

on the snout, much larger than internasals, broader than

long, separated each from the other by two longitudinal

series of small scales; 2/2 canthal scales bordering the

canthus rostralis between intemasal and corresponding

supraocular, i.e., the very large scale behind intemasal

followed by a small scale smaller than adjacent snout

scales between the largest canthal scale and the supra-

oculars respectively; on each side, one elongate loreal

scale between nasal and the upper preocular; 2/2 pre-

oculars above the loreal pit, the upper one visible from

above, both scales elongate and in contact with loreal;

lower preocular fonning the lower margin of loreal pit;

1/1 thin, elongated, crescent-like subocular; 2/2 small

postoculars; 1/1 large supraoculai', broad, 2.2 times as

long as wide, about 1.3 times as wide as intemasal, not

indented by upper head scales; scales on upper snout sur-

face much enlarged, smooth, juxtaposed, subrectangular,

with four scales on a line between the scale separating

the intemasals and a line connecting the anterior margins

of eyes; six cephalic scales on a line between supraocu-

lars, smooth, flat, and juxtaposed; occipital scales flat,

smooth; temporal scales in two or three rows, smooth,

lower ones much enlarged; 9/9 SL, first SL triangular,

rather short, completely separated from the nasal; sec-

ond SL tall, bordering entirely the loreal pit and ante-

riorly in contact with nasal; third SL longest and high-

est, about 1 . 1 times longer than high, in contact on both

sides with subocular; fourth SL barely shorter than third

SL, in contaet with the subocular; fifth SL barely shorter

than fourth one, also in contact with the subocular; 10/11

infralabials, those of the first pair not longitudinally in

contact, shortly separated by the apex of the mental scale,

first-seeond IL in contact with anterior chin shields; four

rows of smooth gular scales; thioat shields irregularly ar-

ranged.

Coloration and pattern

The body is olive-green on third-eleventh DSR, slightly

paler on the bottom of the sides; most dorsal scales dis-

tinctly edged with black producing a reticulate pattern;

46 irregular black crossbars on each side of the body,

either symmetrical, forming black rings or more or less

set off from each other on each side across vertebral

line; scales of the first DSR and lower half of those of

the second DSR pale greenish-yellow with black edges,

producing a pale, diffuse ventrolateral stripe extending

from the area just behind the neck up to vent; scales of

the first DSR with a broad, irregular black edge on then-

anterior lower part. The tail is olive-green on its anterior

half, with scales strongly edged with black, producing a

strongly reticulate pattern, and two or three more or less

distinct black crossbars anteriorly, becoming progres-

sively msty red, strongly reticulate with black.

The head is olive-green above and on the temporal

region, with cephalic, occipital, and temporal scales

strongly edged with black; on each side, another long

streak extends from intemasals to the posterior part of the

head along the inner edge of supraoculars; two oblique

black streaks on the occiput forming an inverted V, its

apex pointing forward; supraoculars olive green, broad-

ly edged with black; some other cephalic and occipital

scales entirely black, so as to produce black blotches

and streaks between supraoculars; upper snout surface

heavily marked with pure black, producing broad edges

around olive-green prefrontals and intemasals; scale be-

hind and between intemasals black; black vertical and

horizontal streaks on the rostral; supralabials yellowish-

green, distinctly paler than upper head surface, strongly

edged with black, with a broader streak at the limit be-

tween third-fourth supralabials; preoculars olive-green

strongly edged with black; a black postocular streak ex-

tending from the eye to the area above the corner of the

mouth on lower postocular and the upper rows of tempo-

rals. Chin and throat pale greenish-yellow; an irregular

dark grey spot on eaeh infralabials of the first pair; all

infralabials edged with black on their posterior margin;

gular scales below the comer of the mouth also narrowly

edged with black.

The venter is pale greenish-yellow; each ventral nar-

rowly edged with black, especially on their central part,

sometimes on the whole of their posterior margin; tips of

ventrals narrowly edged with black on each side, produc-

ing an irregular, zigzag-like black ventral stripe below

the pale ventrolateral stripe. The tail is yellowish-green

or greyish-yellow below on its anterior half, with subcau-

dal scales broadly edged with black producing a conspic-

uous reticulation, turning more or less abruptly to msty

red, strongly reticulated with black.

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Vogel et al.

Diagnosis

A large species of pitviper of the genus Trimeresurus,

characterized by the combination of (1) body elongate,

head long and massive in adults; (2) body deep green

or dark green with conspicuous, black crossbars on the

sides in adults, with a broad, pale ventrolateral stripe, and

without sexual dimorphism in coloration; (3) 21 (excep-

tionally 23) DSR at midbody; (4) first supralabial dis-

tinct from nasal scale; (5) large intemasals, most usually

separated by one scale, only exceptionally in contact;

(6) three supralabials, third, fourth, fifth SL in contact

with subocular, or fifth exceptionally separated by one

scale; (7) supraoculars broad, separated by 3-7 (usually

5-6) scales; (8) tail average, with a ratio TaL/TL between

0.150 and 0.168 in males and 0.128 and 0.160 in females;

(9) 175-191 VEN; (10) 54-68 SC (males: 66-71; fe-

males: 54-68); (11) eye dark grey in life, rather bronze,

brown, dark greyish-brown or golden-brown in preserva-

tive; (12) supralabials and cephalic scales strongly and

broadly edged with black; (13) a black postocular streak;

(14) venter is yellowish-green or pale green, with each

ventral narrowly edged with black posteriorly; and (15)

tail green as the body on its anterior half, becoming more

or less abruptly salmon, pinkish-red, or rusty-red on its

posterior half, strongly and broadly reticulate with black.

Characters separating Trimeresurus sumatranus from

the new species and T. malcolmi are discussed below and

summarized in Table 2.

Description and Variation of T. sumatranus

(Fig. 5-9)

According to Brongersma (1933), Sanders et al. (2002),

Gumprecht et al. (2003), Das (2010), and our material,

this large species reaches a maximum total length of

1,355 mm. Gumprecht et al. (2003) mentioned a total

length of 1,400 nun. Males are seemingly shorter, the

longest male seen by us being only 878 mm long. Adults

reach usually a maximum total length of 90-110 cm.

The body is robust but elongate in both males and in

females, or slightly thinner in males. In adults, the snout

is 24.0-28.0% as long as head or 1.8-2. 6 times as long

as diameter of eye. Eye average, amounting for 0.6- 1.0

times the distance eye-lower edge of the lip in both

sexes. Ratio TaL/TL: 0.128-0.168, with a weak sexual

dimorphism: males: 0.150-0.168; females: 0.128-0.160.

Hemipenis

After Gumprecht et al. (2004: 304: Fig. IV), hemipenes

are long and slender, deeply forked, extending up to

twenty-secondth subcaudal, forked opposite ninth sub-

caudal, smooth at its base and after the forking point, for

about a third of the organ, then strongly papillose and

spinose, with longitudinal folds.

Body scalation

DSR: 21-25 one head length posterior to the head; 21

(exceptionally 23) at midbody; 15 (exceptionally 13 or

17) scales one head length before vent, weakly or dis-

Table 2. Main characters to distinguish between the species of the Trimeresurus sumatranus complex, source specimens from Ap-

pendix I, if not noted different.

Characters

Trimeresurus

gunaleni

spec. nov.

Trimeresurus

sumatranus

Sumatra

Trimeresurus

sumatranus

Borneo

Trimeresurus

sumatranus

Peninsular Malaysia

Trimeresurus

maicoimr

N males/females

5/4

2/13

3/19

2/5

3/4

Middorsal scale rows

21

19

Ventrals males

162-179

179-182

182-185

178-183

169-173

Ventrals females

164-174

175-186

176-191

180-186

168-174

Subcaudals males

71-72

66-70

66-71

69-70

64-81

Subcaudals females

58-66

57-68

54-64

61-66

61-64

Total length

1170

1152

1350

1220

1330

Relative tail length males

0.201-0.210

0.160-0.166

0.154-0.168

0.150-0.161

0.160-0.179"

Relative tail length females

0.144-0.180

0.130-0.159

0.128-0.150

0.134-0.160

0.158^

White lateral line^

Thin

Broad

Broad-

Broad

Absent

Ventrals with dark margins

No

Yes

Subcaudals with dark margins

No

Yes

Tail reddish

No

Yes

Eye in life

Green

Dark grey

'From Stuebing and Inger (1998).

“In two specimens there is no real ventrolateral stripe visible, but the outer row of dorsals is pale.

^Sometimes there is a faint black line below the white lateral line.

■*Holotype and one paratype only according to Loveridge (1938).

'^One paratype only according to Loveridge (1938).

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September 201 4 | Volume 8 | Number 2 | e80

A new species of Trimeresurus from Sumatra

tinctly keeled, smooth on first DSR; VEN: 175-191 (plus

1-2 preventrals); SC: 54-68, all paired, with a sexual di-

morphism (males: 66-71; females: 54-68); anal entire.

In our sample of 44 specimens, we have not examined

any specimen with 23 DSR at midbody but this value

has been recorded in the literature by Brongersma (1933)

from a specimen from Borneo. In our material, two spec-

imens had 13 scale rows before vent and only one had 1 7

rows before vent.

Head scalation

Rostral barely visible from above, triangular, wider than

high; nasals subrectangular, divided; one large, subrect-

angular or nearly square internasal on each side; intema-

sals usually separated by one scale or in contact (in 11/44

examined specimens); two (rarely three) canthal scales

bordering the canthus rostralis, one lai'ger than the large

adjacent snout scales and one (or two) of similar size; two

(exceptionally one) postoculars on each side; one very

large and wide supraocular on each side, about 1.8-2. 2

times as long as wide, 1.0-1. 3 times as wide as interna-

sal, not indented by adjacent cephalic scales; 4-7 much

enlarged scales on upper snout surface on a line between

the scale separating the internasals and a line connect-

ing the anterior margins of eyes, smooth and juxtaposed;

3-7 (usually 5-6) cephalic scales on a line between su-

praoculars, large, smooth, flat, and juxtaposed; occipi-

tal scales larger than cephalic scales, smooth; temporal

scales smooth, large, subequal, in two rows anteriorly,

three rows posteriorly; 8-11 (usually 9-10) supralabi-

als; first SL always separated from nasal; second SL tall,

entirely bordering the anterior margin of the loreal pit,

always in contact with nasal; third SL longest and high-

est, 1.1-1. 4 times as long as high, always in contact with

subocular; fourth SL as long as high, as high or barely

shorter than third SL, always in contact with subocular;

fifth SL usually in contact with subocular, exceptionally

separated by one scale (on one side in 3/44 specimens);

9-15 (usually 10-12) IL; scales of the first pair longitu-

dinally in contact or barely separated by the apex of the

long mental scale; first two or tliree pairs of infralabials

in contact with anterior chin shields; 5-9 rows of smooth

gular scales; throat shields irregularly arranged.

Coloration and pattern

In live adult specimens, the dorsum is yellowish-green,

grass green, deep emerald green, or olive green (deep

green, bluish-green, or dark brown in preservative);

many scales of the body nan'owly edged with black; usu-

ally a series of about 45-50 imegular, black crossbars one

or two dorsal scales in length, reaching downwards to

the third or second dorsal scale rows on each side of the

body. These crossbands are either symmetrical, fonning

black rings, or offset from each other on each side of the

vertebral line. Each crossbar includes one or two entirely

black scales, the other dorsal scales are strongly edged in

Amphib. Reptile Conserv. | amphibian-reptile-conservation.org (16)

black. The interstitial skin between the large dorsal scale

is also black, making an overall distinctly baned and

reticulate pattern. A more or less conspicuous, cream,

greenish-yellow, or pale yellow ventrolateral stripe on

the first and lower half to whole of the seeond dorsal

scale rows, extends from the base of the neck to the vent.

This pale stripe is bordered below by a narrow dark stripe

created by the black edge of the outer tips of the ventral

scales. The background color of the tail is as green as

the body on its anterior half, with scales distinctly edged

with black, producing a strongly reticulate pattern. There

are 2-5 uTegular black crossbars on each side of the tail

which, becomes more or less abruptly greenish-orange,

salmon, pinkish-red, or rusty-red on its posterior half.

The upper head surface and temporal regions are as

green as the body, the sides of the head and temporals

are slightly paler and usually more yellowish-green or

paler green. The rostral is green with black vertical and

horizontal streaks. Scales of the upper snout surface are

also green, broadly edged with black, and may be en-

tirely black with the exception of a large, round green

blotch on each internasal and each scale behind the in-

ternasal. Supraoculars green, broadly edged with black;

many scales of the cephalic and occipital regions black,

producing a pattern of black blotches or streaks. On each

side of the head, a long streak usually extends from in-

temasals to the posterior part of the head along the inner

edge of supraoculars; two oblique black streaks on the

occiput forming an inverted V; its apex pointing forward.

Supralabials greenish-yellow, pale green, yellowish-

green, or bluish-green, paler than the upper head surface,

strongly edged with black, black edges at the limits be-

tween third-fourth and fourth-fifth supralabials broader

and more conspicuous; preoculars and postoculars green

and black or entirely black; a black postocular streak

extends from the eye to the corner of the mouth on the

postoculars or lower postocular and the upper rows of

temporals, more or less broadly blotched with the green

background color. The chin and throat are white, cream,

pale greenish-yellow, or yellowish-green, uniform or

with scattered dark grey dots. The infralabials are white,

pale bright yellowish-green, or pale green, with or with-

out a few dark grey spots, edged with black on their pos-

terior margin; gular scales below the corner of the mouth

nan'owly edged with black. In life, the eye is rather dark,

bronze, brown, dark greyish-brown, or golden-brown.

The venter is yellowish-green, pale greenish-yellow,

or pale green; each ventral is nanowly edged with black

posteriorly, the edge usually broader on their central part

tips of ventrals narrowly edged with dark grey or black,

producing an inegular, zigzag-like dark ventral stripe be-

low the pale ventrolateral stripe. The under surface of the

tail is green or greenish-yellow on its anterior half, with

subcaudal scales broadly edged with black producing a

conspicuous reticulation, becoming more or less abmptly

greenish-orange, salmon or pinkish-red, strongly reticu-

lated with black.

September 2014 | Volume 8 | Number 2 | e80

Vogel et al.

Fig. 8. Live female of Trimeresurus sumatranus from vincity of Padang Panjang, Sumatera Barat Province, Sumatra. Photo: G.

Vogel.

Juveniles show a rather different pattern. The dorsum

is usually bright or grass-green, with only faint and nar-

row black spots or edges of dorsal scales producing faint,

diffuse crossbars. The head is speckled with black dots

but without dark lines and streaks on the edges of scales;

no black edges on the suture of supralabials but with

scattered black spots. The orange, salmon color or rusty

red color of the tail is brighter than in adults.

Distribution (Fig. 8)

Indonesia

Sumatra. Known from the provinces of Sumatera Barat,

Jambi, Bengkulu, and Lampung, in Barisan Range.

Borneo (Kalimantan). Seemingly throughout the island.

Federation of Malaysia

Borneo. Known from the States of Sabah and Sarawak.

West Malaysia. Definitely recorded from the States of

Perak (Sukumaran 2002 as Tropidolaemus wagleri, pers.

comm.), Johore, Pahang, and Trengganu.

Thailand

Recorded only from Yala Province.

In contrast to Sanders et al. (2004), we confirm the occur-

rence of T. sumatranus in extreme southern Thailand and

West Malaysia. Examined specimens present the com-

bination of all scalation and pattern characters, both of

the head and body, in full agreement with the definition

of this species. They all differ from Trimeresurus hageni

and we could not find any reason for not referring them

to T sumatranus. The range of T. sumatranus in Sumatra

is wider than indicated in Sanders et al. (2004) but the

records from the Indonesian islands of Bangka, Belitung,

Nias, Simeulue, and the Mentawai Archipelago (see, for

example, Brongersma 1933; Dring et al. 1990), are now

referred to the T. hageni group.

Natural History

This beautiful species inhabits regions typically covered

with equatorial rainforests, lowland tropical wet forests,

and tropical wet submontane forests, from sea level up

to about 1,000 m. The species shows a predilection for

lowlands in Borneo but, seemingly, only for hilly areas

at elevations between 650 and about 900 m in Sumatra

(Ryabov et al. 2002; Gumprecht et al. 2003; Sanders et

al. 2004). This pitviper is found in tropical forests, along

clearings, in bamboo thickets, mangroves, swamps,

plantations, and cultivated fields such as coffee and tea

estates. However, in Sumatra, all specimens recorded

by Ryabov et al. (2002) and Gumprecht et al. (2003) in

Bengkulu Province (Sumatra) were found in forest, none

in cultivated areas or near villages.

September 201 4 | Volume 8 | Number 2 | e80

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A new species of Trimeresurus from Sumatra

Fig. 9. Live female of Trimeresurus sumatranus from Bengkulu Province, Sumatra. Photo: G. Vogel.

Fig. 10. Live female of Trimeresurus malcolmi from Mount Kinabalu, Sabah, Borneo. Photo: M. Dehling.

September 201 4 | Volume 8 | Number 2 | e80

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Vogel et al.

Fig. 11. Live female of Trimeresurus malcolmi from Mount Kinabalu, Sabah, Borneo. Photo: M. Dehling.

Fig. 12. Live male of Trimeresurus toba from vincity of Padang Panjang, Sumatera Barat Province, Sumatra, a species sympatric

with T. gunaleni spec. nov. Photo: G. Vogel.

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September 201 4 | Volume 8 | Number 2 | e80

A new species of Trimeresurus from Sumatra

TRIGUN0002 Q

Fig. 13. MZB.Ophi.5452 holotype of Trimeresurus gunaleni spec, nov., adult female. Photo: N. Maury.

Fig. 14. Live female of Trimeresurus sumatranus from Bengkulu Province, Sumatra. Photo: N. Maury.

9

TRISUM0001

September 201 4 | Volume 8 | Number 2 | e80

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Vogel et al.

Trimeresurus siimatranus often occurs along the

banks of rivers, ponds, and other watered areas. This di-

urnal and nocturnal species is chiefly arboreal but lives

in the lower vegetation such as in thick bushes, shrubs,

and the tangled lower tree foliage up to 2.5 m above the

ground, where it proves to be a skilled climber. Ryabov et

al. (2002) found specimens basking in early morning. Tr-

imeresurus sumatranus feeds on small mammals, frogs,

lizards, and frogs. It is oviparous, but little is known on

its breeding habits. Ryabov et al. (2002) mentioned a

clutch of 17 eggs that were guarded by the female; we

refer to Ryabov et al. (2002) and Gumprecht et al. (2003)

for additional data on the biology of this species.

In our sample of specimens identified in collections

as Trimeresurus sumatranus, we identified a total of six

specunens that present noteworthy morphological differ-

ences with the species as defined above. We also noted

the same differences in three specimens that were kept

alive. As these differences with T sumatranus are con-

stant, we consider these specimens to be referable to a

distinct species that we here describe as:

Trimeresurus gunaleni spec. nov.

Fig. 3-7, 13

iim;lsid:zoobank.org:act:548DBAC7-D5CC-4D49-9D4F-2A372F0F4520

Trimeresurus sumatranus (nee Coluber sumatranus Raf-

fles, 1822): Sanders et al. (2002: 107, part.; 2004: 722,

part.).

Holotype

MZB.Ophi.5452, adult female, fi-om Mt. Sibayak, ca.

1,500-2,200 m a.s.l., west of Brastagi (Berastagi), Karo

Regency (Kabupaten Karo), Sumatera Utara Province,

Sumatra, Indonesia. Collected by the team of Danny Gu-

nalen, Hidekazu Miyake, Cho Sangyeon, and Moon Suk

Cha.

Paratypes (six specimens)

NHMW 28159:1 (male), ZMB 29642 (male), NHMW

23909:4, NHMW 28159:2 (females), “Padang, Suma-

tra;” NHMB 2599 (male), “Solok, Sumatra;” SMF 52844

(female), “Padang Mountains, Sumatra, 1,700 m,” all

from Sumatera Barat Province, Sumatra.

Non-type material (two live male specimens)

Mt. Singkut, 1,600 m a.s.l., Karo Regency, Sumatra

Utara Province, Sumatra.

Diagnosis

A large species of pitviper of the genus Trimeresurus,

characterized by the combination of (1) body elongate,

head long and massive in adults; (2) an overall green

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coloration with interstitial skin forming irregular, hol-

low, black dorsal crossbands, with a thin, pale ventro-

lateral line; (3) 21 DSR at midbody; (4) first supralabial

totally separated from nasal scale; (5) large internasals,

most usually separated by one scale, only exceptionally

in contact; (6) three supralabials, third, fourth, fifth SL in

contact with subocular; (7) supraoculars large but elon-

gate, separated by 5-7 cephalic scales; (8) tail long, with

a ratio TaL/TL between 0.201 and 0.210 in males and

0.144 and 0.180 in females; (9) 162-179 VEN; (10) 58-

72 SC (males: 71-72; females: 58-66); (11) eye yellow-

ish-green in life and preservative; (12) cephalic scales

strongly and broadly edged with black but not forming

streaks; (13) no black postocular streak; (14) venter

greenish-yellow or pale green, unifomi, with posterior

margin of ventrals paler green; and (15) tail greyish-red,

rusty brown or reddish-brown, mottled with green cross-

bars anteriorly.

Main characters separating T. gunaleni spec. nov.

from other taxa of the complex of T sumatranus are

summarized in Table 2. Trimeresurus gunaleni spec. nov.

mainly differs from T sumatranus by (1) a lower number

of ventrals in males (162-179, x= 168.4 vs. 178-185, x

= 181.5; U = 33.5, P < 0.005) and females (164-171, x

= 169.5 vs. 175-191, 183.3); (2) a higher value of the

ratio TaL/TL in males (0.201-0.210, x = 0.206 vs. 0.150-

0.168, X = 0.161); (3) the color of the tail with hues of

red throughout mottled with green crossbars anteriorly

vs. green as the body on its anterior half, becoming more

or less abruptly red (see above description) posteriorly,

strongly reticulate with black; (4) the color of the eyes:

green or yellowish-green in T. gunaleni spec. nov. vs.

daik brown, dark grey, or bronze in T. sumatranus; (5)

the color of the ventral scales, which are green with a

paler posterior margin in T gunaleni spec. nov. vs. pale

green with a dark grey or black posterior margin in T

sumatranus.

Trimeresurus gunaleni spec. nov. differs from T mal-

colmi by (1) the number of dorsal scales ai'ound mid-

body (21 vs. 19); (2) a higher value of TaL/TL in males

(0.201-0.210, X = 0.206 vs. 0.160-0.179, x = 0.162, .y

= 0.009); (3) the presence of a white lateral stripe in T

gunaleni spec, nov., missing in T. malcolmi; (4) the color

of the tail: greyish-red, rusty brown or reddish-brown,

mottled with green crossbars anteriorly in T. gunaleni vs.

greenish-orange, salmon or pinkish-red, strongly reticu-

lated with black in T malcolmi; below, the tail is yellow-

ish green anteriorly, turning to brown posteriorly in T

gunaleni spec. nov. vs. green or greenish-yellow on its

anterior half, with subcaudal scales broadly edged with

black producing a conspicuous reticulation; (5) the color

of the eyes: yellowish-green in T gunaleni spec. nov. vs.

dark grey in T. malcolmi.

Trimeresurus gunaleni spec. nov. differs from T. hage-

ni by (1) a lower number of ventrals in males (162-179,

.T= 168.4 against 177-189, x= 181.8; U= 139.5, P <

0.001) and females (164-171, x= 169.5 vs.176-196, x =

September 201 4 | Volume 8 | Number 2 | e80

A new species of Trimeresurus from Sumatra

186.7); (2) by the intemasals being separate (in eight out

of nine specimens) against being most usually in contact

in T. hageni (in 66 out of 73 specimens; U = 545.5, P <

0.001); (3) by the number of supralabials, usually being

nine (in 14 out of 18 cases, only exceptionally 8 or 10,

X = 9.11) in T. gunaleni spec. nov. vs. usually 10 or 11

(in 123 of 148 occuiTences, exceptionally 9, 12, or 13, x

= 10.54; U = 637.0, P < 0.001) in T. hageni’, (4) by the

total number of supralabials (on both sides) touching the

subocular, six (in one case in seven, x = 6.1) vs. usually

being 2^ (in 56 out of 74 cases, exceptionally 0, 1,5,

6,x = 2.9; U = 632.0, P < 0.001) in T. hageni’, (5) by the

number of infralabials, usually 11 (in 11 out of 16 occur-

rences, only exceptionally 10 or 12, x = 10.93) vs. usu-

ally 12-14 (in 120 of 148 cases, exceptionally 11, 15, or

16, X = 13.08; t/= 710.0, P< 0.001) in T. hageni’, (6) the

lack of a pale temporal streak in T. gunaleni spec, nov.,

usually present in T. hageni, especially in males (in 27 of

29 males of T. hageni)’, and (7) the missing of dorsolat-

eral white dots in T. gunaleni spec, nov., are usually pres-

ent in T. hageni especially in males (in 27 of 29 males of

T. hageni).

Etymology

The specific nomen is dedicated to Mr. Danny Gunalen,

who was the first to find the species alive and who great-

ly supported the work resulting in the description of this

new species. Suggested common names: English: Gu-

nalen’ s Pitviper. Bahasa Indonesia: Ular Hijau Gunung.

Karo: Nipe Ratah. Padang (Minang): Ular Ijo Babiso.

French: Trimeresure de Gunalen. German: Gunalen’s

Gmbenotter.

Description of the holotype (Fig. 3, 5-7, 13)

Body elongate, compressed; head elongate, distinctly tri-

angular, wide at its base, clearly distinct from the neck,

flattened anteriorly, thick posteriorly, 1 .6 times as long as

wide; snout long, round when seen from above, strongly

obliquely tmncated when seen from the side, with a mod-

erate canthus rostralis, totaling 32.0 % of head length,

and 2.7 times as long as diameter of eye; a large oval nos-

tril piercing in the middle of nasal scale; nostril-loreal pit

distance about 0.5 times the distance between the nostril

and the eye; eye average, totaling 0.65 times the distance

between the lower margin of eye and upper lip border;

tail rather long, tapering, and prehensile.

SVL 995 mm, TaL 195 mm, TL 1,170 mm; largest

head width 35.0 mm; ratio TaL / TL 0.167.

Body scalation

DSR: 21-21-13 scales, rhomboid, distinctly keeled with

the exception of scales of first DSR which are smooth;

171 VEN (+ two preventrals); 60 SC, all paired; anal en-

tire.

Amphib. Reptile Conserv. | amphibian-reptile-conservation.org (22)

Head scalation

Rostral barely visible from above, triangular, much

broader than high; nasals pentagonal, partly divided by a

shallow furrow; on each side, one large, subtriangular in-

temasal, the rounded apex pointing outwards; internasals

separated by one small scale; 2/2 canthal scales border-

ing the canthus rostralis, not larger than the large adja-

cent snout scales on each side, one elongate loreal scale

between nasal and the upper preocular; 2/2 preoculai'S

above the loreal pit, the upper one visible from above,

both scales elongated and in contact with loreal; lower

preocular forming the lower margin of loreal pit; 1/1

thin, elongated, crescent-like subocular; 2/2 small post-

oculars, followed by 2/3 small scales between postocu-

lars and first temporals; 1/1 large, subtriangular, elongate

supraocular on each side, 1.8 times as long as wide, 1.1

times as wide as intemasal, not indented by adjacent ce-

phalic scales; three much enlarged scales on upper snout

surface on a line between the scale separating the inter-

nasals and a fine connecting the anterior margins of eyes,

smooth and juxtaposed; seven cephalic scales on a fine

between supraoculars, smaller than upper snout scales,

smooth, flat, and juxtaposed; occipital scales not larger

than cephalic scales, smooth; temporal scales smooth,

large, subequal, an'anged in two rows anteriorly, three

rows posteriorly; 9/9 supralabials, third-fifth SL in con-

tact with subocular; first SL entirely separated from na-

sal; second SL tall, entirely bordering the anterior margin

of the loreal pit, in contact with nasal; third SL longest

and highest, 1.4/ 1.5 times longer than high; fourth SL tall

and relatively nan'ow, 1.1/1. 2 times higher than long, as

high as third SL; fifth SL relatively narrow; 12/12 IL;

scales of the first pair longitudinally in contact; first three

pairs of infralabials in contact with anterior chin shields;

six rows of smooth gular scales; throat shields irregularly

arranged.

Coloration and pattern

The body is unifonnly deep green (bright emerald green

in fife), with some scales narrowly edged with black,

more strongly on the fore part of the body; a faint, diffuse

pattern of black, hollow crossbars resulting from irregu-

lar areas of interstitial skin around dorsal scales, more

conspicuously visible on the fore part of the body, pro-

ducing about 25 crossbars, three or four DSR long and

separated by one or two scales around which the skin is

grey, reaching downwards the first or second DSR; from

about midbody, the black skin is progressively restricted

to the eighth or ninth DSR, producing irregular dorsal

bars, disappearing entirely before the vent; a naiTow ven-

trolateral stripe, pale blue in fife, white in preservative,

extends from the neck to the vent on the upper edge of

scales of the first DSR, and lower edge of scales of the

second DSR; posterior edge of scales of the first DSR

also pale blue. The tail is greyish-red throughout, with

scales narrowly edged with pale grey and with five pale

green incomplete rings anteriorly.

September 2014 | Volume 8 | Number 2 | e80

Vogel et al.

The head is deep green above (bright emerald green in

life) and on the temporal region, with scales of the snout,

preoculars, supraoculars, cephalic, occipital, and lower

temporal scales narrowly edged with black and surround-

ed with black interstitial skin, producing a conspicuous

pattern of a “mixed” black background with bright green

spots; no cephalic or occipital streaks; supraoculars nar-

rowly edged with black; top of rostral black; anterior

supralabials greenish-yellow, distinctly paler than upper

head surface, others supralabials bright yellow in life;

first-third SL narrowly edged with black posteriorly; no

postocular streak; upper temporals green as the upper

head surface. Chin and throat pale bluish-grey (cream in

preservative); mental and first three infralabials green-

ish-yellow; other infralabials more or less marbled with

greenish-yellow; posterior gular scales dotted with green.

The venter is uniformly yellowish-green, with the

posterior edge of each ventral pale bluish-grey, distinctly

paler than the background color of the venter. The tail is

greenish-yellow on the first two subcaudals then greyish-

red throughout as the upper surface of tail, with scales

narrowly edged with pale grey.

Description of the paratypes

A summary of morphological and meristic data of the

paratypes is given in Table 3. None of the paratype sig-

nificantly differs from the description given for the ho-

lotype.

Description and variation

The maximal confirmed total length known is 1,170 mm

(SVL 995 mm, TaL 195 mm; holotype). The second larg-

est female has a length of 1,154 mm (SVL 972 mm, TaL

182 mm; NHMW 28159:2, from Padang). The largest

known male is 927 mm long (SVL 732 mm, TaL 195

mm; NHMW 28159:1, from Padang). In our sample of

nine specimens, there is a noteworthy difference of size

between males and females (see below, sexual dimor-

phism).

The body is robust but elongate in both males and in

females. In adults, the snout is 28.0-32.8 % as long as

head or 2. 3-2. 8 times as long as diameter of eye. Eye

average, amounting for 0.7-0. 9 times the distance eye-

lower edge of the lip in both sexes. Ratio TaL/TL: 0. 144-

0.210, with a sexual dimorphism (see below).

Hemipenis

Unknown.

Body scalation

DSR: 21-22 one head length posterior to the head, 21 at

midbody, 13-15 scales one head length before vent, dis-

tinctly keeled, smooth on first DSR; VEN: 1 62-1 79 (plus

preventrals), without sexual dimorphism; SC: 58-72, all

paired, with a sexual dimorphism (see below); anal en-

tire.

In our sample of nine specimens, only one specimen

has 22 DSR on the neck, all others have 21 rows. Eur-

theiTnore, all males have 13 rows before the vent but two

females have 15 rows.

Head scalation

As described for the holotype, with the following varia-

tion for major characters: intemasals separated by one

small scale in 8/9 specimens, in contact only in speci-

men ZMB 29642; only two canthal scales on each side

in all specimens, not larger than adjacent snout scales or

slightly smaller, bordering the canthus rostralis between

the internasal and corresponding supraocular; two small

postoculars, in contact with first temporals or followed

by 2-3 small scales between postoeulars and first tem-

porals; one large, elongate, subtriangular supraocular on

each side, 1. 6-2.1 times as long as wide, 1.0-1. 3 times

as wide as internasal, not indented by adjacent cephafic

scales; 3-4 enlarged scales on upper snout surface on a

line between the scale separating the internasals and a

line connecting the anterior margins of eyes, smooth, and

juxtaposed; 5-7 cephalic scales (5: 1/9 specimens; 6: 5/9;

7: 3/9) on a line between supraoculars, smaller than up-

per snout scales, smooth, flat, and juxtaposed; occipital

scales not enlarged and smooth; temporal scales smooth,

large, subequal, in two or three rows; 8-10 supralabials

(8: 2/18 occurrences; 9: 12/18; 10: 4/18); third, fourth,

fifth SL in contact with subocular in all specimens; first

SL always separated from nasal; second SL tall, entirely

bordering the anterior margin of the loreal pit, always in

contact with nasal; third SL longest and highest, 1.2-1. 5

times as long as high; fourth SL higher than long; fifth SL

tall and narrow; 10-12 IL (10 or 11 in most specimens);

scales of the first pair longitudinally in contact; first three

Table 3. Morphological characters of the paratypes of Trimeresurus gunaleni spec. nov. M: male, F: female, for other abbreviations

see Table 1 .

Collection number

Sex

SVL (mm)

TaL (mm)

TaL/ TL

VEN

SC

SL

SL touching

sublabial

Cep

IL

Do

NHMW 28159:1

M

732

195

0.210

162

71

9/9

6

11/10

21

NHMB 2599

M

651

inc.

175

inc.

9/9

6

11/11

21

ZMB 29642

M

638

165

0.205

179

inc.

9/9

6

11/11

21

NHMW 23909:4

F

309

52

0.144

174

58

10/10

7

6

11/11

21

NHMW 28159:2

F

972

182

0.158

169

58

9/10

6

11/11

21

SMF 52844

F

400

88

0.180

164

66

9/9

6

7

9^/ 9^=

21

inc.; Tail incomplete or partly destroyed.

^Destroyed.

Amphib. Reptile Conserv. | amphibian-reptile-conservation.org (23) September 2014 | Volume 8 | Number 2 | e80

A new species of Trimeresurus from Sumatra

pairs of infralabials in contact with anterior chin shields;

5-8 rows of smooth gular scales; throat shields irregu-

larly arranged.

Coloration and pattern

The body is bluish-green or deep green (bright green or

emerald green in life) with most of the dorsal scales nar-

rowly edged with black, usually more strongly on the

anterior part of the body; a faint, diffuse pattern of dark,

hollow crossbars created by irregular areas of black in-

terstitial skin surrounding three or four rows of dorsal

scales, most conspicuous and extensive on the fore part

of the body, separated by one or two scales around which

the skin is grey; these dark crossbands reach downwards

the first or second DSR on the anterior part of the body,

progressively restricted to the upper DSR posteriorly,

producing irregular dorsal bars and disappearing entirely

before the vent; a narrow ventrolateral, stripe, white,

cream, or pale yellow in preservative (cream or pale blue

in life), extends from the neck to the vent on the upper

half of scales of the first DSR and sometimes on low-

er edge of scales of the second DSR; posterior edge of

scales of the first DSR also white or cream (cream or pale

blue in life). The tail is greyish-red or rusty-red through-

out, with scales narrowly edged with cream to pale grey,

and mottled with incomplete cream, pale grey or pale

greenish-grey rings (pale greyish-green in life), present

on the anterior half of the tail or throughout.

The head is deep green above and on the temporal

region; scales of the snout, preoculars, supraoculars,

cephalic, occipital and lower temporal scales narrowly

edged with black and entirely surrounded with interstitial

black skin, producing a conspicuous pattern made of a

black background “mixed” with bright green spots; no

cephalic, occipital, or postocular streaks; supraoculars

narrowly edged with black; top of rostral usually black;

anterior supralabials green or yellowish-green, distinctly

paler than upper head surface, others supralabials yellow,

greenish-yellow or green; first-third SL usually narrowly

edged with black on their posterior edge; upper tempo-

rals green as the upper head surface. Chin and throat

cream (pale bluish-grey in life); mental and first three in-

fralabials greenish-yellow or pale yellowish-green; other

infralabials more or less marbled with greenish-yellow;

posterior gular scales sometimes dotted with greyish-

green or green spots.

Fig. 15. Habitat of Trimeresurus gunaleni spec. nov. Photo: D. Gunalen.

September 201 4 | Volume 8 | Number 2 | e80

Amphib. Reptile Conserv. | amphibian-reptile-conservation.org (24)

Vogel et al.

The venter is uniformly bluish-green or yellowish-

green in preservative (yellow, greenish-yellow, or green

in life), with the posterior edge of each ventral pale blu-

ish-grey or greyish-green, distinctly paler than the back-

ground color of the venter. The tail is greenish-yellow or

green anteriorly on a distance varying from the first sub-

caudals to the middle of the tail then greyish-red or rusty-

red throughout as the upper surface of tail, with scales

narrowly edged with pale grey and with cream, pale grey

or pale greenish-grey blotches (pale greyish-green in

life) corresponding to the rings of the upper surface.

Sexual dimorphism

Males and females differ in the relative length of the tail,

in total length, and in the number of subcaudals:

(1) Strong difference in the ratio TaL/TL:

males: 0.201-0.210 (J =0.206); females: 0.144-0.180

(J =0.162).

(2) Total length:

Largest male: 927 mm vs. largest female 1,170 mm.

(3) Differences in the number of subcaudals:

71-72 (J=71.5) in two males vs. 58-66 (x =60.5) in

four females.

There is no difference in the numbers of ventral scales

or in other scalation characters, nor in pattern or in eye

color.

Distribution

Indonesia

Sumatra. Endemic; Trimeresurus gunaleni spec. nov. is

known only from two provinces: Sumatera Barat (Solok

and Padang Mountains) and Sumatera Utara (Mt. Sibay-

ak, Mt. Sinabung and Mt. Singkut near Berastagi).

This species can be expected in higher elevations all

over the mountainous areas of Sumatra.

Natural History

Trimeresurus gunaleni spec. nov. inhabits regions typi-

cally covered with tropical moist montane forests, from

1,500 m to as high as at least 2,000 m, perhaps as much

as 2,200 m, where it has been observed by local insect

collectors (Figs. 15 and 16). There is no record of popu-

lations lower than 1,500 m. On Mount Sibayak, Danny

Gunalen collected specimens of Trimeresurus hageni at

elevation of 500 m, and Tropidolaemus wagleri at 200 m.

Trimeresurus gunaleni is clearly isolated as a high mon-

tane dweller.

The female holotype of T. gunaleni spec. nov. was col-

lected during the daytime in dense humid montane for-

est scattered with tiny springs. The snake was resting on

the ground under tree roots. In another instance, a male

was seen perched at night on a tree branch at about two

m above the ground. None of the specimens were found

near open water, the biotopes are dense humid montane

forests.

Based on regurgitated prey items and direct observa-

tions in the wild, the diet includes rodents, amphibians,

and lizards (Gonocephalus lacunosus Manthey and Den-

zer, 1991; E. Manik, pers. concun.). In captivity, T. gu-

naleni spec. nov. feeds on mice, birds, and lizards (D.

Gunalen, pers. comm). Reproductive habits are still un-

known.

Discussion

The differences in pholidosis and coloration, together

with the fact that T gunaleni spec. nov. and T. sumatra-

nus are living in close proximity, leaves no doubt that T

gunaleni spec. nov. deserves full species status. Although

T. sumatranus and T gunaleni spec. nov. have been re-

corded from the same mountain ranges in Sumatera Barat

Province, it is not yet known whether these two species

are living in sympatry or syntopy. However T sumatra-

nus seems to live at lower elevations than T. gunaleni

spec. nov. Too little is known about the exact ranges of

both species in western Sumatra to ascertain if there is a

zone of true sympatry.

The recognition of T gunaleni brings the number of

species in the subgenus Parias on Sumatra to three. Su-

matra is inhabited by T sumatranus, T hageni, and T.

gunaleni spec, nov., whereas Borneo is the home of T.

sumatranus and T malcolmi. This latter species is obvi-

Fig. 16. Habitat of Trimeresurus gunaleni spec. nov. Photo: D.

Gunalen.

September 201 4 | Volume 8 | Number 2 | e80

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A new species of Trimeresurus from Sumatra

ously the highland equivalent of T gunaleni spec. nov.

In the Malay Peninsula, only T. sumatranus is known,

although this species is rarely collected there. The rela-

tionship between the Malayan population and the other

two populations of T. sumatranus are not known. In for-

mer reviews of this complex (Sanders et al. 2004), the

existence of T. sumatranus in Peninsular Malaysia was

denied, despite the fact that there are five specimens

available in the collections of the Natural History Mu-

seum of London. Re-examination of these specimens and

new specimens leave no doubt about the occurrence of

T. sumatranus in West Malaysia and extreme southern

Thailand. So far no highland population corresponding

to T. gunaleni spec. nov. or T. malcolmi have been found

in Peninsular Malaysia.

A rather similar scheme of relationships between

zoogeographical entities of the Sunda Shelf can also

be defined in other pitvipers. In the subgenus Popeia of

the genus Trimeresurus, i.e., the complex of Trimeresu-

rus popeiorum Smith, 1937 (see Vogel et al. 2004), the

species T. barati Regenass and Kramer, 1981 is known

from western and south-western Sumatra whereas T. toba

David, Petri, Vogel and Doria, 2009, inhabits mountains

of central northern Sumatra. In Peninsular Malaysia, T

fucatus Vogel, David, and Pauwels, 2004 is widespread

but T. nebularis Vogel, David, and Pauwels, 2004 is cur-

rently considered endemic to the Cameron Highlands.

However, only T sabahi Regenass and Kramer, 1981 is

known in Borneo. As in the subgenus Parias, there is no

species of the subgenus Popeia known from Java. The

situation is shghtly different for the subgenus Craspedo-

cephalus, i.e., the complex of Trimeresurus puniceus (see

David et al. 2006). Here we do have T. puniceus (Boie,

1827) widely distributed in Java and in southern Suma-

tra, but also a distinct species, T andalasensis David, Vo-

gel, Vijayakumar, and Vogel, 2006 in northern Sumatra.

Another species, T wiroti Trutnau, 1981 is known from

Peninsular Malaysia and southern Thailand, whereas T.

borneensis Peters, 1872 is widespread in Borneo.

Trimeresurus hageni and T. puipureomaculatus are

both distributed on Sumatra and Peninsular Malaysia

(David and Vogel 1996) but not in Borneo. The system-

atics of both species is not resolved and there might be

more taxa hidden under these names. The distribution of

T. purpureomaculatus is restricted to mangrove areas.

Lastly, Trimeresurus albolabris lives in the south of Su-

matra (David and Vogel 2000) and on Java (Creer et al.

2003) but is unknown from West Malaysia and Borneo.

These species or complexes of pitvipers show the

close zoogeographic relationships of the islands of Bor-

neo and Sumatra with Peninsular Malaysia. Furthermore,

it can also be seen that Sumatra is split into a northern

and a southern region, with the larger northern region

closely connected to Western Malaysia and Borneo, and

the smaller southern region connected with Java. The

limit between these two regions seems to be located be-

tween Padang and Bengkulu. Previously, these species

Amphib. Reptile Conserv. | amphibian-reptile-conservation.org (26)

complexes were regarded as widely distributed species,

obscuring the zoogeographical relations of these regions.

We are not confident that the taxonomy of the genus

Trimeresurus is fully resolved and previously mentioned

taxa might still prove to be endemic for one of the re-

gions.

The finding of such a large and venomous pitviper as

T gunaleni spec. nov. in a group that was supposed to be

well known is quite surprising. It is hard to understand

that it was overlooked for such a long time despite the

fact that the three specimens in the collection of Vienna

have been available for a long time (collected 1899) and

were already examined by other groups of herpetolo-

gists. The mountainous areas of Sumatra are still very

incompletely known and further research in these areas

is highly desirable.

Acknowledgments. — ^We are grateful to Nathanael

Maury (Singapore) and Ludovic David (Bali) for pic-

tures and information of the live specimens and help in

various aspects.

We want to thank the following persons, who gave

us access to specimens in their care: Colin J. McCarthy

and Patrick Campbell (BMNH), Jens Vindum and Alan

Leviton (CAS), Alan Resetar and Harold Voris (FMNH),

Georges Lenglet (IRSNB), Alain Dubois, Ivan Ineich,

and Annemarie Ohler (MNHN), Heinz Grillitsch, Silke

Schweiger, and Richard Gemel (NHMW), Denis Val-

lan and Raffael Winkler (NHMB), Kiyotaka Hatooka

(OMNH), Esther Dondorp, Pirn Amtzen, and Ronald

de Ruiter (RMNH), Gunther Kohler and Linda Acker

(SMF), Dennis Rodder, and Wolfgang Bohme (ZFMK),

Mark-Oliver R5del and Frank Tillack (ZMB), Jakob

Hallermann (ZMH), Kelvin P. P. Lim (ZRC), and Frank

Glaw and Michael Franzen (ZSM).

We also thank Jakob Hallermann (ZMH) and Frank

Tillack (ZMB) who sent us pictures and data about speci-

mens in their collections. Max Dehling (Koblenz) pro-

vided us with pictures of a live specimen of Trimeresurus

malcolmi, and Jeet Sukumaran (Durham) who provided

us with additional data on the distribution of T suma-

tranus. Wolfgang Bohme (Bonn) provided us with data

from ZFMK specimens.

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Appendix I. Additional specimens examined

Trimeresurus hageni (n=73). INDONESIA. Sumatra. BMNH 89.12.26.20, District of Deli; BMNH 93.6.5.11,

East coast of Sumatra; MNHN 1880.0042, Sumatra; MZB 446, Pulau Batu, west Sumatra; MZB 1740, Padang

Bukit Sebelah; MZB 1892a, b, Ketambe Aceh Tenggara; MZB 1898, Aceh Barut; MZB 2886, a, b Kembang Ma-

nis, Bengkulu; MZB 3716, Kubu Peraka Primer; NHMB 5108, Pelambeng, South Sumatra; NHMB 9423, Suma-

tra; NHMW 23909:1-2, Medan; NHMW 23909:3, Deli; NHMW 28150:1-3; Padang; NHMW 28150:4, Pagay;

NHMW 28155:3, Medan; RMNH 5587A, Deli; RMNH RENA 819 (lectotype). Deli; ZFMK 32508, Sumatra; ZMB

15884, Sukuranda, Oberer Langkat, O-Sumatra; ZMB 29642, Padang; ZMB 32193a, b, Sumatra; ZMB 62699,

Aceh; ZMB 66176, Bengkulu province; ZMH R06937, Serdang; ZSM 109/1927, Goenoeng Rintels, S. Deli; ZSM

202-1979a, Lau Rakit, near Deli; ZSM 202-1979b, Gunoeng Rinteh. Banka. RMNH 4697, Banka; ZSM 365/1907

(4), ZSM 365/1908 (1-3) Simpang, Banka. SINGAPORE. BMNH 80.9.10.6, Singapore. MALAYSIA. West Ma-

laysia. BMNH 1936.9.12.5, “Kualla Taku, Malay Penin;” BMNH 1936.9.91, Perak; BMNH 1967.2290-1, Gu-

nong Benom; CAS 16831, Silensing, Pahang; MNHN 1899.0269, Peninsular Malaysia; MNHN 1974.0044, Kuala

Lumpur; NHMW 28158:1-2, Kedah; PSGV 393, Kuala Lumpur; S 0117 “West Malaysia;” SMF 64464-5, Perak;

ZFMK 16680, Yombak; ZFMK 68522, north of Kuala Lumpur; ZMB 70235, frim, Selangor; ZRC 2.2928, Tasik

Bera, Pahang; ZRC 2.2930, Tembeling, Pahang; ZRC 2.2932, Bukit Lagong Forest, Selangor; ZRC 2.2933-4, Ulu

Langat, Selangor; ZRC 2.2935, Cameron Highlands, Pahang; ZRC 2.2943, Negeri Sembilan, Gunung Angsi; ZRC

2.5362, Bellum, Perak; ZRC 2.5397, Kepong, Frim, Selangor. THAILAND. BM 1936.9.12.4, Belong, Yala; BM

1988.858-62, Trang; IRSNB 3059 Belong, no locality: ZFMK 18835, no locality; ZFMK 21497, Sunda Islands.

Amphib. Reptile Conserv. | amphibian-reptile-conservation.org (28)

September 201 4 | Volume 8 | Number 2 | e80

Vogel et al.

Gernot Vogel was bom in Heidelberg, Germany and received a Ph.D. in Chemistry. Dr. Vogel is now working

as a chemist at an international company, doing research and registration of plant protection products in Hirsch-

berg, close to Heidelberg. Beginning as a reptile keeper. Dr. Vogel developed a great interest in the snake fauna

of the Oriental region. He has concentrated his work on southwestern China, Indonesia, and tropical India. Dr.

Vogel has revised large snake genera with a wide distribution, and this has been achieved through international

collaborations with, for example, the institutes of Chengdu and Kunming in China, with Lipi and KPH Salvator

in Indonesia, and with various Indian groups including ARRS and ANET. His research is based on specimen

collections all over the world and on field research in the regions cited above. Dr. Vogel has authored or co-

authored the following books: The Snakes of Sumatra', The Snakes of Sulawesi', Amphibians and Reptiles of

Mount Kinabalu', and parts I to III in the Terralog Series on the venomous snakes of the world.

Patrick David (born in 1959 near Paris, France), received a Ph.D. in polymers chemistry at the University

of Paris-Orsay. Dr. David developed an early interest in herpetology as a reptile keeper and then turned to the

systematics of Asian reptiles. He is, or has been a member of several international herpetological societies.

Dr. David has been involved for nearly 25 years, mostly with Gernot Vogel, in systematic research on several

groups of Asian reptiles, especially the Trimeresurus-complox and the genera Oligodon, Amphiesma, Xeno-

chrophis, and Cyrtodactylus. His geographic areas of interest include India, Thailand, China, Vietnam, and

especially Laos and Sumatra (Indonesia). Dr. David has also addressed problems of nomenclature affecting

various taxa of snakes and lizards. As of July 2014, Dr. David is the author or co-author of 121 publications,

including five monographs or books. He has co-authored the description of a total of 31 new species of am-

phibians and reptiles. Along with Gemot Vogel, Dr. David is preparing a monograph on Asian pitvipers and

the snake fauna of Sumatra.

Irvan Sidik was bom in Bandung, West Java Province, Indonesia. Irvan obtained a Masters of Science degree

in the field of phylogenetics at the Institute Technology of Bandung. Since 1992 Irvan has worked as a staff

researcher in the laboratory of herpetology at the Museum Zoologicum Bogoriense, Indonesian Institute of

Sciences (LIPI) in the Cibinong Science Center. Beginning as an auxiliary field survey researcher, and then

as a local CITES officer, Irvan became interested and developed a great interest in the snakes of the region

of Sundaland. Irvan has continued with more scholarly work on the mountainous areas of the western part

of Indonesia. Irvan’s research is based on museum collections of specimens and field research in Indonesia’s

regions mentioned above. Irvan has been involved in several international research collaborations, and is cur-

rently working with the University of Texas at Arlington, USA on research of amphibians and reptiles in the

mountains of Java and Sumatra. Irvan has published on the herpetofauna of Kalimantan and his first book was

about snakes that are traded in Indonesia (CITES appendices I, II, and III) written in Indonesian. Currently,

Irvan is studying the phylogeography of the reed snake genera Calamaria for his Ph.D. at the University of

Brawijaya, Malang.

In accordance with the International Code of Zoological Nomenclature new rules and regulations (ICZN 2012), we have deposited this paper in publicly accessible institutional libraries.

The new species described herein has been registered in ZooBank (Polaszek 2005a, b), the official online registration system for the ICZN. The ZooBank publication LSID (Life Science

Identifier) for the new species described here can be viewed through any standard web browser by appending the LSID to the prefix “http://zoobank.org/”. The LSID for this publication

is:um:lsid:zoobank.org:pub:27336534-BAFC-40BE-84F7-43E0334596CD.

Separate print-only edition of paper(s) (reprint) are available upon request as a print-on-demand service. Please inquire by sending a request to: Amphibian & Reptile Conservation

(amphibian-reptile-conservation.org; arc.publisher@gmail.com).

Amphibian & Reptile Consen’ation is a Content Partner with the Encyclopedia of Life (EOL); http:///www.eol.org/ and submits information about new species to the EOL freely.

Digital archiving of this paper are found at the following institutions: ZenScientist (http://www.zenscientist.com/index.php/filedrawer); Ernst Mayr Library, Museum of Comparative Zool-

ogy, Harvard University, Cambridge, Massachusetts (USA); Florida Museum of Natural History, Gainesville, Florida (USA).

Complete journal archiving is found at: ZenScientist (http://www.zenscientist.com/index.php/filedrawer); Florida Museum of Natural History, Gainesville, Florida (USA).

Citations

ICZN. 2012. Amendment of Articles 8,9,10,21 and 78 of the International Code of Zoological Nomenclature to expand and refine methods of publication. Zootaxa 3450: 1-7.

Polaszek A et al. 2005a. Commentary: A universal register for animal names. Nature 437: All .

Polaszek A et al. 2005b. ZooBank: The open-access register for zoological taxonomy: Technical Discussion Paper. Bulletin of Zoological Nomenclature 62(4): 210-220.

September 201 4 | Volume 8 | Number 2 | e80

Amphib. Reptile Conserv. | amphibian-reptile-conservation.org (29)

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptiie Conservation

8(2) [General Section]: 30-35.

Short Communication

Field surveys in Western Panama indicate populations of

Atelopus varius frogs are persisting in regions where

Batrachochytrium dendrobatidis is now enzootic

^Rachel Perez, ^’^Corinne L. Richards-Zawacki, ^Alexander R. Krohn, ^Matthew Robak,

"^Edgardo J. Griffith, "^Heidi Ross, ^Brian Gratwicke, ^Roberto Ibanez, and ^ *Jamie Voyles

^Department of Biology, New Mexico Institute of Mining and Technology, Socorro, New Mexico 87801, USA ^Department of Ecology and

Evolutionary Biology, Tulane University, New Orleans, Louisiana 70118, USA ^Department of Environmental Science Policy and Management,

University of California, Berkeley, Berkeley, California 94720, USA '^El Valle Amphibian Conservation Center, El Valle, PANAMA ^Smithsonian

Conservation Biology Institute, Washington D.C., USA ^Smithsonian Tropical Research Institute, PO. Box 0843-03092, Balboa, Ancon, PANAMA

Key words. Chytridiomycosis, Central America, Harlequin frog, golden frog, threatened species, emerging infec-

tious disease, chytridiomycosis, Batrachochytrium dendrobatidis

Citation: Perez R, Richards-Zawacki CL, Krohn AR, Robak M, Griffith EJ, Ross H, Gratwicke B, Ibanez R, Voyles J. 2014. Field surveys in Western

Panama indicate populations of Atelopus varius frogs are persisting in regions where Batrachochytrium dendrobatidis is now enzootic. Amphibian &

Reptile Conservation 8(2) [General Section]: 30-35 (e85).

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: 18 June 2014; Accepted: 25 August 2014; Published: 11 November 2014

The stunning Harlequin frogs of the genus Atelopus,

once common, are now among the most imperiled of

all amphibian species (La Marca et al. 2005; Zippel et

al. 2006). Of 88 described Atelopus species in Central

and South America, 65 (74%) are Critically Endangered

(La Marca et al. 2005). The most pressing threat to these

frogs is ch54ridiomycosis, a fungal disease caused by Ba-

trachochytrium dendrobatidis (hereafter “5<i”) and as-

sociated with die-offs of amphibians around the world

(Berger et al. 1998; Longcore et al. 1999; Kilpatrick et al.

2010). In Western Panama, an epidemic wave spreading

from west to east caused mass mortality events, result-

ing in catastrophic losses in amphibian diversity (Lips

et al. 2006; Brem and Lips 2008; Woodhams et al. 2008;

Kilbum et al. 2010), including declines in t\wQQ Atelopus

species: A. varius, A. zeteki, and A. chiriquiensis.

Relatively few studies have been conducted on the

amphibian co mm unities of Western Panama following

ch54ridiomycosis outbreaks. Those few investigations

that have focused on understanding community compo-

sition where Bd is now enzootic report differential sur-

vival among host species (e.g., Brem and Lips 2008),

with some species putatively driven to local extinction

(Gagliardo et al. 2008). In particular, Atelopus species

were considered to be highly vulnerable to disease-in-

duced extinction for multiple reasons. First, A. varius

was used as an “indicator species” to monitor declines

and thus helped to document Bd invasion and character-

ize 5J-related amphibian losses (Brem and Lips 2008).

Second, A. zeteki has been repeatedly tested in controlled

laboratory infection experiments and found to be highly

susceptible to lethal ch)4ridiomycosis (Bustamente et al.

2010; DiRenzo et al. 2014; Ellison et al. 2014). Third,

recent immunogenetics research suggests that A. zeteki

adaptive immune responses are suppressed by Bd (El-

lison et al. 2014). Thus, Atelopus species have become

important focal species in the study of ch)4ridiomycosis

dynamics and have also provided motivation for progres-

sive conservation action (Gagliardo et al. 2008).

In October 2012, we revisited study sites that were

surveyed fox Atelopus from 2001 to 2004 (Richards and

Knowles 2007) and we established new study sites in

hopes of discovering extant populations of Critically

Endangered Atelopus species. Using a measuring tape,

a 200 m transect was developed and marked with flag-

ging tape every 10 m. For our surveys, 2-3 observers

walked these transects slowly, searching for amphibians

according to established visual encounter survey proto-

cols (e.g.. Lips 1999). We captured all post-metamorphic

amphibians we encountered using a fresh pair of gloves

or inverted plastic bag to minimi ze transmission or infec-

tion and followed strict field hygiene protocols (Phillot

et al. 2010). We noted the time and location of capture,

identified the species, sex and age class, and measured

Correspondence. Email: amie.voyles@gmail.com (Corresponding author, Jamie Voyles); tel: (720) 883-2341; fax: (208) 885-7905.

Amphib. Reptile Conserv.

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November 2014 | Volume 8 | Number 2 | e85

Perez et al.

snout-to-vent length and body mass. We also eollected

skin swab samples for all amphibians using standard-

ized swabbing techniques (Hyatt et al. 2007). We pre-

served the skin swab samples (by freezing at -20 °C) to

test for Bd infection using quantitative polymerase chain

reaction (qPCR; Boyle et al. 2005; Hyatt et al. 2007).

For the qPCR assay, we analyzed all samples in tripli-

cate with an internal positive control (Hyatt et al. 2007)

and used a dilution set of plasmid standards (obtained

from Pisces Molecular, Boulder, Colorado) to quantify

pathogen load. We converted plasmid copy numbers to

zoospore copy numbers using the line of best fit (r^ >

0.999) from a linear regression of log (plasmids) vs. log

(zoospores) (t4 = 210.6, P < 0.0001) that we obtained

by running the plasmid standard set alongside a series

of standards containing known quantities of zoospores

(obtained from Alex Hyatt, Australian Animal Health

Laboratory). If one of three replicate wells turned up

positive, we checked Cycle Threshold (Ct) value to de-

termine whether non-amplification in two of three wells

could have been caused by a low-level infection (near the

detection threshold) and verified that the qPCR was not

inhibited (IPC amplified normally). In cases of inhibition

or Ct values far from the detection threshold, we re-ran

and considered them positive if Bd was detected in any of

the three re-run wells.

We surveyed 16 field sites from 2012 to 2013, 10 of

which were sites where Atelopus species were found in

2004 (Richards and Knowles 2007), prior to the chy-

tridiomycosis epidemic. The remaining six sites were

chosen based on other biologists’ sightings of Atelopus

Figure 1. A female Harlequin frog, Atelopus varius. This spe-

cies, classified as Critically Endangered by lUCN, has been

found in small numbers in the mountains of Western Panama.

Figure 2. A pair of Atelopus varius in amplexus, found in the mountains of Western Panama.

Amphib. Reptile Conserv.

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November 2014 | Volume 8 | Number 2 | e85

Atelopus varius populations persisting after chytridiomycosis

Table 1. Summary of infection prevalence of Batrachochytrium dendrobatidis from amphibians found at three sites

where Atelopus varius still persist.

Site (Elevation)

Species

Prevalence

N

Lower 95%

Upper 95%

Highland 1 (735 m)

Atelopus varius

0%

0/1

0

0.975

Colostethus flotator

0%

0/1

0

0.975

Craugastor bransfordii

0%

0/1

0

0.975

Craugastor crassidigitus

0%

0/1

0

0.975

Craugastor fitzingeri

0%

0/1

0

0.975

Lithobates warszewitschii

60%

3/5

0.147

0.947

Pristimantis cerasinus

100%

1/1

0.025

1

Pristimantis cruentus

0%

0/2

0

0.842

Pristimantis museosus

0%

0/1

0

0.975

Pristimantis ridens

0%

0/1

0

0.975

Sachatamia albomaculata

25%

2/8

0.0715

0.591

Terahyla spinosa

0%

0/1

0

0.975

Glass frog metamorph

0%

0/1

0

0.975

Highland 2 (521 m)

Atelopus varius

0%

0/6

0

0.459

Colostethus flotator

0%

0/7

0

0.41

Colostethus panamensis

25%

1/4

0.073

0.524

Pristimantis ridens

0/1

0

0.975

Rhaebo haemititicus

42%

1/5

0.057

0.437

Silverstoneia flotator

33%

1/3

0.008

0.906

Smilisca spp.

100%

3/3

0.292

1

Lowland 1 (0 m)

Atelopus varius

0%

0/1

0

0.975

Craugastor bransfordii

0%

0/2

0

0.842

Craugastor crassidigitus

0%

0/1

0

0.975

Craugastor longirostris

0%

0/1

0

0.975

Craugastor fitzingeri

0%

0/2

0

0.842

Dendrobates auratus

0%

0/3

0

0.708

Dendrobates minnutus

0%

0/1

0

0.975

Diasperous spp.

0%

0/3

0

0.708

Pristimantis caryophyllaceus

0%

0/1

0

0.975

Rhinella alata

0%

0/10

0

0.308

Silverstoneia flotator

0%

0/1

0

0.975

(e.g., Hertz et al. 2012) or predicted habitat suitability

in species distribution models. We found persisting pop-

ulations of varius at three of 16 (18.7%) field sites

(Fig. 1, Table 1). At one site, we found one juvenile A.

varius, five adult males, and two adult females, including

one pair in amplexus (Fig. 2). We found individual adult

males at each of the other two respective sites (Table 1).

We have intentionally only provided general site infor-

mation, rather than precise site coordinates, due to the

risk of illegal animal collections.

We confirmed that Bd is present in two of these three

populations based on detection of Bd on skin swabs If om

other species (e.g., Lithobates warszewitschii and Sach-

atamia albomaculatd), but none of the Atelopus samples

were Bd positive on these transects (Table 1). These sites

vary in elevation from 45 to 750 m and all three are in

areas where fungal epidemics were associated with mas-

Amphib. Reptile Conserv.

sive amphibian declines from 2004 to 2006 (Lips et al.

2006; Brem and Lips 2008; Kilbum et al. 2010). We did

not detects, zeteki ox A. chiriquiensis at any of our study

sites. Although suiwey efforts for these species are still

underway, the absence of these species is concerning be-

cause we know that they previously had restricted ranges

(Zipple et al. 2006) and at least A. zeteki is known to be

highly susceptible to chytridiomycosis in laboratory in-

fection experiments (Bustamente et al. 2010; DiRenzo et

al. 2014; Ellison et al. 2014).

We found that A. varius is persisting in multiple sites

following a chytridiomycosis outbreak in western Pana-

ma. Furthermore, our positive qPCR results suggest that

these populations have survived despite the presence of

Bd. Prior to this study. Hertz et al. (2012) was the only

study to document sightings of A. varius in the wild in

Panama since 2006. Those observations were made in

32

November 2014 | Volume 8 | Number 2 | e85

Perez et al.

2009 at a site in Santa Fe National Park. These popula-

tions may be persisting for a wide range of biotic (e.g.,

changes in host behavioral, innate or acquired immune

responses, anti-5(i microbial communities) or abiotic

(e.g., environmental/thermal conditions) reasons. How-

ever, because there have been few coordinated efforts to

locate new populations, resurvey historical localities, or

test for Bd infections, the question of how these popu-

lations have persisted — and whether any other Atelopus

populations have survived — remains to be unraveled. We

believe that the lack of post-decline survey effort has not

been so much an oversight, but likely a consequence of

the enormity of the challenge of monitoring these spe-

cies while simultaneously establishing conservation pro-

grams to abate the threat of chytridiomycosis to entire

amphibian communities.

Post-decline surveys are critical for conservation of

Atelopus species, as well as for other neotropical am-

phibians. Documenting rediscovered species is critically

important for informing conservation and management

initiatives (Minteer et al. 2014) and, in this case, could

be accomplished with photographs, rather than collecting

the individuals. Moreover, understanding the variables

that permit some populations to persist while others die

out will be critical to conservation, especially since sev-

eral species are being bred in captivity (e.g., A. varius

and A. zeteki) with the expectation of one day return-

ing them to the wild (Gagliardo et al. 2008; Zippel et al.

2011). Our discovery of extant populations of A. varius

in 5(i-enzootic areas underscores the importance of con-

tinued monitoring for species presumed to be “extinct in

the wild,” even after long periods without any sightings.

Acknowledgments. — Disney Worldwide Wildlife

Fund, Association of Zoos and Aquariums (AZA) Con-

servation Endowment Fund, Riverbanks Conservation

Support Fund, Bay and Paul Foundations, Minnesota

Zoo Ulysses S. Seal Conservation Grant, Maryland Zoo

Conservation Fund, the Chicago Board of Trade Endan-

gered Species Fund, The AZA Amphibian Taxon Advi-

sory Group, Stone Center for Latin American Studies,

and Project Golden Frog funded this research. We thank

the Smithsonian Tropical Research Institute for allowing

access to their facilities. We thank Maggie Unkefer, Clif-

ford Richardson, Kelly Cruz and the Cruz family, Daniel

Medina, Karina Klonoski, and Simone Des Roches for

their support. This work was conducted under lACUC

NMT: 2013-1, lACUC Tulane: 0453, STRI lACUC:

2012-0901-2015, ANAM SE/AH-4-12, SE/AH-1-13,

SE/AH-1-12, and SE/AH-4-13.

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Rachel Perez is a M.S. student in the department of biology at New Mexieo Teeh. She reeeived her B.S.

at University of California, Riverside. Her main researeh interests are in the areas of epidemiology, eeoim-

munology, and amphibian eonservation. She is eurrently investigating innate defenses and mierohabitat

eonditions of neotropieal amphibians.

Corinne Richards-Zawacki is a professor in the Eeology and Evolutionary Biology department at Tulane

University. Her researeh lies at the interseetion of eeology and evolutionary biology in that she approaehes

questions about how ehanges in elimate and habitat shape population and eommunity proeesses in a way

that explieitly eonsiders their evolutionary implieations. The questions she asks address (1) the effeets

of elimate and landseape ehanges on speeies distributions and diversity, (2) how reproduetive isolation

evolves during speeiation, and (3) how elimate and host/pathogen evolution shape the dynamies of wild-

life diseases. She is passionate about amphibian eonservation and has 12 years of experienee working on

eonservation-oriented projeets in Panama. Mueh of her work in Panama has foeused on the Critieally En-

dangered Panamanian golden Ifogs. Her lab has also published studies related to the eaptive management

of amphibian speeies threatened by Bd.

Alexander Krohn is a eurrent Ph.D. eandidate at the University of California, Berkeley, in the department

of Environmental, Seienee, Poliey and Management. His dissertation foeuses on the eonvergent evolution

of melanism in desert reptiles, but he has been interested in herpetology, tropieal eeology, and eonservation

sinee he was in middle sehool.

Matthew Robak is broadly interested in amphibian eonservation. He is eurrently a Ph.D. eandidate at

Tulane University where he is researehing how differenees in temperature affeet amphibians’ immune

responses to Batrachochytrium dendrobatidis exposure.

Edgardo J. Griffith is a world-renowned leader in amphibian eonservation in Panama and one of the

founders of the El Valle Amphibian Conservation Center (EVACC). Edgardo is one of the pioneers that has

proven that multi-speeies ex situ amphibian eonservation is the only option we have at the moment to fight

the dramatie amphibian deelines in Panama. His work has been featured in several eonservation books,

doeumentaries, and peer-reviewed seientifie papers. Edgardo has 14 years working with the Panamanian

amphibians, ineluding the Panamanian golden frog in situ and breeding them ex situ.

Heidi Ross is the projeet direetor at the El Valle Amphibian Conservation Center (EVACC). Heidi and

her husband were awarded the San Diego Medal of International Conservation in 2012 for their work at

EVACC. Heidi has 10 years of experienee ex situ experienee with Panamanian amphibians leading to the

breeding of nine of the 12 priority speeies in the EVACC eolleetion, ineluding the Panamanian golden frog.

Amphib. Reptile Conserv.

34

November 2014 | Volume 8 | Number 2 | e85

Perez et al.

Brian Gratwicke is a conservation biologist that leads the amphibian eonservation program at the Nation-

al Zoo (Washington, DC, USA). Brian also leads the Panama Amphibian Reseue and Conservation Projeet.

Previous eonservation experienee ineludes work on freshwater eeology in Afriea, tropieal marine eeology

in the Caribbean, and tiger eonservation efforts in Asia. He has published more than 25 peer-reviewed

papers and book ehapters and was a eontributor to Hotspots Revisited. Brian reeeived a Ph.D. in zoology

from Oxford University. He obtained his baehelor’s and master’s degrees in zoology and fisheries eeology,

respeetively, from the University of Zimbabwe.

Roberto Ibanez is the in-eountry Direetor of the Panama Amphibian Reseue and Conservation Projeet,

based at the Smithsonian Tropieal Researeh Institute. He has been part-time Assoeiate Professor at the Uni-

versidad de Panama (1996—2014), and professor during de Panama Field Study Semester of MeGill Uni-

versity (2003—2013). He reeeived his B.S. in Biology from the Universidad de Panama, and his Master’s

degree and Ph.D. in Zoology from the Eeology and Evolutionary Biology Department at the University of

Conneetieut. He has studied the amphibians and reptiles of Panama for more than 30 years. He has pub-

lished more than 30 peer-reviewed papers, more than 20 notes, and a book guide to the amphibians of the

lowlands of eentral Panama. He is a distinguished researeher of the national researeh system of Panama.

Jamie Voyles is eurrently an Assistant Professor at New Mexieo Teeh working on emerging infeetious

diseases in wildlife. She eonduets eh54ridiomyeosis researeh in Central Ameriea and in California. She

is a member of multiple working groups investigating disease-related amphibian deelines. She is aetively

involved in eonservation initiatives, sueh as Amphibian Reseue and Conservation Projeet, and eontributes

to amphibianreseue.org and AmphibiaWeb.

Amphib. Reptile Conserv.

35

November 2014 | Volume 8 | Number 2 | e85

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptiie Conservation

8(2) [General Section]: 36-43.

Confirmation of introduced Louisiana pinesnakes, Pituophis

ruthveni, in Fiorida based on molecular analyses

^’*Kenneth L. Krysko, ^Dustin C. Smith, ^Michael R. Rochford, "^Guy N. Kieckhefer III,

and ^’^Leroy P. Nunez

^Florida Museum of Natural History, Division of Herpetology, 1659 Museum Road, University of Florida, Gainesville, Florida 32611, USA ^Zoo

Miami, 12400 Southwest 152nd Street, Miami, Florida 33177, USA ^University of Florida, Fort Lauderdale Research and Education Center, 3205

College Avenue, Fort Lauderdale, Florida 33314-7719, USA ‘^1113 Wyoming Street, Belgrade, Montana 59714, USA

Abstract — As more wide-range phylogenetic studies are available, the opportunity arises to

compare DNAfrom these data sets to suspected introduced individuals in order to confirm species

identification and determine their geographic origins. Two recently collected Pituophis specimens

in Miami-Dade County, Florida, were examined using molecular analyses. Maximum likelihood

and Bayesian inference methods place our specimens within the P. catenifer sayi I P ruthveni

clade. Additional morphological evidence support their identification as the Louisiana pinesnake,

Pituophis ruthveni StuW 1929, a species indigenous to a small area in western Louisiana and eastern

Texas and candidate for listing by the U.S. Fish and Wildlife Service. Although P ruthveni is viewed

as a distinct species from P catenifer sayi based on allopatry and differences in color pattern, no

molecular evidence was found supporting the recognition of P ruthveni as a separate species.

However, adding other mtDNA and nuclear DNA genes might provide needed data for distinguishing

between these two named taxa.

Key words. DNA, exotics, ND4, mitochondrial, mtDNA, nonnative, phylogenetics, Squamata, species

Citation: Krysko KL, Smith DC, Rochford MR, Kieckhefer III GN, Nunez LR 2014. Confirmation of introduced Louisiana pinesnakes, Pituophis ruth-

veni, in Florida based on molecular analyses. Amphibian & Reptile Conservation 8(2) [General Section]: 36-43 (e86).

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: 19 August 2014; Accepted: 27 October 2014; Published: 19 November 2014

Introduction

Introduced species (e.g., stages 2-5 after Colautti and

Macisaae 2004) are those transferred from their native

range into a new nonindigenous area. Over the past cen-

tury it has become increasingly clear how disruptive

human-eaused biological introductions have been to the

planet. While not all introduced species eause obvious

harm, some introdueed speeies can eventually become

eeonomic threats and lead to serious conservation prob-

lems (Simberloff et al. 1997). As of 2005, it was estimat-

ed that the eost of environmental damages, losses, and

control due to introdueed speeies exeeeded $120 billion

per year in the United States alone (Pimentel et al. 2005).

Prior to 2011, the state of Florida had 137 doeumented

introdueed reptile and amphibian taxa (56 being estab-

lished), which ranks highest in the world (Krysko et al.

2011a, 2012). Invasion pathways in Florida include (few-

est to highest numbers) biological control, zoos, cargo/

plant shipments, and the pet trade.

Pinesnakes, bullsnakes, and gophersnakes {Pituophis

Holbrook 1842) are large (up to 254 em total length)

eonstrictors native to North Ameriea, charaeterized by

disproportionately small heads, four prefrontal seales,

and a large rostral plate that extends upwards between

the intemasals (Conant and Collins 1991). Based primar-

ily on moleeular data using Parsimony and Maximum

Likelihood analyses with 893 base pairs (bp) of the nieo-

tinamide adenine dinucleotide dehydrogenase subunit 4

(ND4) region (Rodriguez-Robles and De Jesus-Eseobar

2000), the P. melanoleucus species complex contains

three eurrently recognized species; P. melanoleucus

(sensu strieto; Pinesnakes; with three subspecies P. m.

lodingi, P m. melanoleucus, P m. mugitus), P cateni-

fer (gophersnakes and bullsnakes; with six subspecies P.

c. affinis, P c. annectens, P c. catenifer, P c. deserti-

Correspondence. Email: * kenneyk@ufl.edu (Corresponding author, KLK); ^dustsmi@miamidade.gov; ^ mikerochford@hotmail.com',

^gnkieck@gmail. com ; fherizinosaur@yahoo. com

Amphib. Reptile Conserv.

36

November 2014 | Volume 8 | Number 2 | e86

Krysko et al.

Figure 1. Map of Zoo Miami bounded in green. Note that major roadways, residential areas, and undeveloped proteeted lands

surround zoo property. Dots represents loeations of Pituophis found on zoo property; yellow = UF-Herpetology 157954 (gravid

female) and red = UF-Herpetology 163092 (adult male).

cola, P c. pumilis, P. c. sayi), and P. ruthveni (Louisi-

ana pinesnake). Pituophis melanoleucus (Daudin 1803)

occurs in the eastern United States from southern New

Jersey south to extreme southern peninsular Florida (i.e.,

Miami-Dade County; Krysko et al. 2011b) and west

to Kentueky and southeastern Louisiana (Rodriguez-

Robles and De Jesus-Eseobar 2000). This species lacks

a dark line from the eye to the angle of the jaw, has a

dorsal pattern either absent (uniform blaek), obseure, or

whitish to brownish with 23-30 distinct dark dorsal body

blotches that are elearly separated from each other both

anteriorly and posteriorly along the body and tail (Knight

1986; Powell et al. 1998; Reiehling 1995; Thomas et al.

1976). Pituophis catenifer oceurs from the Paeifie Ocean

east to Wiseonsin, Illinois, and Texas, and from Canada

south to Mexieo (Rodriguez-Robles and De Jesus-Eseo-

bar 2000; Powell et al. 1998). This speeies typically has

a dark line from the eye to the angle of the jaw, and a yel-

low or cream-eolored dorsal pattern with 41-79 distinet

dark dorsal blotehes that are clearly separated from eaeh

other both anteriorly and posteriorly along the body and

tail (Knight 1986; Powell et al. 1998; Reiehling 1995;

Thomas et al. 1976). Pituophis ruthveni occurs in allo-

patric populations in western-central Louisiana to eastern

Texas (Ealy et al. 2004; Powell et al. 1998). This species

sometimes laeks a dark line from the eye to the angle of

the jaw, and has a pale brown dorsal pattern with 28-38

dark dorsal blotches; near the head the blotches obscure

the ground coloration, whereas near the tail they are dis-

tinctly separated from eaeh other (Knight 1986; Pow-

ell et al. 1998; Reiehling 1995; Stull 1929; Thomas et

al. 1976). Although P. ruthveni is nested within a elade

eontaining only P. c. sayi, it is recognized as a separate

species because it oceurs in ahopatric populations and

is somewhat diagnosable using eolor pattern charaeters

(Collins 1991; Knight 1986; Reiehling 1995; Rodriguez-

Robles and De Jesus-Escobar 2000; Thomas et al. 1976).

Pituophis ruthveni is also a eandidate for listing as an

imperiled species by the U.S. Fish and Wildlife Serviee

(2013).

The last known Pituophis melanoleucus from ex-

treme southern peninsular Florida (UF-Herpetology

45970) was eohected in 1980 in a disturbed pineland

(with Casuarina and Schinus) in Cutler Ridge, Miami-

Dade County, and because of ongoing dense urbaniza-

tion this speeies is believed to be extirpated along the

Atlantie Coast Ridge (Krysko et al. 2011b). In 2010, two

Pituophis were eohected on the Atlantie Coast Ridge at

Zoo Miami, Miami-Dade County; one was found in an

undeveloped area and another near publie aeeess. Based

on color pattern alone, these snakes were suspected to be

introduced P. ruthveni and reported to represent the first

known vouehers for this speeies in Florida (Krysko et al.

2011a). Many doeumented introduetions categorize spe-

eies based on sometimes vague superficial morphology,

such as color patterns, which may or may not be arbitrary

human constructs. However, as more wide-range phylo-

genetie studies are conducted and published, the oppor-

tunity arises for other researchers to eompare DNA from

known data sets to suspected introdueed individuals in

Amphib. Reptile Conserv.

37

November 2014 | Volume 8 | Number 2 | e86

Verification of introduced Pituophis ruthveni in Florida

Figure 2. Well-dQYQlopQd Pituophis embryo (UF-Herpetology 164295) oviposited from wild colleeted gravid female (UF-Herpe-

tology 157954) in Miami, Miami-Dade County, Florida.

order to confirm species identification as well as deter-

mine their geographic origins. In this paper, we conduct

molecular analyses of Pituophis in a coalescent frame-

work to confirm species identity and phylogenetic place-

ment of our two specimens, followed by more detailed

examination of morphology and color pattern.

Material and Methods

Site description and specimen acquisition

Zoo Miami is situated at 12400 SW 152* Street, Miami,

Miami-Dade County, Florida, USA (Fig. 1 ; 25.61 1926°N,

80.398042°W, Datum WGS84, elev. 2 m). The property

consists of ca. 300 ha, 106 ha of which are undeveloped

managed lands, predominantly of pine rockland habitat.

Zoo Miami property is surrounded by a mixture of natu-

ral areas, disturbed areas, and a county park, followed by

dense urbanization.

On 16 May 2010 at 1645 h, an didult Pituophis (gravid

female, 1,302 mm SVL, 1,486 mm TL; UF-Herpetolo-

gy 157954; see Fig. 86 in Krysko et al. 2011) was col-

lected in a service area behind a large animal exhibit

(25.60395°N, 80.4006°W). This snake was observed

by zoo staff the previous day along an adjacent public

walkway, but was not captured. This snake was retained

in captivity and oviposited three eggs on 22 June 2010.

The eggs were viewed with a light on 28 June 2010; all

three eggs contained a dark blood spot, but only one egg

had an obvious network of veins developing. The first

two eggs failed to develop and were discarded on 06 July

2010. The third egg had an unpleasant odor and was fro-

zen on 14 September 2010; it was dissected on 20 Sep-

tember 2010 and revealed a well-developed embryo (UF-

Herpetology 164295; Fig. 2).

On 25 December 2010 at 1215 h, another adult (male,

1,425 mm SVL, 1,635 mm TL) Pituophis (UF-Herpe-

tology 163092) was collected in an undeveloped area

(25.60304°N, 80.40295°W), across a large man-made

lake and 0.26 km southwest of the first snake.

The well-developed embryo, shed skins from the two

adults, and digital images were deposited in the Divi-

sion of Herpetology, Florida Museum of Natural His-

tory, University of Florida. The female (UF-Herpetology

157954) is currently housed at the Memphis Zoo, and the

male (UF-Herpetology 163092) is housed at Zoo Miami.

Laboratory techniques

DNA isolations were obtained using QIAquick PCR Pu-

rification Kit and DNeasy Blood and Tissue Kit (Qiagen

Sciences, LLC). Using total cellular DNA as a template

and Polymerase Chain Reaction (PCR) methodology

(Saiki et al. 1988), mitochondrial DNA (mtDNA) was

amplified and sequenced for the ND4 region follow-

ing Rodriguez-Robles and De Jesus-Escobar (2000).

The ND4 region includes a section of the 3 ’ end of the

ND4 gene, and two subsequent transfer ribonucleic ac-

ids (tRNA”‘®, tRNA^®^), which were sequenced using the

primers ND4 and Leu (Arevalo et al. 1994). PCR was

conducted in 25 pi reactions: 9.5 pi H^O, 12.5 pi GoTaq®

Master Mix (Promega Corp, Madison, Wisconsin, USA),

1.0 pi each primer (10 pM), and 1.0 pi DNA template.

PCR parameters included initial denaturing at 94 °C for

three min, followed by 35 cycles of amplification: de-

Amphib. Reptile Conserv.

38

November 2014 | Volume 8 | Number 2 | e86

Krysko et al.

98

93

CA_SanBenito_Lamprope/f/5 9efi//a_HWG1485

TX_Culberson_Bogerfop/?/s suboct//ans_CME116

CA_Riverside_Anzo^s elegans_W^Z^

■ OHOttawaPanfPerophis U'U/p/>JUS_CAS1 84362

100

94

— FL_Hi llsborough_P/fiiop/7/s melanoletJCLis mugitus_W\yZ^ 3063

- FL^Waku na^Pituophis melanoleucus mugitusJJSN M2 1 1 452

\^C_Brur\5'Mck_Pituophis melanoleucus mela^oleucus_W\VZ^502^9

•“I NJ_Cumberland_P/fuop/i/s meianoleucus melanoieucus_MVZ225b20

- NJ_Burlington_P/f/;op/7/s meianoleucus melanoleucus_W\WZ22b52)

I — Pituophis meianoleucus lod^nglJH\NG2Q5^

L Pituophis meianoleucus lodlngl_HV\lG2652

Pituophis deppei jani

q:

100

Mexico_Michoacan_P/^yop/7is deppei deppei_2

Mexico_Durango_P/fL/op/i/s deppei b'eppe/_1

.^0

100

■ Mexico_BajaCal_P;Yuop^/5 vertebralisJSB

Wiex\co_Ba\aCa\_Pituophis vertebra!is_JAR7S

— Mexico_BajaCal_P/fuopYj/s catenifer bimans_^ 1

97r

93

97i

C

100

Pituophis vertebralis_40

- AZ_Oooh\se_Pituophis cafe/?/feraffi;?/s_MVZ1 37697

■ AZ_Maricopa_P/fi/opft/s catemfer affln/s_MVZ162369

- CO_Garfield_P/f£/opft/s catenifer deserf/co/a_MVZ150216

NM_Luna_P/fyop/?/5 catenifer 51 1

TX_JeffDavis_P/fmp/?/s catenifer sayi^MVZ 150218

MO^SaintLouis^P/Yuoph/s cafefT/fersayi^MVZ226247

£

cc

o

LA_Bienville_P/f£iop/7/s ruthveni_32

— OK_Cleveland_P/ftiop/7/s catenifer sayi^37

*- LA_Bienville_P/fL/opYi/s ruthvenl_33

— LA_Bienville_P/fi/opY?/s lvthveniJ3^

— CO_Jefferson_P»ft/opft/s cafef?rfe/'say/_35

74

FL_MiamiDade_UF1 57954

FL MiamiDade UF163092

100r

99

C

CD

£

2

a;

‘5;

CO

ti

a;

- CA_Napa_Pifuopft/s catenifer catenifer

- CA_Alameda_Pifuopft(s catenifer catenifer_JAR77

100p CA_Mendocino_P/fuopftis catenifer catenifer_CAS20A258

' N V^Mineral Pjfuoppjs catenifer deserticolaJtJNZt 37577

v3>

5

CD

O

CD

<2 d;

2

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100 r

■ Gualemala_P/?t/op/7/s//r?eaf/coWs g/jbson/_CJF1500

Guatemala^Prfuop/iis lineaticollis g/ijso/7/^CJF1501

98r

99

97

— UT_Ulah_P/fi;op/7/s catenifer deserticoia

CA Kern P/fyopWs catenifer deserficola_RSR 115

Mexico_BajaCal_P/ft/op/7fs catenifer b;mar/s_10

- CA_Monterey_Prtuopb/s catenifer catenifer_JAR75

Mexico_BajaCal_P/ft/op/7/s cafeY7/ferft;//g'//7afi/s

Pituophis catenifer bimaris_‘]3

— W\ex\co_BajaOa\_Pituophis cafenife/'d/Vr>ar/5_12

— CA_SanDiego_P/ft/oppys catenifer annectens_M^/Z^ 50206

84l- CA_SanDiego_P/fL/op/7/s catenifer annecteas_MVZ149983

CASantaBarbaraPyfyop/r/s catenifer pumilus_29

CA_SantaBarbara_P/fi;op/7/s catenifer pumllus_30

88

.CO

"o

.O

03

.g

q:

0.02

Figure 3. Maximum Likelihood phylogeny for Pituophis (Squamata: Colubridae) snakes, ineluding the two known P. ruthveni

(highlighted in yellow, UF-Herpetology 157954 and 163092) eolleeted in Miami, Miami-Dade County, Florida. Note that values (>

50%) above nodes represent bootstrap support. Inset photograph of UF-Herpetology 157954 by Dustin C. Smith.

naturing at 94 °C for one min, annealing at 52 °C for

one min, and extension at 72 °C for one min, followed

by a final extension at 72 °C for seven min . Three pi of

each PCR product were electrophoresed on a 1% agarose

gel, visualized with GelRed™ staining (Biotium Inc.,

Hayward, California, USA), and compared with a DNA

standard. Sequence files from the automated sequencer

(Genomics Division, Interdisciplinary Center for Bio-

technology Research, University of Florida) were as-

sembled and edited as necessary with Geneious software

(ver. 6.1, created by Biomatters. Available from http://

www.geneious.com).

Phylogenetic analyses. — DNA sequence data were

downloaded from GenBank for 46 snakes, including 42

Pituophis, and one of each Lampropeltis getula, Panthe-

rophis vulpinus, Bogertophis subocularis, and Arizona

elegans incorporating the original data set from Rodri-

guez-Robles and De Jesus-Escobar (2000) and current

taxonomy after Pyron and Burbrink (2009). GenBank

Accession numbers for our two Pituophis specimens

(UF-Herpetology 157954 and 163092) are KJ938643

and KJ938644, respectively.

A total of 48 specimens with 875 base pairs (bp) of se-

quence data were analyzed. Relationships among mtDNA

haplotypes were estimated using both Maximum Likeli-

hood (ML) and Bayesian Inference (BI) methods. ML

was conducted with the General Time Reversible model

with gamma distributed rate heterogeneity (GTR + F)

and 1,000 nonparametric bootstrap replicates (Felsen-

stein 2004) to assess node support using RAxML-HPC

BlackBox (Stamatakis 2006; Stamatakis et al. 2008)

from the CIPRES Science Gateway (Miller et al. 2010).

BI was conducted using BEAST (ver. 1.8; Drummond

and Rambaut 2007) from the UF-HPC Galaxy instance

(http://hpc.ufl.edu; Blankenberg et al. 2010; Giardine et

al. 2005; Goecks et al. 2010). The Bayesian Information

Criterion in jModelTest (ver. 2.1.4; Darriba et al. 2012;

Guindon and Gascuel 2003) determined the best-fit nu-

cleotide substitution model to be Hasegawa, Kishino,

and Yano with a proportion of invariant sites and gamma

distributed rate heterogeneity (HKY + I + T). A relaxed

Amphib. Reptile Conserv.

39

November 2014 | Volume 8 | Number 2 | e86

Verification of introduced Pituophis ruthveni in Florida

99

99|

1

99 1

CA_SanBeii1fl_LaffipiTope!fe9eliy/a_HWG1485

CARiverside _Artzofiseiega;is_HVZ137665

TX_Culberson_Sogdop/i/s sui5«ci/)0fi5_CMEI 1 6

0N_0tl3wa_PaafteropWs Ki/lpiii[;s_CAS184362

Pifyop/jiscfeppe/^ani

Meaco_MichDacan_fl'(i/opljis [teppef rfeppa_2

Meaco_Durarigo_filuflp/?(s deppei cteppe/_1

GualeTOla_P('t[j<}pfi's//nea!('M!fe giiMf]f_CJF1501 1

Guale™ia_P('!uopfe /meatofe g/iso/jf_CJF1500 1

P. deppei

P lineat/collis

FLHillsboroi^hPjfijtipte me/araletos rm;^(us_MVZFC13063

FL_Wakulla_R'!uoph(s melano'ei/ws ff!!/jifws_USNM21 1452

NC_Bruisv.'icli_P/fi;w/i/s m. meiar:!>lwMS_MVZ150219

NJ_C jmbeHand_Pft:;!)p/jis m. »7;e/a/!o/ei/cus_MVZ225520

NJ_Burlinglon_Pituophis_rr_rrelanoleu(!us_MVZ225521

Pifiypp/:is ;ne/a™/e(;c!;s /t>dingf_HWG2651

P^uophk mknolsucus Wng[_HIWG2652

CA_Alameda_Pwwp/;is ca/en/fe; cafefflfer_JAR77

CA_Napa_PtftJopPis cafeiiife^ cafemfer

CA_MerdocinD_Pm.^fe cateiffer cafenffer_CAS20l258

NV Mineral PifuopA/s Mte/i;fefdeseriia)Ja_MVZ137577

CAKemPdmphis catew/fef dssert'coteRSR1 15

UT_Ulah_P[i'iropft*s' caimkr desejl'ico/a

Mexico_BajaCal_Rli;!)p/)(5 calmer lymans_IO

CA_Monterey_Pi7t/(3pfjJS cafe/?ter cafenffe,''_JAR75

Mexico_BajaCal_fl'lrjt)pn(s rafenifer Mgmfus

Mexico_BajaCal_RK;op/)(5 cafepder lxmans_l2

Prfuop/iis calender toa;;s_1 3

CA_SanDiego_Pflw>pfiis ralenife'aifleciefJs_HVZ15CI206

CA_SanDifigo_Pf!uophis t;atenite'3iinecitefls_HVZ1499B3

CA_SantaBaitara_Pfltjopft,ficafeni/efpOT/ius_2&

CA_SantaBaitara_Ptopft,isiafe.ri//efpum/lusJ

MeBCO_BajaCal_PfW&pP(S i«/telrelis_38

Me)dco_BajaCal_fl'l[jt)pP(s rafemfer !xmans_1 1

Meaco_BajaCal Rlijoprts i«rteljraI('s_JAR78

Prfvop/iisverfebra/IsJO

AZ_Mana)pa_P/f(/(!pf]ys :a!e;!ffer affiffl5_WVZ1 62369

AZ_Cochise_P/!(/opf?ls calaiifer 3]^nis_MVZ137697

CO_6arfield_PJ!'i/op/;fsc3teajferdese.^'coia_UVZ150216

TX_Js1fDavis_P/fuopf}rsc3te,r!/fersayi_MVZ15D218

NM_Ljna_Pikipi)fs ca?ei'w'fera;i?f]fs_H8S151 1

MO_SaintLouis_P/luoptecaIer:/'fersay(_MVZ226247

LA,_Bienville_Pt[/opft/s futhveniJ2

OK_C!evelard_P(‘f[/op^fs catepifersayi_37

LA,_8ienwille_PfT[/opf]/'s

LA,_Bienwille_PrT[/opfi/s rutoiJS

CO_Jefferson_R1[jopb(sca!a,7ifer53ji_35

FL_MmiDade_UF163092

FL MiamiOade UF157954

P- catenifer

c.

-c

s

a;

I'

to

6

a;

Figure 4. Bayesian Inference phylogeny for Pituophis (Squamata: Colubridae) snakes, including the two known P. ruthveni (high-

lighted in yellow, UF-Herpetology 157954 and 163092) collected in Miami, Miami-Dade County, Florida. Note that values (> 95%)

above nodes represent posterior probabilities. Inset photograph of UF-Herpetology 163092 by Dustin C. Smith.

phylogenetics method was used without relying on a

potentially arbitrary molecular clock (Zuckerkandl and

Pauling 1965) that might incorporate uncertainty in the

tree estimation process (Drummond et al. 2006). An un-

correlated lognormal relaxed clock with coalescent con-

stant population size (Kingman 1982), estimated base

frequencies, randomly generated starting tree, and nor-

mal distribution for the ucld.mean parameter priors were

used. Two independent runs were performed consisting

of three heated and one cold Markov Chain Monte Carlo

(MCMC) estimated for 40 million generations, with ev-

ery 1,000th sample being retained. Both MCMC runs

were analyzed independently (to confirm chains were

converging and not sampling local optima) using Tracer

(ver. 1.6) for ESS values >200, as well as for a split stan-

dard deviation less than 0.005 for -InL tree values among

chains that indicate parameter stationarity was achieved.

Trees sampled prior to stationarity were discarded as

bum-in, which occurred prior to five million generations.

Trees from both independent MCMC mns were com-

bined and burn-in was removed using LogCombiner (ver.

1.8), the best statistically supported tree (i.e.. Maximum

clade credibility tree) with mean heights was obtained

using TreeAnnotator (ver. 1.8), and a phylogenetic hy-

pothesis with posterior probabilities was created using

FigTree (ver. 1.4).

The most credible inferences of phylogenetic relation-

ships were confined to nodes where nonparametric boot-

strap values > 70% and posterior probability (Pp) was >

95% (Hillis and Bull 1993; Felsenstein 2004).

Morphology and color pattern

We determined sex, snout-vent length (SVF), tail length,

number of ventrals, subcaudals, supralabials, infralabi-

als, preoculars, postoculars, temporals, loreals, and dor-

sal scale rows; and color pattern of dorsum and venter.

We compared these data to those found in the literature.

Results

Phylogenetic analyses. — Both MF and BI methods

produced identical phylogenetic groupings (Figs. 3 and

4). Although some of these clades are organized differ-

ently in relation to one another the monophyly of Pituo-

phis is well supported, which is congment with the find-

ings by Pyron and Burbrink (2009), though the latter

study used only single samples for each species. Both of

Amphib. Reptile Conserv.

40

November 2014 | Volume 8 | Number 2 | e86

Krysko et al.

our two Pituophis specimens have the same mtDNAhap-

lotype, and both phylogenetic methods place them within

the P. catenifer sayi / P. ruthveni clade.

Morphological data for UF -Herpetology 157954 in-

clude 226 ventrals, 55 subcaudals, 8/8 (left/right) supral-

abials, 11/11 infiralabials, 1/1 preoculars, 7/7 postoculars,

4 temporals, 1/1 loreals, 27-30-24 dorsal scale rows,

34 body blotches, 8 tail blotches, parietal stripe present,

and heavily patterned venter. Data for UF-Herpetology

163092 include 212 ventrals, 57 subcaudals, 8/8 (left/

right) supralabials, 11/11 infralabials, 1/1 preoculars, 7/7

postoculars, 4 temporals, 1/1 loreals, 27-31-23 dorsal

scale rows, 32 body blotches, and 11 tail blotches.

Discussion

Our ML and BI phylogenies produced identical mam

phylogenetic groupings (Figs. 3 and 4) as those found

in the ML analysis by Rodriguez-Robles and De Jesus-

Escobar (2000). However, we found no support for some

relationships, and no support values are provided on the

original ML tree by Rodriguez-Robles and De Jesus-

Escobar (2000). Our two Pituophis specimens were

placed within a well-supported P. catenifer sayi / P. ruth-

veni clade, the same group of specimens (except for our

Florida specimens) uncovered by Rodriguez-Robles and

De Jesus-Escobar (2000). Pituophis catenifer sayi and

P ruthveni were also found to be sister taxa based on

a combined mtDNA and single nuclear (nDNA) (Pyron

and Burbrink 2009) and phenetic moiphological similar-

ity (Reichling 1995) analyses. Nonetheless, we found

no molecular support for the recognition of P ruthveni

as a separate species. One of the limitations of our and

Rodriguez-Robles and De Jesus-Escobar ’s (2000) mo-

lecular studies is the use of only a single locus (ND4 re-

gion), and adding additional mtDNA and unlinked nDNA

genes might provide needed data for distinguishing be-

tween these two named taxa. Pituophis ruthveni is cur-

rently recognized as a separate species because it occurs

in allopatric populations and is believed to be diagnos-

able using color pattern characters, the most diagnostic

being 28-38 dark dorsal body blotches and the blotches

obscuring the ground coloration anteriorly (Collins 1991 ;

Reichling 1995; Rodriguez-Robles and De Jesus-Esco-

bar 2000). Our two Pituophis specimens exhibit these

three characters, thus we categorized them as P. ruthveni.

Before our specimens were found, Pituophis ruthveni

was not known to be kept at Zoo Miami, therefore this

species is not representative of a zoo-mediated introduc-

tion pathway and was likely released by an outside per-

son. Other species such as the Reticulated python, Ma-

layopython reticulatus (see Kaiser et al. 2013; Reynolds

et al. 2014), and Pacific Coast giant musk turtle, Stauro-

typus salvinii, are other examples of reptile species that

have been illegally released on zoo property, the latter

possibly established (Smith et al. 2011). Although we are

currently uncertain if P ruthveni is established in the vast

protected undeveloped habitats surrounding public ac-

cess areas, an adult male and gravid female were found

suggesting reproduction might have taken place in the

wild.

Acknowledgments . — We are thankful to everyone

who assisted in this study: Ryan Zach, Kevin Kopf, and

Oscar Rodriguez for collecting specimens; Steve Reich-

ling for providing data; David L. Reed and Gustav Pau-

lay for laboratory space; and R. Alexander Pyron and

Craig Hassapakis for reviewing this paper.

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Kenneth L. Krysko is a senior herpetologist at the Florida Museum of Natural History, University of

Florida (UF), where he has helped eurate the international researeh and teaehing colleetions for more than

15 years. He studies ecology and molecular systematics on both native and normative herps and produced

Florida’s first Atlas of amphibians and reptiles. His research on introduced species has made UF-Herpe-

tology the information center for invasive species in Florida. He recently published a large monograph

documenting species introductions as far back as 1863 and illustrated that Florida has the worst invasive

reptile problem in the entire world. He received his B.S. in Biological Sciences from Florida State Univer-

sity, M.S. in Biological Sciences from Florida International University, and Ph.D. in Wildlife Ecology and

Conservation from the University of Florida.

Dustin C. Smith is a conservation and research specialist at Zoo Miami. He graduated with a B.A. in 2001

from the University of South Florida. He has been involved with research projects in Costa Rica, Panama,

and throughout the southeastern US, and is currently working on projects in Puerto Rico and Belize. Most

of his current research involves native and non-native amphibians and reptiles in southern Florida.

Michael R. Rochford is the Invasive Species Coordinator at the University of Florida’s Fort Lauderdale

Research and Education Center, where he has worked for the last eight years. He received his B.S. in Biol-

ogy from Kansas State University in 2004. He has worked extensively on radio-telemetry and diet studies

of Burmese pythons in Elorida as well as on projects to assess populations of American alligators and

American crocodiles. In addition to his research and conservation interests, he also has strong interest in

finding and photographing herpetofauna of the United States and Mexico.

Trey “Guy” Kieckhefer graduated with a B.S. in Biology from Eckerd College in Saint Petersburg, Elor-

ida. He worked for the University of Elorida studying invasive Burmese pythons in the Everglades, and

currently works at the Tavernier Science Center in the Elorida Keys for the National Audubon Society. He

conducts surveys for prey fishes in the Everglades and submerged aquatic vegetation. He is interested in

Ichthyology, Ornithology, exotic species, and the overall health of the Everglades ecosystem.

a Leroy P. Nunez is a collections assistant within the Division of Herpetology at the Elorida Museum of

Natural History, University of Elorida (UE). He obtained an Associate’s Degree in Zoology from Santa Ee

College and a Bachelor’s Degree in Biology from UE. He is currently working on his Master’s Degree in

Interdisciplinary Ecology at the School of Natural Resources and Environment at UF. His current research

involves studying phytogenies of invasive squamates in Florida and his future research projects will focus

on phylogenetics and developmental evolutionary biology.

Amphib. Reptile Conserv.

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November 2014 I Volume 8 I Number 2 I e86

CONTENTS

Special Section

Alejandro Valencia-Zuleta, Andres Felipe Jaramillo-Martinez, Andrea Echeverry-Bocanegra, Ron-

ald Viafara-Vega, Oscar Hernandez- Cordoba, Victoria E. Cardona-Botero, Jaime Gutierrez-

ZuNiGA, AND Fernando Castro -Herrera — Conservation status of the herpetofauna, proteeted areas, and

eurrent problems in Valle del Cauea, Colombia 1

Alejandro Valencia-Zuleta et al. — Supplemental Material SI

General Section

Gernot Vogel, Patrick David, and Irvan Sidik — On Trimeresurus sumatranus (Raffles, 1822), with the des-

ignation of a neotype and the description of a new species of pitviper from Sumatra (Squamata: Viperidae:

Crotalinae) 1

Rachel Perez, Corinne L. Richards-Zawacki, Alexander R. Krohn, Matthew Robak, Edgardo J.

Griffith, Heidi Ross, Brian Gratwicke, Roberto Ibanez, and Jamie Voyles — Field surveys in Western

Panama indicate populations of Atelopus varius frogs are persisting in regions where Batrachochytrium

dendrobatidis is now enzootic 30

Kenneth L. Krysko, Dustin C. Smith, Michael R. Rochford, Guy N. Kieckhefer III, and Leroy P. Nu-

nez — Confirmation of introduced Louisiana pinesnakes, Pituophis ruthveni, in Florida based on molecular

analyses 36

Table of Contents Back cover

Cover: Bothrocophias colombianus, by Jorge Alberto Zuniga Baos and Luis Enrique Vera Perez; Allobates juanii, by

Alvaro Velasquez; Bolitoglossa savagei and Podocnemis lewyana, by Juan David Jimenez Bolano; Caecilia thomp-

soni, by Esteban Alzate; Anolis macrolepis, by Fernando Castro.

Instructions for Authors: Located at the Amphibian & Reptile Conservation website:

http://amphibian-reptile-conservation.org/submissions.html

volume 8

2014

NUMBER 2