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A distinctive new species of moss salamander (Caudata: Plethodontidae:

Nototriton) from an imperiled Honduran endemism hotspot

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 Zootaxa 2434: 1–16 (2010) ISSN 1175-5326 (print edition)
www.mapress.com / zootaxa/ Article ZOOTAXA
Copyright © 2010 · Magnolia Press ISSN 1175-5334 (online edition)

A distinctive new species of moss salamander (Caudata: Plethodontidae:

Nototriton) from an imperiled Honduran endemism hotspot

JOSIAH H. TOWNSEND1,2,4, J. MICHAEL BUTLER3, LARRY DAVID WILSON2 & JAMES D. AUSTIN3
1
School of Natural Resources and Environment, University of Florida, Gainesville, Florida 32611
2
Instituto Regional de Biodiversidad (IRBio), Centro Zamorano de Biodiversidad, Escuela Agrícola Panamericana Zamorano,
Departamento de Francisco Morazán, Honduras
3
Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, Florida 32611
4
Corresponding author. E-mail: jtwnsnd@ufl.edu

Abstract

A new species of Nototriton is described from Refugio de Vida Silvestre Texiguat in northern Honduras, and is presented
in a new phylogenetic hypothesis for the genus Nototriton. This species differs from all other congeners by a number of
morphological characteristics, including but not limited to large nares, syndactylous hands and feet with pointed toe tips,
and a pale ventral surface with light mottling. Mitochondrial sequence divergence distinguishes this species from all
other Nototriton (3.6–6.0% at 16S and 10.2–15.1% at cytochrome b). Refugio de Vida Silvestre Texiguat harbors a
remarkable herpetofauna, including 20% of all endemic amphibian and reptile species in Honduras, among which are
five species (one anuran, one lizard, and three snakes) known only from the immediate vicinity of the type locality of the
new species. Despite the evolutionary and conservation importance of this reserve, it remains one of the most imperiled
cloud forests in Honduras due to persistent illegal logging and forest clearing.

Key words: Bolitoglossinae, conservation, cloud forest, endemic, Nototriton tomamorum sp. nov., Nuclear Central
America, tropical salamanders, Texiguat, mtDNA, 16S, cytochrome b

Resumen

Se describe una nueva especie de Nototriton del Refugio de Vida Silvestre Texiguat en el Norte de Honduras, y es
presentada en una nueva hipótesis filogenética para el genero Nototriton. Esta especie se diferencia de todos sus
congéneres por un número de características morfológicas, incluyendo, pero no limitadas a, fosas nasales grandes, manos
y pies sindáctilos con las puntas los dedos en punta y superficie ventral pálida con ligeras manchas. Las secuencias
mitocondriales demuestran la diferencia entre esta especie y todas las demás especies en el género Nototriton (3.6–6.0%
en 16S y 10.2–15.1% en citocromo b). El Refugio de Vida Silvestre Texiguat alberga una herpetofauna endémica
destacable, incluyendo 20% de las especies anfibios y reptiles endémicos en Honduras, entre los cuales hay cinco
especies (un anuros, una lagartija, y tres serpientes) que ocurren únicamente en las inmediaciones de la localidad tipo de
la nueva especie. A pesar de la importancia evolutiva y para conservación de esta reserva, sigue siendo uno de los
bosques nublados en mayor peligro en Honduras debido a la persistente tala ilegal y desforestación del bosque.

Palabras claves: Bolitoglossinae, conservación, bosque nublado, endémico, Nototriton tomamorum sp. nov.,
Centroamerica Nuclear, salamandras tropicales, Texiguat, ADNmt, 16S, citocromo b

Introduction

The highlands of Central America are home to a suite of small lungless salamanders (Plethodontidae:
Cryptotriton, Dendrotriton, Nototriton, Oedipina) characterized by conserved morphology that has masked
considerable evolutionary diversification (Wake 1987). Adding to the challenge posed by the often

Accepted by M. Vences: 12 Mar. 2010; published: 23 Apr. 2010 1

unremarkable external morphology of these amphibians is their small size and cryptic nature. Most species are
known only from a handful of specimens, in many cases limited to the type specimen or type series (Wake &
Elias 1983; Good & Wake 1993; McCranie et al. 2008). The majority of species in the four aforementioned
genera were historically considered members of the genus Chiropterotriton (Wake & Elias 1983), and it was
not until 2000 that all four currently accepted genera were recognized (García-París & Wake 2000).
The genus Nototriton comprises thirteen species of miniature salamanders that inhabit Central American
highland forests (AmphibiaWeb 2009). Papenfuss & Wake (1987) and Savage (2002) recognized three species
groups within the genus: the N. barbouri group, the N. picadoi group, and the N. richardi group. These three
groups exhibit two discrete patterns of geographic distribution: the N. barbouri group (five species) is
restricted to the highlands of Honduras and Guatemala, and the N. picadoi group (six species) and N. richardi
group (two species) are found in the highlands of Costa Rica, with a single species in the N. picadoi group, N.
saslaya, endemic to a cloud forest in northern Nicaragua (Köhler 2002).
Honduras is home to a diverse endemic salamander fauna (McCranie & Wilson 1993a, 2002; McCranie et
al. 2008; Townsend et al. 2009), with four of the five currently recognized species of Nototriton in the N.
barbouri group being found there (Fig. 1): N. barbouri (Schmidt), N. brodiei Campbell & Smith, N. lignicola
McCranie & Wilson, and N. limnospectator McCranie, Wilson & Polisar (McCranie 2009). These species, as
well as the majority of endemic Honduran amphibians, are restricted to geographically small highland forests
that are threatened by deforestation and conversion to subsistence agriculture (Wilson & McCranie 2004a,
2004b).
In April 2008, as part of a country-wide on-site assessment of habitat and localities known to support
populations of critically endangered amphibians in Honduras, the authors and a field team visited Refugio de
Vida Silvestre (RVS) Texiguat at the western end of the Cordillera Nombre de Dios in northern Departamento
de Yoro, Honduras. This small cloud forest supports one of the richest endemic herpetofaunas in Honduras,
including a population of salamanders assigned to Nototriton barbouri (McCranie & Wilson 2002; Wilson &
McCranie 2004b), and is considered one of the most at-risk protected areas in Honduras due to a lack of
enforcement and consistent loss of habitat since at least 1991 (McCranie et al. 1993a; McCranie & Castañeda
2004a, 2004b). During our 2008 visit to RVS Texiguat, we collected a single specimen of an undescribed
species of Nototriton that possesses a series of morphological characteristics unique among Honduran
congeners. Comparison of morphological and mitochondrial DNA sequence data confirms that this specimen
represents a distinctive and undescribed species. Below we describe this new species, present a new
phylogenetic hypothesis for the genus Nototriton, and emphasize the need for immediate efforts to further
document and conserve the biodiversity of RVS Texiguat.

Material and methods

Taxa and samples utilized in this study, with their associated locality and museum and GenBank accession
numbers, are summarized in Table 1. Institutional abbreviations follow those standardized by the American
Society of Ichthyologists and Herpetologists (http://www.asih.org/codons.pdf); numbers from the field series
of the first author (JHT) and Ileana R. Luque (IRL) represent voucher specimens deposited at UF in May
2009. Morphological measurements were taken with precision digital calipers and a stereomicroscope with an
optical micrometer; measurements are rounded to the nearest 0.1 mm. Abbreviations used for morphological
measurements are as follows: snout to posterior edge of vent, SVL; axilla–groin length, AG; trunk width at
midbody, TW; head length from tip of snout to gular fold, HL; head width taken at maximum, HW; tail length,
TL; hind limb length, HLL; forelimb length, FLL; combined forelimb and hind limb lengths, CLL; hind foot
length, HFL; hind foot width, HFW; and nares length, NL. Morphological comparisons of the aforementioned
characters are presented in Table 2. Comparative morphological data for other species of Nototriton,
Cryptotriton, and Dendrotriton are taken from Good & Wake (1993), Campbell & Smith (1998), Lynch &
Wake (1978), McCranie et al. (1998), Ehmcke & Clemen (2000), McCranie & Wilson (2002), Savage (2002),
Vásquez-Almazán et al. (2009), and Wake & Campbell (2000).

2 · Zootaxa 2434 © 2010 Magnolia Press TOWNSEND ET AL.

TABLE 1. Samples used in sequence divergence and phylogenetic analyses, with GenBank accession and museum
voucher numbers.
Taxon Locality GenBank accession numbers GenBank voucher
(cob; 16S) specimens
1 N. tomamorum Honduras: Yoro: Texiguat GU971732; GU971731 UF 155377
2 N. barbouri Honduras: Yoro: Macuzal GU971734; GU971733 JHT 2420
3 N. sp. Honduras: Atlántida: Quebrada de Oro AF199136; AF199201 USNM 339712
4 N. sp. Honduras: Atlántida: Cerro Búfalo AF199137; none USNM 497552
5 N. sp. Honduras: Yoro: Texiguat AF199138; none USNM 509333
6 N. brodiei Guatemala: Izabal: Sierra de Caral AF199139; AF199202 UTA A-51490
7 N. lignicola Honduras: Francisco Morazán: Cataguana GU971736; GU971735 JHT 2122
8 N. lignicola Honduras: Olancho: La Muralla AF199141; AF199204 USNM 497540
9 N. lignicola Honduras: Olancho: La Muralla AF199142; none USNM 497550
10 N. limnospectator Honduras: Santa Bárbara: El Ocotillo AF199143; none MVZ 225866
11 N. limnospectator Honduras: Comayagua: Cerro Azul Meámbar GU971738; GU971737 IRL 070
12 N. richardi Costa Rica: San José: Cascajal de las Nubes AF199147; none MVZ 194887
13 N. richardi Costa Rica: San José: Cascajal de las Nubes AF199148; none MVZ 194885
14 N. richardi Costa Rica: San José: Cascajal de las Nubes AF199146; AF199206 UCR 12057
15 N. picadoi Costa Rica: Cartago: Tapantí AF199145; none MVZ 203745
16 N. picadoi Costa Rica: Cartago: Tapantí AF199144; AF199205 MVZ 225899
17 N. guanacaste Costa Rica: Guanacaste: Volcán Cacao AF199140; AF199203 MVZ 207106
18 N. gamezi Costa Rica: Alajuela: Monteverde AF199135; AF199200 MVZ 207122
19 N. abscondens Costa Rica: San José: Cascada de La Paz AF199130; AF199199 UCR 12071
20 N. abscondens Costa Rica: Alajuela: Vara Blanca AF199132; none MVZ 181351
21 N. saslaya Nicaragua: Atlantíco Norte: Saslaya None; GU981761 MVZ 230241
O. gephyra Honduras: Yoro: Texiguat AF199162; AF199218 USNM 530582

Template DNA was extracted from muscle tissue using the QIAGEN PureGene DNA Isolation Kit
following manufacturer’s instructions. Two mitochondrial gene fragments were amplified: a 692 base pair
segment of cytochrome b (cob) gene using primers MVZ15-L and MVZ18-H (Moritz et al. 1992) and a 550
base pair fragment of the 16S large subunit RNA (16S) using 16Sar-L and 16Sbr-H (Palumbi et al. 1991).
Unincorporated nucleotides were removed from PCR product with ExoSAP-IT, and product was sequenced
using forward and reverse primers following standard sequencing protocols and sequences electrophoresed on
an ABI 3130xl (Applied Biosystems, Inc.).
Sequence data for 16S and cob was combined with available samples of Nototriton from GenBank (Table
1) and used in sequence divergence analysis. Sequences were aligned and pairwise comparisons of corrected
sequence divergence were generated using Mega 4.0 (Tamura et al. 2007), using the Kimura-2 parameter
(K2p; Kimura 1980) to provide relatively conservative estimates of pairwise sequence divergence. A dataset
limited to taxa with data available for both 16S and cob and one outgroup taxon (Oedipina gephyra) was used
for phylogenetic analyses. Sequences were trimmed to 497 bp for 16S and 385 bp for cob to match the length
of available sequences from multiple genera.
Bayesian and maximum likelihood (ML) phylogenetic analyses were performed on the 882 bp combined
dataset. We partitioned the dataset by gene (16S) and codon position (1st, 2nd, 3rd), with best-fit models for
each partition determined using the Akaike Information Criterion implemented in MrModeltest2.2 (Nylander
2004). The model GTR+I+Γ was selected for 16S; for cob GTR+Γ was selected for 1st position, K(80)+I+Γ
for 2nd position, and HKY for 3rd position. Bayesian analysis was performed using MrBayes 3.1.2
(Huelsenbeck and Ronquist 2001), and consisted of two parallel runs of four Markov chains (three heated, one
cold) run for 20 x 106 generations and sampled every 1000 generations, with a random starting tree and the

NEW SPECIES OF HONDURAN NOTOTRITON Zootaxa 2434 © 2010 Magnolia Press · 3

first 4000 samples discarded as burnin. The remaining 32,002 post-burnin trees from both runs were used to
generate a 50% majority rule consensus tree. Maximum likelihood analysis was performed in PhyML 3.0
(Guindon & Gascuel 2003), with 1000 bootstrap replicates run on the combined dataset under the GTR
substitution model with four relative substitution rate categories and program-estimated base frequencies,
transition/transversion ratios, and gamma shape parameter.

FIGURE 1. Distribution of Nototriton in eastern Nuclear Central America; shaded areas >1000 m elevation; open star =
type locality of N. tomamorum sp. nov., N. sp.; solid star = N. stuarti; open triangles = N. brodiei; closed triangles = N.
limnospectator; solid square = N. barbouri; open diamonds = N. lignicola; solid diamond = N. saslaya.

Environmental Vulnerability Scores (EVS) were calculated using the methodology developed and refined
by Wilson & McCranie (1992, 2003, 2004a), were generated by taking the total of three rankings: 1) extent of
geographic range, 2a) degree of specialization of reproductive mode for amphibians or 2b) the degree of
persecution by humans for reptiles, and 3) extent of ecological distribution in Honduras; EVS scores from 10–
13 indicate medium vulnerability, and scores from 14–19 are high vulnerability (Wilson & McCranie 2003).
Proposed Red List categorization follows the criteria utilized by the IUCN (2001; 2009). Definitions of forest
formations follow Holdridge (1967) and McCranie & Wilson (2002).

Nototriton tomamorum new species

(Figs. 2, 3)

Holotype: A female (UF 155377) from 2.5 km NNE of La Fortuna (15°25.965’N, 87°18.556’W), 1550 m,
Refugio de Vida Silvestre Texiguat, Departamento de Yoro, Honduras, collected 9 April 2008 by J. Slapcinsky
and the field team of J. M. Butler, L. P. Ketzler, N. M. Stewart, J. H. Townsend, and L. D. Wilson. Original
field number JHT 2437.
Diagnosis. A small member of the genus Nototriton (SVL=26.9 mm; Table 2) based on having 13 costal
grooves (>16 costal grooves in Oedipina), hands and feet longer than broad (hands and feet broader than long
in Bolitoglossa), and nares that are smaller than most Cryptotriton and Dendrotriton (Fig. 3A, B; 0.018 NL/

4 · Zootaxa 2434 © 2010 Magnolia Press TOWNSEND ET AL.

TABLE 2. Morphological and morphometric comparison of species of Nototriton; see Material and Methods section for
explanation of abbreviations; SVL is given in mm. Comparative morphological data for species other than N.
tomamorum are from Good & Wake (1993), Campbell & Smith (1998), McCranie et al. (1998), Ehmcke & Clemen
(2000), Wake & Campbell (2000), Köhler (2002), McCranie & Wilson (2002), Savage (2002).
Species SVL HW/SVL TL/SVL HLL/SVL FLL/SVL HFW/SVL NL/SVL MT VT
N. tomamorum 26.9 0.145 0.911 0.197 0.160 0.037 0.018 26 11
sp. nov.
N. barbouri group
N. barbouri 30.2–39.9 0.104–0.132 1.031–1.398 0.153–0.200 0.142–0.174 0.037–0.060 0.005–0.011 41–54 12–23
N. brodiei 33.2–34.5 0.120 1.420–1.440 0.166–0.180 0.148–0.151 0.040–0.060 0.004–0.005 60–62 23–24
N. lignicola 28.3–33.9 0.103–0.118 0.840–1.059 0.158–0.181 0.137–0.160 0.029–0.040 0.006–0.009 46–54 16–24
N. limnospectator 33.0–38.2 0.095–0.118 1.027–1.297 0.164–0.211 0.156–0.183 0.048–0.061 0.003 42–55 16–26
N. stuarti 32.6 0.138 1.264 0.178 0.172 0.049 0.012 36 20
N. richardi group
N. richardi 21.0–24.0 0.126–0.150 1.072–1.482 0.174–0.187 0.140–0.146 0.044–0.050 0.012–0.016 24 16
N. tapanti 23.5 0.129 1.205 0.174 0.147 0.041 0.013 31 14
N. picadoi group
N. abscondens 21.0–33.0 0.121–0.143 1.013–1.365 0.157–0.206 0.148–0.183 0.058–0.071 0.003–0.014 27–38 8–11
N. gamezi 23.6–26.2 0.130–0.140 1.10–1.30 0.160–0.170 0.152 0.061 0.007 24–35 12–17
N. guanacaste 18.2–29.7 0.145–0.157 1.210–1.337 0.185–0.201 0.170–0.179 0.066–0.072 0.004–0.009 25–49 10–18
N. major 37.9 0.113 1.441 0.174 0.148 0.059 0.003 33 18
N. picadoi 20.0–32.0 0.123–0.144 1.203–1.424 0.180–0.189 0.169–0.182 0.060–0.070 0.010–0.016 – –
N. saslaya 28.1–34.6 0.133–0.155 0.883–1.255 0.217–0.244 0.194–0.210 0.075–0.091 0.002–0.003 17–22 3–11

SVL; 0.020–0.029 NL/SVL in Cryptotriton [except some individuals of C. veraepacis] and Dendrotriton).
Cryptotriton veraepacis has nares ranging from 0.017–0.027 NL/SVL (mean 0.022), and can be differentiated
from N. tomamorum by having a uniformly dark gray ventral surface (ventral surface pale with gray flecks in
N. tomamorum; Fig. 2). Generic placement in Nototriton is also strongly supported by sequence data from the
mitochondrial genes 16S and cob (Table 3; Fig. 4). This new species is distinguished from all other
Nototriton, except N. richardi and N. tapanti, by having syndactylous hands and feet (Fig. 3C, D; hands and
feet with free, differentiated toes in all other species) and relatively large nares (Fig. 3A, B; 0.018 NL/SVL
versus 0.010–0.016 in N. picadoi, 0.003–0.014 in N. abscondens, 0.012 in N. stuarti, 0.005–0.011 in N.
barbouri, 0.006–0.009 in N. lignicola, 0.004–0.009 in N. guanacaste, 0.004–0.005 in N. brodiei, 0.003 in N.
limnospectator, 0.003 in N. major, and 0.002–0.003 in N. saslaya). Nototriton tomamorum can be further
differentiated from members of the N. barbouri group by having a broader head (0.145 HW/SVL versus 0.138
in N. stuarti, 0.104–0.132 in N. barbouri, 0.120 in N. brodiei, 0.103–0.118 in N. lignicola, and 0.095–0.118 in
N. limnospectator) and fewer maxillary teeth (26, versus 36 in N. stuarti, 41–54 in N. barbouri, 42–55 in N.
limnospectator, 46–54 in N. lignicola, and 60–62 in N. brodiei), from members of the N. picadoi group by
having a relatively shorter tail (0.911 TL/SVL, versus 1.441 in N. major, 1.123–1.344 in N. picadoi, 1.013–
1.365 in N. abscondens, 1.210–1.337 in N. guanacaste, and 1.10–1.30 in N. gamezi) and narrower feet (0.037
HFW/SVL, versus 0.058–0.071 in N. abscondens, 0.059 in N. major, 0.060–0.070 in N. picadoi, and 0.066–
0.072 in N. guanacaste), and from N. saslaya by having shorter forelimbs (0.160 FLL/SVL, versus 0.194–
0.210 in N. saslaya) and hind limbs (0.197 HLL/SVL, versus 0.217–0.244 in N. saslaya) and narrower feet
(0.037 HFW/SVL, versus 0.075–0.091 in N. saslaya). The new species also differs from N. richardi and N.
tapanti in having a pale ventral surface mottled with gray chromatophores (ventral surface brown with dark
flecks in N. richardi and dark brown in N. tapanti), by having a tail that is shorter than the snout-vent length
(0.91 TL/SVL, versus 1.072–1.482 in N. richardi and 1.205 in N. tapanti), longer forelimbs (0.160 FLL/SVL,
versus 0.140–0.146 in N. richardi and 0.147 in N. tapanti), longer hind limbs (0.197 HLL/SVL, versus 0.174–
0.187 in N. richardi and 0.174 in N. tapanti), and narrower feet (0.037 HFW/SVL, versus 0.044–0.050 in N.

NEW SPECIES OF HONDURAN NOTOTRITON Zootaxa 2434 © 2010 Magnolia Press · 5

richardi and 0.041 in N. tapanti). This new species is also well differentiated from all other species of
Nototriton based on mitochondrial sequence data (Table 3), and is 3.6–6.0% divergent on 16S and 10.2–
15.1% divergent on cob from all other congeners.

FIGURE 2. Dorsal and ventral aspects of the holotype of Nototriton tomamorum sp. nov.

FIGURE 3. Dorsal (A) and lateral (B) views of the head, and dorsal (C) and ventral (D) aspects of the right hind foot of
Nototriton tomamorum sp. nov., showing the lack of separation or differentiation in the toes, and lack of subdigital pads.

6 · Zootaxa 2434 © 2010 Magnolia Press TOWNSEND ET AL.

NEW SPECIES OF HONDURAN NOTOTRITON Zootaxa 2434 © 2010 Magnolia Press · 7
 Description of holotype. Nototriton tomamorum is known only from a single, presumably female (mental
gland and cloacal papillae absent) specimen, preserved with its mouth open and tongue extended, and is a
relatively small member of the genus (SVL=26.9 mm, total length=51.4 mm) with a slender body and reduced
limbs. Its head is rounded and slightly broader than the body, and the nostrils are relatively large (NL/
SVL=0.018), and the snout is rounded and of moderate length (Fig. 3A, B). The nasolabial protuberances are
apparent but not well developed, and barely extend below the upper lip line. The eyes are relatively large and
protuberant, and the parotoid glands appear large but not well defined. The teeth are exceedingly small; there
are approximately 26 maxillaries, 4 premaxillaries set slightly forward from the maxillary teeth, and 11
vomerines; vomerine teeth are arranged in two short medially-positioned arches. The limbs are short (CLL/
SVL=0.36), with the adpressed limbs being separated by approximately 5.5 costal grooves. The hands and
feet are narrow and have poorly-developed, poorly-differentiated digits that are fused and lack subdigital pads
(Fig. 3C, D). The free tips of digits III on the hands and III and IV on the feet are pointed, and digits I, II and
IV on the hands and I, II, and V on the feet are very short and essentially completely fused, being demarcated
by shallow grooves on the dorsal side of the feet. The relative length of the digits is I<IV<II<III on the hands
and V<I<II<IV<III on the feet. The tail is slightly shorter than the body (TL/SVL=0.91), with a slight basal
constriction most apparent on the ventral side, and is slightly compressed laterally (tail depth 1.17 times tail
width at level of basal constriction).
Measurements of holotype (in mm). SVL 26.9; AG 15.2; TW 3.6; HL 4.8; HW 3.9; TL 24.5; HLL 5.3;
FLL 4.3; CLL 9.6; HFL 1.7; HFW 1.2; NL 0.5; eye length 1.6; eye width 1.2; interorbital distance 1.4;
anterior rim of orbit to snout 1.1; distance separating internal nares 0.8; distance separating external nares 1.8;
snout projection beyond mandible 0.6; tip of snout to axilla 7.7; distance from axilla to groin 15.2; snout to
anterior edge of vent 24.9; tail depth at basal constriction 2.8; tail width at basal constriction 2.4.
Coloration of holotype. Dorsal surfaces of head, body, and tail medium grayish brown, with profuse pale
chromatophores laterally, becoming less abundant dorsolaterally. The head has some pale mottling on the top
of the snout, and two irregular lines of pale chromatophores extending from the lateral region above the
forelimbs onto the posterior portion of the head and parotoid glands. There is a very thin, pale middorsal
stripe, with a herringbone pattern with lines extending from the middorsal stripe posteriorly. There is an
indistinct dark dorsolateral stripe starting about one-third the way down the trunk and extending onto the
proximal one-third of the tail. Ventral surface of head, body, and tail cream, mottled with dark gray
chromatophores, becoming somewhat more profuse toward the distal end of the tail.
Etymology. The specific name “tomamorum” means “belonging to the Tomams.” Tomams are the highest
level of deities recognized in the belief system of the indigenous Tolupan of Honduras, of which there are
four: Tomam Pones Popawai (Grandfather Tomam), his wife Tomam Pones Namawai (mother of all that
exists), and their children Tomam the Elder and Tomam the Younger (Chapman 1992). This name is given in
recognition that the Tolupan are the traditional inhabitants of this area and that this new species is known only
from the Cordillera Nombre de Dios, or “Name of God Mountains,” a name which, ironically, was given by
15th century Spanish explorers.
Natural history. The single known specimen of Nototriton tomamorum was collected during the daytime
from leaf litter packed onto a rock ledge alongside a small creek at about 1550 m elevation in the Lower
Montane Wet Forest formation (Fig. 5). This species is presumably endemic to the vicinity of the type locality,
and is likely restricted to the Lower Montane Wet Forest around the neighboring peaks of Cerro San Francisco
and Cerro Texiguat. The locality can be characterized as a narrow, steep-sided canyon with riparian forest, and
more xeric upland pine-oak forest dominating the terrain above the canyon. The pine-oak forest transitions to
mesic cloud forest (or remnant cloud forest) uphill from this locality. This species is sympatric with a
congener, N. barbouri, and three other plethodontids: Bolitoglossa dofleini, Bolitoglossa porrasorum and
Oedipina gephyra. The canyon where the holotype of N. tomamorum was collected is also the type locality of
the hylid frog Isthmohyla insolita, and a known locality for the anurans Lithobates maculatus, Plectrohyla
guatemalensis, and Ptychohyla spinipollex.
Conservation status. Based on the criteria of the IUCN (2001), Nototriton tomamorum should be
classified as Critically Endangered (B1ab[iii]+2ab[iii]) due to the extremely limited extent of its known

8 · Zootaxa 2434 © 2010 Magnolia Press TOWNSEND ET AL.

distribution and the direct threat to the remaining habitat around that locality. The primary threat to the type
and only known locality for this species is from illegal timber extraction and wholesale slash-and-burn
clearing of forest for bean cultivation once it has been depleted of valuable hardwoods, such as mahogany
(Fig. 6).

FIGURE 4. Bayesian phylogram based on combined dataset of cob and 16S, showing phylogenetic hypothesis for
Nototriton and placement of N. tomamorum (bold) as a weakly support sister taxon to the N. barbouri group. Bayesian
posterior probability values shown above and maximum likelihood bootstrap support values shown below branches;
bootstrap values less than 50% are not shown.

Systematic relationships. Nototriton tomamorum bears the strongest morphological resemblance to

members of the N. richardi group (Table 2), which are endemic to the central highlands of Costa Rica. The
two species, N. richardi and N. tapanti, are the most morphologically distinctive members of the genus,
differing from all other Nototriton by having syndactylous feet, relatively large nostrils, and frontal processes
of the premaxillary arising separately from the maxillary (Good & Wake 1993; Papenfuss & Wake 1987;
Savage 2002). According to Savage (2002), N. richardi is distributed from 1370 to 1800 m elevation along the
Atlantic slope of the Cordillera Central, and N. tapanti at a single locality at 1300 m elevation in the northern
portion of the Cordillera de Talamanca. These localities are situated in the central montane spine of Costa
Rica in Lower Central America and are separated widely from the type locality of N. tomamorum by over 670
airline km and much of the central highlands of Honduras and the lowland Nicaraguan Depression. Despite
the morphological similarities between N. tomamorum and members of the N. richardi group, our
phylogenetic analysis (Fig. 4) does not support the inclusion of N. tomamorum in the N. richardi group.
Preliminary phylogenetic analysis of fragments from two mitochondrial genes (16S and cob) weakly supports
(posterior probability = 0.55, bootstrap support = 58%) N. tomamorum as sister taxon to the rest of the N.
barbouri group (Fig. 4). We also recovered N. richardi as the weakly supported sister (posterior probability =
0.58, bootstrap support = 59%) to the N. picadoi group, a finding consistent with previous phylogenetic
hypotheses for Nototriton and N. richardi (García-París & Wake 2000; Wiens et al. 2007; Adams et al. 2009).

NEW SPECIES OF HONDURAN NOTOTRITON Zootaxa 2434 © 2010 Magnolia Press · 9

FIGURE 5. Riparian habitat at type locality of Nototriton tomamorum, 1550 m elevation; the holotype was collected
from a leaf litter pack in a rock wall along the margins of this stream.

10 · Zootaxa 2434 © 2010 Magnolia Press TOWNSEND ET AL.

FIGURE 6. A. Deforestation in the core zone of RVS Texiguat, 1710 m elevation; recently planted beans in the
foreground, with rapidly progressing land clearing visible on the far ridges in the highest part of the reserve; B. Recently
cut mahogany logs at the end of the road accessing the vicinity of the type locality of Nototriton tomamorum; these logs
were exposed the day we arrived and were partially covered up the second day our team was in the area in an attempt to
conceal them. C. Recent deforestation less than 1 km upstream from the type locality of Nototriton tomamorum and the
endemic treefrog Isthmohyla insolita; two of our co-workers are pictured in the lower-central portion of the photograph
for scale.

Sequence data from the holotype of Nototriton tomamorum are at least 12.5% (16S) and 20.6% (cob)
divergent from representatives of morphologically-similar genera of Central American plethodontids
(Cryptotriton, Dendrotriton, and Oedipina), but less than 6.0% (16S) and 15.1% (cob) divergent from other
species of Nototriton, supporting placement of N. tomamorum in this genus (Table 3). Within the genus
Nototriton, the new species demonstrates differing patterns of sequence divergence on the two genes sampled.
For 16S, N. tomamorum is closest to N. limnospectator (3.6–3.8%), a biogeographically-logical situation
given the relative geographical proximity of RVS Texiguat to the type locality of N. limnospectator,
approximately 100 airline km to the WSW in Parque Nacional Montaña de Santa Bárbara; however N.
tomamorum is 12.4% divergent from N. limnospectator for cob, and for that gene is closer to the Costa Rican
species N. abscondens (10.2%) and N. gamezi (11.5%) than to any other member of the N. barbouri group
(Table 3).
Given the morphological differentiation and lack of strong phylogenetic support, we refrain from
assigning N. tomamorum to any recognized species group pending acquisition and analysis of additional
molecular data from mitochondrial and nuclear genes.
Our phylogenetic analyses included taxa and populations of Nototriton not available in previous studies
(García-París & Wake 2000; Wiens et al. 2007), including a sample from the vicinity of the type locality of N.
barbouri (“Portillo Grande, Yoro, Honduras” [Schmidt 1936]). Inclusion of these data reveals the taxon N.
barbouri to be paraphyletic (Fig. 4), with strong support for N. barbouri sensu stricto as the sister species of

NEW SPECIES OF HONDURAN NOTOTRITON Zootaxa 2434 © 2010 Magnolia Press · 11

N. limnospectator and for the remaining populations assigned to N. barbouri (N. sp. in Fig 4) from the
Cordillera Nombre de Dios to be sister to N. brodiei from NE Guatemala and NW Honduras. We are currently
preparing an intergrative systematic revision of the N. barbouri group that will use molecular and
morphological analyses to resolve the taxonomic status of populations assigned to N. barbouri.

TABLE 4. Summary of conservation priority amphibian and reptiles from Reserva de Vida Silvestre Texiguat; distribution is
characterized as Endemic (restricted to Honduras), NCA (restricted to Nuclear Central America), or Widespread (range extending
outside of NCA); IUCN Red List status abbreviations are CR (Critically Endangered), EN (Endangered), VU (Vulnerable); methods
for calculating Environmental Vulnerability Score are detailed in the Materials and Methods section.

Taxa Distribution IUCN Red List Status Environmental Vulnerability Score

Caudata
Bolitoglossa porrasorum Endemic EN 15
Nototriton barbouri Endemic EN 15
Nototriton tomamorum Endemic CR 17
Oedipina gephyra Endemic EN 16
Anura
Craugastor aurilegulus Endemic EN 14
Craugastor saltuarius Endemic CR 16
Craugastor stadelmani Endemic CR 15
Duellmanohyla salvavida Endemic CR 12
Isthmohyla insolita Endemic CR 16
Incilius leucomyos Endemic EN 11
Plectrohyla guatemalensis NCA CR 9
Ptychohyla spinipollex Endemic EN 11
Squamata
Anolis kreutzi Endemic CR 16
Anolis purpurgularis Endemic EN 15
Anolis yoroensis Endemic EN 14
Bothriechis marchi Endemic EN 16
Celestus scansorius Endemic EN 14
Cerrophidion godmani Widespread VU 12
Drymobius chloroticus Widespread VU 11
Geophis damiani Endemic CR 15
Mastigodryas dorsalis NCA VU 12
Ninia pavimentata NCA EN 12
Omoadiphas texiguatensis Endemic CR 14
Rhadinaea godmani NCA VU 9
Rhadinaea kinkelini NCA VU 12
Rhadinaea tolpanorum Endemic EN 15
Sphenomorphus incertus NCA VU 12

Discussion

The herpetofauna of the RVS Texiguat is characterized by a remarkable degree of endemism (McCranie &
Castañeda 2004a; Wilson & McCranie 2004b). Currently, 39 species are known from 1550 m in elevation and

12 · Zootaxa 2434 © 2010 Magnolia Press TOWNSEND ET AL.

above in this refuge. Of this number, 27 are considered to be conservation priority species (Table 4), defined
as those species judged to occupy one of three IUCN categories Critically Endangered, Endangered, or
Vulnerable. Nine of the 27 are considered Critically Endangered, 12 are Endangered, and six are Vulnerable;
19 (70.4%) of these species (four salamanders, seven anurans, four lizards, and four snakes) are endemic to
Honduras (Table 4). Of the remainder, six are endemic to Nuclear Central America and two have widespread
distributions. As indicated by Townsend & Wilson (In press), 91 species of amphibians and reptiles are
endemic to Honduras, thus the Texiguat figure is 20.9% of the total. Therefore, RVS Texiguat qualifies as a
major center of endemism in Honduras, as well as in the whole of eastern Nuclear Central America (as
defined by Campbell, 1999). The Environmental Vulnerability Scores, as tabulated by Townsend and Wilson
(in press) and indicated in Table 4, indicate that the values for the Texiguat conservation priority species range
from 9 to 17. Most of these species involved (16) are considered high vulnerability species (59.3%), nine
(33.3%) are medium vulnerability species, and only two (7.4%) have scores lying at the upper end of the
range for low vulnerability species (Table 4). In summary, of the 27 species judged to be of conservation
priority, 19 are endemic to Honduras and 16 are of high vulnerability. Based on the analysis above, Refugio de
Vida Silvestre Texiguat is an area of immense importance for the conservation of the Honduran herpetofauna,
especially its endemic component.
Unfortunately, little attention has been paid to RVS Texiguat by conservation authorities past its initial
designation as a protected area in 1987 (Wilson et al. 2001). Since 1991, ten species (including the present
one) with type localities in the area have been described: Anolis kreutzi, A. purpurgularis, A. yoroensis,
Celestus scansorius, Geophis damiani, Isthmohyla insolita, Nototriton tomamorum, Oedipina gephyra,
Omoadiphas texiguatensis, and Rhadinaea tolpanorum (McCranie et al. 1993a, b; Wilson et al. 1998;
McCranie & Castañeda 2004a). Given the pattern of these finds, we expect that additional new taxa remain to
be described from RVS Texiguat. During this period, significant environmental damage has been observed,
most of it apparently as a consequence of the illegal extraction of mahogany and the subsequent conversion of
the remaining forest to cropland for use in subsistence agriculture (Fig. 6). Most recently during our visit in
April 2008, we observed that a large stretch of forest upstream from the type locality of Nototriton
tomamorum and Isthmohyla insolita had been clear-cut down to the streamside and planted to beans (Fig. 6C).
Damage to the forests in this refuge also has occurred by natural means. The stream that constitutes the
type locality of Nototriton tomamorum and Isthmohyla insolita (Fig. 5) was severely damaged by Hurricane
Mitch in 1998. At that time, the habitat supporting I. insolita was scoured by high waters in the narrow
canyon. The moss masses clinging to the branches of small trees overhanging the stream, within which the
adults laid their eggs, were stripped away. Our trip in 2008 evidenced that the moss has yet to be reestablished
as we found it in 1991 when I. insolita was discovered. Whether the reestablishment of these conditions will
occur is doubtful. Isthmohyla insolita has been located in one or two nearby streams (McCranie & Castañeda
2007), but the same anthropogenic activities detailed above also are occurring there.
It is apparent that the natural habitat of this hotspot for herpetofaunal endemism will continue to succumb
to the forces of habitat destruction unless steps are taken to protect it. Therefore, we use this opportunity not
only to describe yet another endemic species from this reserve, but to attempt to raise RVS Texiguat’s
international profile and issue an urgent call for its conservation. We offer the following recommendations,
and emphasize the need for immediate and sustained action to prevent further loss of habitat:

• Carry out intensive reconnaissance of RVS Texiguat on the Caribbean side of Cerro Texiguat and Cerro San
Francisco, to search for and identify previously unknown areas supporting populations of conservation
priority species. The majority of work in RVS Texiguat has taken place in the vicinity of La Fortuna,
which is accessed by means of a logging road established sometime before 1991. The Caribbean slope of
RVS Texiguat remains virtually unexplored and has no known access routes, and so exploratory work is
needed immediately to determine if this areas supports target species.
• Establish a habitat and species monitoring program led by Honduran biologists and local guardarecursos.
This program has been initiated in cooperation with PROLANSATE (the organization tasked with
managing RVS Texiguat). Initial plans are to hire six guardarecursos in early 2010 and begin training and
capacity building for development of long-term monitoring teams.

NEW SPECIES OF HONDURAN NOTOTRITON Zootaxa 2434 © 2010 Magnolia Press · 13

• Identify means of mitigating illegal logging and subsequent habitat destruction on the southern side of the
RVS Texiguat. The extraction of high-value hardwoods from RVS Texiguat by way of the drier southern
side of the mountain is well established and obviously influences the local economy; the subsequent
clearing of relatively large plots of forest for cultivation of beans appears to be the work of relatively few
individuals. While legal action against these actors is unlikely, identification of the most influential of
these stakeholders and initiation of a dialogue with them is the first step toward mitigating the destruction
of remaining habitat in the vicinity of La Fortuna.
• Initiate public outreach and environmental education projects in buffer zone communities. Awareness of the
unique nature and value of the biodiversity RVS Texiguat, or even the very existence of the reserve as a
legal entity, is virtually non-existent. Any scientific-based conservation efforts need to be supported by
efforts to empower local actors through education and capacity building, in an attempt to engender a local
desire to conserve the natural resources of RVS Texiguat for the foreseeable future.

If these steps are not taken, it is likely that this significant center of herpetofaunal endemism will succumb
to the pressures of severe, and largely unnecessary, environmental degradation. Given that fully one-fifth of
Honduras’ endemic herpetofauna occurs in RVS Texiguat, this simply cannot be allowed to happen.

Acknowledgments

Fieldwork was supported by a grant from the Critical Ecosystem Partnership Fund. Centro Zamorano de
Biodiversidad provided us with a vehicle and other logistical support in 2008, and we especially thank Jorge
Iván Restrepo, Arie Sanders, José Mora, Suyapa Meyer, and Fredy Membreño for facilitating this support.
Research and exportation permits were furnished by Carla Cárcamo de Martínez and Iris Acosta O.
(Departamento de Areas Protegidas y Vida Silvestre) and fieldwork was completed under permits AFE-
COHDEFOR Resolucion GG-MP-055-2006 and Dictamen DAPVS 0091-2006. Field assistance in April
2008 was provided by Lorraine Ketzler, John Slapcinsky, and Nathaniel Stewart. We thank Mario García-
París and David Wake for providing unpublished sequence data for Nototriton saslaya for use in our
comparative dataset, Ileana Luque-Montes for translating the abstract, and John Slapcinsky for assisting with
photography of the holotype. We also thank David Wake and an anonymous reviewer, whose
recommendations helped to greatly improve this paper.

Literature cited

Adams, D.C., Berns, C.M., Kozak, K.H. & Wiens, J.J. (2009) Are rates of species diversification correlated with rates of
morphological evolution? Proceedings of the Royal Society B, 276, 2729–2738.
AmphibiaWeb (2009) Berkeley, California: AmphibiaWeb: Information on amphibian biology and conservation [web
application]. Available: http://amphibiaweb.org/. Accessed 12 Dec 2009.
Campbell, J.A. (1999) Distribution patterns of amphibians in Middle America. In: Duellman, W.E. (Ed.), Patterns of
Distribution of Amphibians: a global perspective. Johns Hopkins University Press, Baltimore, Maryland, pp. 111–
210.
Campbell, J.A. & Smith, E.N. (1998) New species of Nototriton (Caudata: Plethodontidae) from eastern Guatemala.
Scientific Papers of the Natural History Museum of the University of Kansas, 6, 1–8.
Chapman, A.M. (1992). Masters of Animals: Oral Traditions of the Tolupan Indians, Honduras. Gordon and Breach
Scientific Publishers, Philadelphia, Pennsylvania. xxi + 259 pp.
Ehmcke, J. & Clemen, G. (2000) Teeth and their sex-dependent dimorphic shape in three species of Costa Rican
plethodontid salamanders (Amphibia: Urodela). Annals of Anatomy, 182, 403–414.
García-París, M. & Wake, D.B. (2000) Molecular phylogenetic analysis of relationships of the tropical salamander
genera Oedipina and Nototriton, with descriptions of a new genus and three new species. Copeia, 2000, 42–70.
Good, D.A. & Wake, D.B. (1993) Systematic studies of the Costa Rican moss salamanders, genus Nototriton, with
descriptions of three new species. Herpetological Monographs, 7, 131–159.

14 · Zootaxa 2434 © 2010 Magnolia Press TOWNSEND ET AL.

Guindon S. & Gascuel, O. (2003) A simple, fast, and accurate algorithm to estimate large phylogenies by maximum
likelihood. Systematic Biology, 52, 696–704.
Holdridge, L.R. (1967) Life Zone Ecology. Revised Edition. Tropical Science Center, San José, Costa Rica. 206 p.
Huelsenbeck, J.P. & Ronquist, F.R. (2001) MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics
(Oxford), 17, 754–755.
IUCN. (2001) IUCN Red List Categories and Criteria: Version 3.1. IUCN Species Survival Commission. IUCN, Gland,
Switzerland and Cambridge, United Kingdom.
IUCN. (2009) IUCN Red List of Threatened Species. Version 2009.2 [web application]. Available: http://
www.iucnredlist.org. Accessed 12 Dec 2009.
Kimura, M. (1980) A simple method for estimating evolutionary rate of base substitutions through comparative studies
of nucleotide sequences. Journal of Molecular Evolution, 16, 111–120.
Köhler, G. (2002) A new species of salamander of the genus Nototriton from Nicaragua (Amphibia: Caudata:
Plethodontidae). Herpetologica, 58, 205–210.
Lynch, J.F. & Wake, D.B. (1978) A new species of Chiropterotriton (Amphibia: Caudata) from Baja Verapaz,
Guatemala, with comments on relationships among Central American members of the genus. Contributions in
Science, Natural History Museum of Los Angeles County, 294, 1–22.
McCranie, J.R. (2009) Amphibians and reptiles of Honduras. Listas Zoológicas Actualizadas UCR. Museo de Zoología
UCR, San Pedro, Costa Rica. Available from (http://museo.biologia.ucr.ac.cr/Listas/LZAPublicaciones.htm).
Accessed 5 January 2010.
McCranie, J.R. & Wilson, L.D. (1993) A review of the Bolitoglossa dunni group (Amphibia: Caudata) from Honduras
with the description of three new species. Herpetologica, 49, 1–15.
McCranie, J.R., Wilson, L.D. & Williams, K.L. (1993a) A new species of Oedipina (Amphibia: Caudata:
Plethodontidae) from northern Honduras. Proceedings of the Biological Society of Washington, 106, 385–389.
McCranie, J.R., Wilson, L.D. & Williams, K.L. (1993b) New species of tree frog of the genus Hyla (Anura: Hylidae)
from northern Honduras. Copeia, 1993, 1057–1062.
McCranie, J.R., Wilson, L.D. & Polisar, J. (1998) Another new montane salamander (Amphibia: Caudata:
Plethodontidae) from Parque Nacional Santa Bárbara, Honduras. Herpetologica, 54, 455–461.
McCranie, J.R., Vieites, D.R. & Wake, D.B. (2008) Description of a new divergent lineage and three new species of
Honduran salamanders of the genus Oedipina (Caudata, Plethodontidae). Zootaxa, 1930, 1–17.
McCranie, J.R. & Castañeda, F.E. (2004a) A new species of snake of the genus Omoadiphas (Reptilia: Squamata:
Colubridae) from the Cordillera Nombre de Dios in northern Honduras. Proceedings of the Biological Society of
Washington, 117, 311–316.
McCranie, J.R. & Castañeda, F.E. (2004b) Notes on the second specimens of Geophis damiani Wilson, McCranie and
Williams and Rhadinaea tolpanorum Holm and Cruz D. (Colubridae). Herpetological Review, 35, 341.
McCranie, J.R. & Castañeda, F.E. (2007) Guía de Campo de los Anfibios de Honduras. Bibliomania!, Salt Lake City,
Utah, 304 pp.
McCranie, J.R. & Wilson, L.D. (2002) The Amphibians of Honduras. Society for the Study of Amphibians and Reptiles,
Contributions in Herpetology, 625 pp.
Moritz, C., Schneider, C.J. & Wake, D.B. (1992) Evolutionary relationships within the Ensatina eschscholtzii complex
confirm the ring species interpretation. Systematic Biology, 41, 273–291.
Nylander, J.A.A. (2004) MrModeltest v2. Program distributed by the author. Evolutionary Biology Centre, Uppsala
University.
Palumbi, S.R., Martin, A.P., Romano, S., Mcmillan, W.O., Stice, L. & Grabowski, G. (1991) The simple fool’s guide to
PCR. Special Publication of the Department of Zoology, University of Hawaii, Honolulu. 46 p.
Papenfuss, T.J. & Wake, D.B. (1987) Two new species of plethodontid salamanders (genus Nototriton) from Mexico.
Acta Zoologica Mexicana, 21, 1–16.
Savage, J.M. (2002) The Amphibians and Reptiles of Costa Rica: A Herpetofauna Between Two Continents, Between
Two Seas. University of Chicago Press. Chicago, Illinois. xx + 934 pp.
Schmidt, K.P. (1936) New amphibians and reptiles from Honduras in the Museum of Comparative Zoology. Proceedings
of the Biological Society of Washington 49, 43–50.
Tamura, K., Dudley, J., Nei, M. & Kumar, S. (2007) MEGA4: Molecular Evolutionary Genetics Analysis (MEGA)
software version 4.0. Molecular Biology and Evolution, 24, 1596–1599.
Taylor, E.H. (1949) New salamanders from Costa Rica. University of Kansas Science Bulletin, 33, 279–288.
Townsend, J.H., Butler, J.M., Wilson, L.D. & Austin, J.D. (2009) A new species of salamander in the Bolitoglossa dunni
group (Caudata: Plethodontidae: Bolitoglossinae) from Parque Nacional Montaña de Yoro. Salamandra, 45, 95–
105.
Townsend, J.H. & Wilson, L.D. (In press) Conservation of the Honduran herpetofauna: issues and imperative. In:
Wilson, L.D., Townsend, J.H., & Johnson, J.D. (Eds.). Conservation of Mesoamerican Amphibians and Reptiles.
Eagle Mountain Publishing LC, Eagle Mountain, Utah, pp. 402–429.

NEW SPECIES OF HONDURAN NOTOTRITON Zootaxa 2434 © 2010 Magnolia Press · 15

 Vásquez-Almazán, C.R., Rovito, S.M., Good, D.A. & Wake, D.B. (2009) A new species of Cryptotriton (Caudata:
Plethodontidae) from eastern Guatemala. Copeia, 2009, 313–319.
Wake, D.B. (1987) Adaptive radiation of salamanders in Middle American cloud forests. Annals of the Missouri
Botanical Garden, 74, 242–264.
Wake, D.B. & Campbell, J.A. (2000) A new species of diminutive salamander (Amphibia: Caudata: Plethodontidae:
Nototriton) from the Montañas del Mico of Guatemala. Proceedings of the Biological Society of Washington, 113,
815–819.
Wake, D.B. & Elias, P. (1983) New genera and a new species of Central American salamanders, with a review of the
tropical genera (Amphibia, Caudata, Plethodontidae). Contributions in Science, Natural History Museum of Los
Angeles County, 345, 1–19.
Wiens, J.J., Parra-Olea, G., García-París, M. & Wake, D.B. (2007) Phylogenetic history underlies elevational biodiversity
patterns in tropical salamanders. Proceedings of the Royal Society B, 274, 919–928.
Wilson, L.D. & McCranie, J.R. (1992) Status of amphibian populations in Honduras. Unpublished Report to the
Declining Amphibian Populations Task Force, 15 August 1992. 14 p.
Wilson, L.D. & McCranie, J.R. (1998) Amphibian population decline in a Honduran national park. Froglog, 25, 1–2.
Wilson, L.D., McCranie, J.R. & Espinal, M.R. (2001) The ecogeography of the amphibians and reptiles of Honduras and
the design of herpetological reserves. In: Johnson, J.D., Webb, R.G., & Flores-Villela, O.A. (Eds.). Mesoamerican
Herpetology: Systematics, Zoogeography, and Conservation. Centennial Museum, Special Publication No. 1, The
University of Texas at El Paso, pp. 109–158.
Wilson, L.D. & McCranie, J.R. (2003) Herpetofaunal indicator species as measures of environmental stability in
Honduras. Caribbean Journal of Science, 39, 50–67.
Wilson, L.D. & McCranie, J.R. (2004a) The conservation status of the herpetofauna of Honduras. Amphibian and Reptile
Conservation, 3, 6–33.
Wilson, L.D. & McCranie, J.R. (2004b) The herpetofauna of the cloud forests of Honduras. Amphibian and Reptile
Conservation, 3, 34–48.
Wilson, L.D., McCranie, J.R. & Williams, K.L. (1998) A new species of Geophis of the sieboldi group (Reptilia:
Squamata: Colubridae) from northern Honduras. Proceedings of the Biological Society of Washington, 111, 410–
417.

16 · Zootaxa 2434 © 2010 Magnolia Press TOWNSEND ET AL.

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