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37
Amphib. Reptile Conserv. December 2018 | Volume 12 | Number 2 | e167
Amphibian & Reptile Conservation
12(2) [Special Section]: 37–54 (e167).
urn:lsid:zoobank.org:pub:440CB3D6-450A-463B-B3D3-1CCBCBD8670E
Two new species of Chiropterotriton (Caudata:
Plethodontidae) from central Veracruz, Mexico
1Mirna G. García-Castillo, 2Ángel F. Soto-Pozos, 3J. Luis Aguilar-López, 4Eduardo Pineda,
and 5Gabriela Parra-Olea
1,2,5Departamento de Zoología, Instituto de Biología, Universidad Nacional Autónoma de México, AP 70-153, Tercer Circuito Exterior s/n, Ciudad
Universitaria, México, Distrito Federal, MÉXICO 3,4Red de Biología y Conservación de Vertebrados, Instituto de Ecología, A.C., Carretera Antigua
a Coatepec No. 351, El Haya, CP. 91070, Xalapa, Veracruz, MÉXICO
Abstract.—The lungless salamanders of the tribe Bolitoglossini show notable diversication in the Neotropics,
and through the use of molecular tools and/or new discoveries, the total number of species continues to
increase. Mexico is home to a great number of bolitoglossines primarily distributed along the eastern, central,
and southern mountain ranges where the genus Chiropterotriton occurs. This group is relatively small, with 16
described species, but there remains a considerable number of undescribed species, suggested by the use of
molecular tools in the lab more than a decade ago. Most of these undescribed species are found in the state
of Veracruz, an area characterized by diverse topography and high salamander richness. Described herein
are two new species of Chiropterotriton based on molecular and morphological data. Both new species were
found in bromeliads in cloud forests of central Veracruz and do not correspond to any previously proposed
species. Phylogenetic reconstructions included two mitochondrial fragments (L2 and COI) and identied are
two primary assemblages corresponding to northern and southern species distributions, concordant with
previous studies. The two new species are closely related but morphologically and molecularly differentiated
from other species of the C. chiropterus group.
Keywords. Salamanders, bolitoglossines, bromeliads, phylogenetics, cryopreservation, living tissue, biobanking
Citation: García-Castillo MG, Soto-Pozos ÁF, Aguilar-López JL, Pineda E, Parra-Olea G. 2018. Two new species of Chiropterotriton (Caudata:
Plethodontidae) from central Veracruz, Mexico. Amphibian & Reptile Conservation 12(2) [Special Section]: 37–54 (e167).
Copyright: © 2018 García-Castillo et al. This is an open-access article distributed under the terms of the Creative Commons Attribution-NonCommer-
cialNoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any medium, provided
the original author and the ofcial and authorized publication sources are recognized and properly credited. The ofcial and authorized publication
credit sources, which will be duly enforced, are as follows: ofcial journal title Amphibian & Reptile Conservation; ofcial journal website <amphibian-
reptile-conservation.org>.
Received: 30 October 2018; Accepted: 01 December 2018; Published: 31 December 2018
Ofcial journal website:
amphibian-reptile-conservation.org
Introduction
Due to their unique topography and geological history, the
Mexican highlands have played an important role in the
evolution of plethodontid salamanders (Wake and Lynch
1976; Darda 1994). Particularly, the tribe Bolitoglossini
(Wake 2012) underwent an adaptive radiation and
diversication in the mountainous regions of Mexico
(Wake and Lynch 1976; Wake 1987), resulting in 40% of
the representative biodiversity of the group (AmphibiaWeb
2018). With the aid of molecular tools and recent expedition
activity, the number of described species has increased in
recent years (Parra-Olea et al. 2016; García-Castillo et al.
2017; Sandoval-Comte et al. 2017).
In Mexico, plethodontid richness is concentrated in
regions with rugged topography and a corresponding
great diversity of habitats and microhabitats (Wake et al.
1992; Rovito et al. 2009). These characteristics are found
in the central region of Veracruz, where two important
mountain systems converge: the Trans Mexican Volcanic
Belt (TMVB) and the Sierra Madre Oriental (SMO). The
state of Veracruz has the second highest salamander di-
versity in Mexico with 36 species, after Oaxaca with 42
species (Parra-Olea et al. 2014).
The genus Chiropterotriton includes 16 described
species with seven populations suggested as candidate
species in previous phylogenetic analyses: C. sp. C, C.
sp. F, C. sp. G, C. sp. H, C. sp. I, C. sp. J, and C. sp.
K (Darda 1994; Parra-Olea 2003). Two of the described
species, C. lavae (Taylor) and C. chiropterus (Cope), and
two candidate species (C. sp. C and C. sp. H) occur in
Veracruz (Fig. 1). Describe herein are two new species
Correspondence. 1 biol.mirnagarcia@gmail.com 2 angelfsotop90@gmail.com 3 jlal.herp@gmail.com 4 eduardo.pineda@inecol.mx
5 gparra@ib.unam.mx (Corresponding author)
38
Amphib. Reptile Conserv. December 2018 | Volume 12 | Number 2 | e167
García-Castillo et al.
of Chiropterotriton based on analysis of two mitochon-
drial fragments (L2 and COI) and differing morphologi-
cal characteristics. Specimens were discovered during
recent expeditions in the mountainous regions of central
Veracruz but could not be assigned to any current species
due to their unique morphological and genetic differen-
tiation. Furthermore, these proposed new salamanders do
not belong to any candidate species postulated by Darda
(1994) and Parra-Olea (2003).
Methods
Molecular Analyses
Genomic DNA was extracted from liver, intestine, and
tail tissue samples from 38 Chiropterotriton individuals
and Aquiloeurycea cephalica and Parvimolge townsendi
using a DNeasy tissue kit (Qiagen, Valencia, California,
USA). Amplied two mitochondrial fragments using
primers LX12SN1 and LX16S1R for L2 (partial 12S ri-
bosomal subunit, the tRNA, and large subunit16S; Zhang
et al. 2008) and dgLCO and dgHCO for COI (Meyer
2003). PCR conditions were as follows: L2, 35 cycles of
96 ºC (120 s), 55 ºC (60 s), and 72 ºC (300 s), and COI,
35 cycles of 94 ºC (30 s), 50 ºC (30 s), and 72 ºC (45 s).
PCR products were cleaned with ExoSap-IT (USB Cor-
poration, Cleveland, Ohio, USA) and sequenced with a
BigDye Terminator v3.1 cycle sequencing kit (Applied
Biosystems, Foster City, California, USA). Products
were puried using Sephadex G-50 (GE Healthcare) and
an ABI3730 capillary sequencer to run sequences. Ad-
ditionally, 13 Chiropterotriton sequences were obtained
from previous studies (Parra-Olea 2003; Rovito et al.
2015) to complete the study. Voucher information for all
sequences are shown in Table 1.
Sequencher 5.0.1 (Gene Codes Corporation) was used
to edit and assemble sequences and Muscle 3.8 (Edgar
2004) to align fasta les. Mesquite v3.40 (Maddison and
Maddison 2018) was applied to review and concatenate
data matrices and calculate Kimura 2-parameter (K2P)
corrected genetic distances (Table 2). DNA substitution
models were calculated using PartitionFinder v1.0 (Lan-
fear et al. 2012) under the Bayesian information crite-
rion (BIC), and estimated a maximum likelihood (ML)
tree from RAxML v8.2 (Stamatakis 2014) with 1,000
bootstrap replicates and a GTR+G substitution model.
Additionally, MrBayes v3.2 (Huelsenbeck and Ronquist
2001) was applied for Bayesian analysis with 20 million
generations, sampling every 1,000 generations, and four
chains used to construct a majority consensus tree. Tracer
v.1.7 (Rambaut et al. 2018) was administered to check
stationarity and convergence of chains. Lastly, both phy-
Fig. 1. Map of sampled localities for phylogenetic analyses of the genus Chiropterotriton. White circles correspond to: 1) C. priscus,
2) C. miquihuanus, 3) C. infernalis, 4) C. cieloensis, 5) C. cracens, 6) C. multidentatus (Cd. Maiz), 7) C. multidentatus (Rancho
Borbotón), 8) C. multidentatus (Sierra de Álvarez), 9) C. magnipes, 10) C. mosaueri, 11) C. chondrostega, 12) C. terrestris, 13) C.
arboreus, 14) C. dimidiatus, 15) C. chico, 16) C. sp. G, 17) C. orculus, 18) C. sp. I, and 19) C. sp. K.
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Amphib. Reptile Conserv. December 2018 | Volume 12 | Number 2 | e167
Two new Chiropterotriton from central Veracruz, Mexico
Table 1. Voucher information, localities, GenBank accessions, coordinates and elevation data from specimens used for phylogenetic
analyses. Collection abbreviations: CARIE, Colección de Referencia de Anbios y Reptiles del Instituto de Ecología, A.C.; IBH,
Colección Nacional de Anbios y Reptiles, Instituto de Biología, UNAM; MVZ, Museum of Vertebrate Zoology, University of
California, Berkeley, California, USA. NOTE: Asterisks indicate data inferred indirectly from the available information.
Species Voucher
Number Locality 16S GenBank COI
GenBank Latitude Longitude Elevation
m asl
C. arboreus IBH28191
Hidalgo: 6.8 km SW (by rd)
of Zacualtipan on road to
Tianguistengo
MK335386 MK335232 20.702 -98.667 2029
C. aureus IBH31040
Veracruz: 6.5 km N from Atzalan,
ejido de desarrollo urbano
Quetzalcoatl
MK335395 MK335241 19.843 -97.231 1249
C. aureus IBH31041
Veracruz: 6.5 km N from Atzalan,
ejido de desarrollo urbano
Quetzalcoatl
MK335398 MK335244 19.843 -97.231 1249
C. aureus IBH31042
Veracruz: 6.5 km N from Atzalan,
ejido de desarrollo urbano
Quetzalcoatl
MK335396 MK335242 19.843 -97.231 1249
C. aureus IBH31043
Veracruz: 6.5 km N from Atzalan,
ejido de desarrollo urbano
Quetzalcoatl
MK335394 MK335240 19.843 -97.231 1249
C. aureus IBH31044
Veracruz: 6.5 km N from Atzalan,
ejido de desarrollo urbano
Quetzalcoatl
MK335397 MK335243 19.843 -97.231 1249
C. chico MVZ200679 Hidalgo: 3.8 km S Mineral del
Chico AY522471 – 20.180 -98.731 2630
C. chiropterus CARIE0777 Veracruz: Huatusco MK335407 MK335253 19.185 -96.959 1280
C. chiropterus CARIE0719 Veracruz: Huatusco MK335408 – 19.185 -96.959 1280
C. chondrostega IBH30098
Hidalgo: 1.0 km S (by rd) of La
Encarnacion on road to MX85,
Parque Nacional los Marmoles
MK335383 MK335229 20.866 -99.219 2471
C. cieloensis IBH28181
Tamaulipas: 0.2 km E (by air) of
Rancho El Cielo, 6.9 km NNW (by
air) of the center of Gomez Farías,
Reserva de la Biosfera El Cielo
MK335385 MK335231 23.100 -99.190 1174
C. cracens IBH28192
Tamaulipas: Road from Alta Cima
to San Jose, 1.3 km NE (by air) of
San Jose, Reserva de la Biosfera
El Cielo
MK335384 MK335230 23.059 -99.226 1320
C. dimidiatus IBH28196
Hidalgo: 4.1 km S (by rd) of
Mineral del Chico on the road
to Pachuca, Parque Nacional El
Chico
MK335390 MK335236 20.198 -98.727 2768
C. infernalis MVZ269665
Tamps: Cueva del Brinco, Conrado
Castillo, ca. 43.5 km SW (by rd) of
Ejido Guayabas
MK335382 MK335228 23.959 -99.474 1920
C. lavae IBH22369 Veracruz: 200 m N hwy 140 at
La Joya MK335393 MK335239 19.614 -97.030 2060
C. magnipes IBH28176
Hidalgo: “El Coní,” 900 m SSE of
the center of Durango, Municipio
Zimapan, Parque Nacional los
Marmoles
MK335387 MK335233 20.888 -99.226 2234
C. miquihuanus IBH30329
Nuevo León: 1.8 km S (by rd) of
La Encantada on road from La
Bolsa to Zaragoza
MK335381 MK335227 23.893 -99.803 2803
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Amphib. Reptile Conserv. December 2018 | Volume 12 | Number 2 | e167
García-Castillo et al.
Species Voucher
Number Locality 16S GenBank COI
GenBank Latitude Longitude Elevation
m asl
C. mosaueri IBH28179
Hidalgo: “El Coní,” 900 m SSE
of center of Durango, Municipio
Zimapan, Parque Nacional los
Marmoles
MK335388 MK335234 20.888 -99.226 2234
C. multidentatus IBH28177
San Luis Potosí: Cueva el
Madroño, 900 m NW (by air)
of the entrance to Valle de los
Fantasmas on MX70, Sierra de
Alvarez
MK335416 – 22.071 -100.614 2297
C. multidentatus IBH30102
San Luis Potosí: Cueva el
Madroño, 900 m NW (by air) of
entrance to Valle de los Fantasmas
on MX70, Sierra de Alvarez
MK335417 – 22.071 -100.614 2297
C. multidentatus IBH28193
San Luis Potosí: 26.2 km E (by rd)
of the center of Ciudad del Maíz
on MX80, at turnoff to RMO Las
Antenas San Luis Potosí
MK335412 – 22.487 -99.473 1223
C. multidentatus IBH30104
San Luis Potosí: 26.2 km E (by rd)
of the center of Ciudad del Maíz
on MX80, at turnoff to RMO Las
Antenas San Luis Potosí
MK335414 – 22.487 -99.473 1223
C. multidentatus IBH28194
San Luis Potosí: 26.2 km E (by
rd) of center of Ciudad del Maíz
on MX80, at turnoff to RMO Las
Antenas San Luis Potosí
MK335413 – 22.487 -99.473 1223
C. multidentatus IBH23111 San Luis Potosí: Rancho
Borbortón MK335415 – 22.116 -100.601 2098
C. nubilus IBH31045 Veracruz: 8.2 km W of Xico,
Coxmatla MK335405 MK335251 19.433 -97.080 2023
C. nubilus IBH31046 Veracruz: 8.2 km W of Xico,
Coxmatla MK335399 MK335245 19.433 -97.080 2023
C. nubilus IBH31048 Veracruz: 8.2 km W of Xico,
Coxmatla MK335402 MK335248 19.433 -97.080 2023
C. nubilus IBH31049 Veracruz: 8.2 km W of Xico,
Coxmatla MK335403 MK335249 19.433 -97.080 2023
C. nubilus IBH31050 Veracruz: 8.2 km W of Xico,
Coxmatla MK335400 MK335246 19.433 -97.080 2023
C. nubilus IBH31052 Veracruz: 8.2 km W of Xico,
Coxmatla MK335401 MK335247 19.433 -97.080 2023
C. nubilus IBH31053 Veracruz: 4 km W of Xico, road to
Xico Viejo MK335404 MK335250 19.439 -97.043 1583
C. nubilus CARIE0739 Veracruz: Bosque Banderilla,
Banderilla MK335411 – 19.586 -96.946 1580
C. nubilus CARIE0740 Veracruz: Bosque Rancho Viejo,
Tlalnehuayocan MK335406 MK335252 19.521 -96.984 1520
C. nubilus CARIE1162 Veracruz: Rancho la Mesa,
Banderilla KP886894 – 19.582 -96.945 1577
C. orculus IBH30765 Estado de México: Amecameca,
road to Popocatepetl volcano MK335391 MK335237 19.072 -98.711 2800*
C. orculus IBH30746 Estado de México: Amecameca,
road to Popocatepetl volcano MK335392 MK335238 19.072 -98.711 2800*
Table 1 (continued). Voucher information, localities, GenBank accessions, coordinates and elevation data from specimens used for
phylogenetic analyses. Collection abbreviations: CARIE, Colección de Referencia de Anbios y Reptiles del Instituto de Ecología,
A.C.; IBH, Colección Nacional de Anbios y Reptiles, Instituto de Biología, UNAM; MVZ, Museum of Vertebrate Zoology, University
of California, Berkeley, California, USA. NOTE: Asterisks indicate data inferred indirectly from the available information.
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Amphib. Reptile Conserv. December 2018 | Volume 12 | Number 2 | e167
Two new Chiropterotriton from central Veracruz, Mexico
Species Voucher
Number Locality 16S GenBank COI
GenBank Latitude Longitude Elevation
m asl
C. priscus IBH22367 Nuevo León: 19.4 Km W 18 de
Marzo, Cerro Potosi MK335380 MK335226 24.891 -100.208 2600
C. terrestris GP215 Hidalgo: 5.3 km N hwy 105 at
Zacualtipan. MK335389 MK335235 20.674 -98.696 1860
C. sp. CMVZ163635 Veracruz: 3.2 km S Puerto del Aire AY522453 – 18.670 -97.338 2406*
C. sp. CIBH 14317 Veracruz: 3.0 km S Puerto del Aire AY522454 – 18.670 -97.338 2400
C. sp. FIBH30112
Puebla: 7.1 km N (by rd) of the
center of Cuetzalan on road to
Yohualichán
MK335410 MK335255 20.050 -97.500 965
C. sp. FMVZ178706 Puebla: 3.9 km S Xicotepec de
Juárez AY522477 – 20.246 -97.854 1135
C. sp. FMVZ200723
Puebla: Xicotepec de Juárez, 3.3
km S of Hotel M. Ranchito on
Mexico Hwy. 130, 21 km E on
road to La Union
AY522478 – 20.246 -97.854 1152
C. sp. FMVZ178707 Puebla: 3.9 km S Xicotepec de
Juárez AY522479 – 20.246 -97.854 1135
C. sp. GMVZ178700 Puebla: 4 km S Chignahuapan AY522480 – 19.801 -98.030 2750
C. sp. GMVZ178703 Puebla: 4 km S Chignahuapan AY522481 – 19.801 -98.030 2750
C. sp. HIBH22568 Veracruz: Microondas las Lajas KP886893 – 19.593 -97.095 3127
C. sp. IMVZ201387 Puebla: Santa Cruz de
Texmalaquilla AY522488 – 18.942 -97.287 3100*
C. sp. IMVZ201389 Puebla: Santa Cruz de
Texmalaquilla AY522487 – 18.942 -97.287 3100*
C. sp. J IBH30099 Oaxaca: San Bernardo, 4.8 km SW
(by rd) of La Esperanza on MX177 MK335409 MK335254 18.015 -96.660 1672
C. sp. KMVZ173231 Oaxaca: Cerro San Felipe AY522493 – 17.160 -96.661 3010*
Aquiloeurycea
cephalica IBH30253
Hidalgo: 1.0 km S (by rd) of La
Encarnación on road to MX85,
Parque Nacional los Mármoles
MK335378 – 20.866 -99.219 2407
Parvimolge
townsendi IBH31063 Veracruz: 4 km W Xico, road to
Xico Viejo MK335379 MK335225 19.439 -97.043 1583
Table 1 (continued). Voucher information, localities, GenBank accessions, coordinates and elevation data from specimens used for
phylogenetic analyses. Collection abbreviations: CARIE, Colección de Referencia de Anbios y Reptiles del Instituto de Ecología,
A.C.; IBH, Colección Nacional de Anbios y Reptiles, Instituto de Biología, UNAM; MVZ, Museum of Vertebrate Zoology, University
of California, Berkeley, California, USA. NOTE: Asterisks indicate data inferred indirectly from the available information.
Table 2. Sequence divergence with Kimura two-parameter distances for 16S (left) and COI (right).
C. aureus C. chiropterus C. lavae C. nubilus C. sp. C C. sp. F C. sp. G C. sp. H C. sp. I C. sp. J C. sp. K
C. aureus – 5%/10% 10%/13% 4%/7% 9%/– 6%/11% 6%/– 9%/– 9%/– 5%/8% 5%/–
C. chiropterus 5%/10% – 7%/16% 3%/10% 7%/– 6%/13% 7%/– 7%/– 7%/– 1%/5% 6%/–
C. lavae 10%/13% 7%/16% – 8%/15% 1%/– 9%/13% 7%/– 1%/– 1%/– 8%/15% 7%/–
C. nubilus 4%/7% 3%10% 8%/15% – 7%/– 5%/12% 7%/– 8%/– 7%/– 3%/8% 6%/–
C. sp. C 9%/– 7%/– 1%/– 7%/– – 9%/– 7%/– 2%/– 1%/– 7%/– 7%/–
C. sp. F 6%/11% 6%/13% 9%/13% 5%/12% 9%/– – 8%/– 9%/– 8%/– 7%/12% 7%/–
C. sp. G 6%/– 7%/– 7%/– 7%/– 7%/– 8%/– – 7%/– 7%/– 7%/– 4%/–
C. sp. H 9%/– 7%/– 1%/– 8%/– 2%/– 9%/– 7%/– – 1%/– 8%/– 7%/–
C. sp. I 9%/– 7%/– 1%/– 7%/– 1%/– 8%/– 7%/– 1%/– – 7%/– 6%/–
C. sp. J 5%/8% 1%/5% 8%/15% 3%/8% 7%/– 7%/12% 7%/– 8%/– 7%/– – 7%/–
C. sp. K 5%/– 6%/– 7%/– 6%/– 7%/– 7%/– 4%/– 7%/– 6%/– 7%/– –
42
Amphib. Reptile Conserv. December 2018 | Volume 12 | Number 2 | e167
García-Castillo et al.
logenetic methods were ran through the CIPRES data
portal (Miller et al. 2010).
Morphological Analyses
Analysis compared new taxa morphology with phyloge-
netically and geographically related species (see Results).
Further comparisons included measurements taken from
seven adult specimens of the two new species, twelve C.
chiropterus, nineteen C. lavae, twenty C. orculus (Cope),
and published measurements of C. dimidiatus (Taylor)
from García-Castillo et al. (2017) [Table 3; Appendix 1].
Male and female comparisons were completed separately
due to sexual dimorphic differences.
Basic characters and measurements follow the for-
mat used by Lynch and Wake (1989): snout-vent length
(SVL), tail length (TL), axilla-groin distance (AX), fore-
limb length (FLL), hind limb length (HLL), snout to gular
fold distance (head length, HL), head width at the angle
of the jaw (HW), head depth (HD), shoulder width (SW),
internarial distance (IN), and right foot width (FW). In ad-
dition, twelve measurements were taken from holotypes:
anterior rim of orbit to snout, eyelid length, eyelid width,
horizontal orbital diameter, interorbital distance, length of
third (longest) toe, length of fth toe, projection of snout
beyond mandible, snout to anterior angle of the vent,
snout to forelimb, tail depth at the base, and tail width at
the base (All measurements are given in mm, except tooth
counts and adpressed limbs). Maxillary plus premaxillary
(MT+PMT) and vomerine teeth (VT) were recorded for
all specimens. Finally, measurements were documented
for the limb interval (LI) as the number of costal folds
between adpressed limbs (positive values as grooves and
negative values as the overlap between limbs). Descrip-
tions are based on the color catalogue from Köhler (2012).
Results
Mitochondrial DNA (mtDNA) dataset included the 16
described species of Chiropterotriton plus seven previ-
ously proposed candidate species (Darda 1994; Parra-Olea
2003). Obtained were a 1,477-bp matrix for ribosomal
12S, tRNA, and 16S genes (including gaps) and 658 bp
for COI gene. The estimated substitution models were as
follows: GTR+G for 12S, tRNA, 16S, the 3rd codon posi-
tion of COI, K80+G for the 1st codon position of COI, and
HKY+1 for the 2nd codon position of COI. Our concat-
enated phylogeny has a similar topology as shown in pre-
vious studies (Darda 1994; Parra-Olea 2003; Rovito and
Parra-Olea 2015; García-Castillo et al. 2017), which show
two main groups, a northern and southern species groups.
(Fig. 2). The northern assemblages have a distribution
from central Mexico in Hidalgo to Nuevo Leon, the most
northern limit for the genus, and include the following
species: C. terrestris (Taylor), C. chico García-Castillo
et al., C. infernalis Rovito and Parra-Olea, C. chondro-
stega (Taylor), C. mosaueri (Woodall), C. priscus Rabb,
C. miquihuanus Campbell et al., C. magnipes Rabb, C.
cracens Rabb, C. cieloensis Rovito and Parra-Olea, C.
arboreus (Taylor), and C. multidentatus (Taylor). Where-
as, the southern assemblages (PP = 1.0, BS = 100) occur
from central Mexico in Hidalgo to the south in Oaxaca
and only have four described species: C. dimidiatus,
C. orculus, C. lavae, and C. chiropterus. However, this
clade includes seven previously proposed candidate spe-
cies: C. sp. G, C. sp. K, C. sp. H, C. sp. I, C. sp. C, C. sp.
F, and C. sp. J (Fig. 2). Results support the distinctive-
ness of two additional taxa genetically divergent from all
others and correspond to specimens collected in central
Veracruz. One occurs in only one locality (Atzalan) on
the western side of Sierra de Chiconquiaco, but the sec-
ond was found in six localities (Coxmatla, Xico, Ban-
derilla, Cinco Palos, La Cortadura, and Tlalnehuayocan)
on the eastern slope of Cofre de Perote (Fig. 2). There
is no molecular data for the Cinco Palos and La Corta-
dura populations, but these specimens were assigned to
the new taxa according to morphological characters and
concordant geographical distributions (Fig. 1).
The new taxa are phylogenetically related to C. chi-
ropterus, C. sp. J, and C. sp. F. The genetic distance
(K2P) between specimens from Atzalan and their closely
related taxa are as follows: C. chiropterus 5% (16S) and
10% (COI), C. sp. J 5% (16S) and 8% (COI), C. sp. F 6%
(16S) and 11% (COI), and the Cofre de Perote specimens
(average for all 4 populations) 4% (16S) and 7% (COI).
The genetic distance between specimens from Cofre de
Perote (all 4 populations) and their closely related taxa
are as follows: C. chiropterus 3% (16S) and 10% (COI),
C. sp. J 3% (16S) and 8% (COI), C. sp. F 5% (16S) and
12% (COI), and Atzalan specimens 4% (16S) and 7%
(COI) [Table 2]. According to the phylogenetic analysis,
C. chiropterus and C. sp. J are sister taxa (PP = 1.0, BS
= 100) with 1% (16S) and 5% (COI) genetic divergence
between them, and a sister clade to specimens from Co-
fre de Perote, although with little support (PP = 0.61, BS
= 40). These three taxa are the sister group to specimens
from Atzalan (PP = 1.0, BS = 100). Chiropterotriton sp.
F is the sister taxon of all the aforementioned taxa (PP =
1.0, BS = 100) [Fig. 2]. Given the molecular evidence
and morphological comparisons, proposed herein are the
Atzalan and Cofre de Perote populations as new species.
Systematics
Chiropterotriton aureus sp. nov.
urn:lsid:zoobank.org:act:A288BF9A-589E-42D5-8675-2AA9E6E55865
Atzalan Golden Salamander
Salamandra Dorada de Atzalan
(Figs. 3A, 4A, and 4B)
Holotype. IBH 31042, an adult male from Atzalan, Vera-
cruz, 6.5 km N from Atzalan, ejido de desarrollo urbano
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Amphib. Reptile Conserv. December 2018 | Volume 12 | Number 2 | e167
Quetzalcóatl, Mexico, 1,249 meters (m) above sea level
(asl), 19.843138N, 97.231194W. Collected on 11 July
2016 by Ángel F. Soto-Pozos, M. Delia Basanta, Omar
Becerra-Soria, and Mirna G. García-Castillo.
Paratypes. Three specimens from Atzalan, Veracruz,
Mexico. All females: IBH 31041, 31043–44, 6.5 km N
from Atzalan, ejido de desarrollo urbano Quetzalcóatl,
Atzalan, Veracruz, Mexico.
Referred specimens. IBH 31040, 6.5 km N from At-
zalan, ejido de desarrollo urbano Quetzalcóatl, Atzalan,
Veracruz, Mexico.
Diagnosis. A plethodontid salamander assigned to the
genus Chiropterotriton due to its small size, slender
body, shape of hand and feet digits (relatively long outer
digit), relatively long tail, presence of sublingual fold,
and based on mtDNA sequence data. Phylogenetically
related to C. nubilus, C. chiropterus, C. sp. F, and C. sp. J
(Fig. 2). Chiropterotriton aureus differs from C. nubilus
in being shorter (SVL 28.5 in one male, mean 26.8 in fe-
males of C. aureus vs. 29.4 in one male, 30.5 in females
of C. nubilus) with a shorter head (HL 6.4 in one male,
mean 6.0 in females of C. aureus vs. 6.6 in one male,
7.4 in females of C. nubilus), narrower head in females
(mean HW 3.6 in females of C. aureus vs. 4.4 in females
of C. nubilus), relatively shorter limbs in females (mean
LI 2.3 in females of C. aureus vs. 1.5 in females of C. nu-
bilus), and smaller feet (FW 2.4 in one male, mean 1.8 in
females of C. aureus vs. 2.6 in one male, 2.3 in females
of C. nubilus). Digits are narrower at the tip and with
Two new Chiropterotriton from central Veracruz, Mexico
Fig. 2. Bayesian analysis tree for mitochondrial loci. Numbers above branches correspond to posterior probability, and numbers
below branches are bootstrap values from maximum likelihood analysis. Asterisks indicate signicant support (posterior probability,
PP > 0.95 and bootstrap, BS > 70) in both analyses. The topology is grouped into northern and southern assemblages according to
species distributions.
44
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García-Castillo et al.
Table 3. Mean ± standard deviation (above) and range (below) of morphometric variables from males and females of C. aureus,
C. chiropterus, C. dimidiatus, C. lavae, C. nubilus, and C. orculus. Measurements are given in millimeters (mm), except TL/SLV
(proportional value), LI (limb interval), and tooth counts. NOTE: Data taken from García-Castillo et al. 2017.
Males C. aureus
N=1
C. chiropterus
N=8
C. dimidiatus*
N=15
C. lavae
N=10
C. nubilus
N=1
C. orculus
N=10
SVL 28.5 37.5±0.98
(36.1–38.8)
24.7±0.97
(23.3–26.7)
32.4±0.92
(31.0–33.8) 29.4 35.9±1.36
(33.6–38.9)
TL 36.5
47.3±3.24
(42.6–52.3)
N=7
22.0±1.72
(18.5–24.1)
38.5±2.11
(36.2–42.3) 40.2
36.6±2.87
(33.3–41.0)
N=9
TL/SLV 1.28
1.25±0.08
(1.13–1.38)
N=7
0.89±0.08
(0.7–1.0)
1.2±0.06
(1.11–1.27) 1.37
1.02±0.08
(0.86–1.15)
N=9
AX 15.5 19.6±0.59
(18.7–20.8)
13.1±0.75
(11.7–14.0)
16.2±0.87
(14.7–17.4) 15.9 18.6±1.04
(17.1–20.5)
FLL 5.9 9.1±0.44
(8.2–9.5)
4.5±0.34
(3.8–5.0)
9.3±0.59
(8.4–10.2) 6.4 8.9±0.65
(7.4–9.6)
HLL 7.5 10.3±0.47
(9.5–10.8)
5.2±0.34
(4.9–5.9)
9.9±0.72
(8.5–11.0) 7.1 9.3±0.64
(8.2–10.4)
HL 6.4 8.1±0.41
(7.7–8.9)
5.3±0.32
(4.8–5.8)
7.5±0.33
(7.2–8.1) 6.6 7.4±0.47
(6.7–8.1)
HW 4.0 5.6±0.22
(5.4–6.0)
3.5±0.21
(3.0–3.7)
4.9±0.31
(4.5–5.6) 4.0 5.0±0.35
(4.5–5.5)
HD 1.8 2.7±0.07
(2.6–2.8)
1.8±0.09
(1.7–2.0)
2.5±0.19
(2.3–2.9) 2.0 2.4±0.13
(2.2–2.7)
SW 3.4 4.0±0.35
(3.2–4.4)
2.9±0.29
(2.3–3.6)
3.1±0.30
(2.6–3.5) 3.4 3.4±0.30
(3.1–4.0)
IN 1.0 1.9±0.13
(1.7–2.1)
1.2±0.08
(1.0–1.3)
2.3±0.20
(1.9–2.5) 1.2 2.2±0.19
(1.9–2.5)
FW 2.4 3.7±0.33
(3.3–4.4)
1.7±0.20
(1.4–2.1)
3.7±0.39
(3.1–4.2) 2.6 3.2±0.22
(2.8–3.5)
LI 2.0 0.3±0.53
(-0.5–1.0)
3.9±0.35
(3.0–4.0)
-0.6±0.52
(-1.0–0.0) 2.0 1.9±0.88
(0.0–3.0)
PMT+MT 14.0 16.3±3.69
(11.0–21.0)
9.4±2.59
(5.0–14.0)
10.3±3.62
(3.0–15.0) 20.0 10.9±2.47
(7.0–14.0)
VT 15.0 10.6±1.06
(9.0–12.0)
5.7±1.35
(4.0–8.0)
8.9±1.10
(7.0–10.0) 10.0 8.6±1.90
(5.0–11.0)
Females C. aureus
N=3
C. chiropterus
N=4
C. dimidiatus*
N=15
C. lavae
N=9
C. nubilus
N=2
C. orculus
N=10
SVL 26.8±0.86
(26.0–27.7)
33.5±2.55
(30.7–36.7)
25.8±1.56
(23.1–29.1)
31.6±2.46
(27.9–34.9)
30.5±3.89
(27.7–33.2)
39.0±2.70
(34.9–43.0)
TL 31.1±1.41
(30.1–32.1)
39.5±2.35
(37.0–42.6)
22.4±1.85
(19.9–25.2)
32.5±4.89
(25.7–40.1)
34.3±5.16
(30.6–37.9)
39.2±3.64
(34.7–44.7)
N=9
TL/SLV 1.16±0.00
(1.16–1.16)
1.19±0.12
(1.01–1.26)
0.87±0.06
(0.7–1.0)
1.0±0.10
(0.85–1.15)
1.12±0.03
(1.10–1.14)
1.02±0.08
(0.87–1.12)
N=9
AX 15.0±0.49
(14.7–15.6)
18.5±2.27
(15.4–20.7)
14.8±1.24
(12.6–17.3)
16.3±1.68
(13.9–18.5)
16.4±2.69
(14.5–18.3)
21.2±1.58
(18.6–23.2)
FLL 5.3±0.42
(4.8–5.6)
7.8±0.48
(7.1–8.2)
4.3±0.43
(3.8–5.1)
8.2±0.72
(7.1–9.5)
6.5±0.28
(6.3–6.7)
8.9±0.63
(7.6–10.0)
HLL 6.7±0.35
(6.4–7.1)
8.9±0.31
(8.4–9.1)
5.0±0.47
(4.4–6.1)
8.8±0.73
(7.5–9.8)
7.2±0.14
(7.1–7.3)
9.5±0.57
(8.6–10.4)
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Two new Chiropterotriton from central Veracruz, Mexico
Females C. aureus
N=3
C. chiropterus
N=4
C. dimidiatus*
N=15
C. lavae
N=9
C. nubilus
N=2
C. orculus
N=10
HL 6.0±0.31
(5.7–6.3)
7.3±0.56
(6.5–7.8)
5.1±0.34
(4.5–5.6)
7.0±0.42
(6.3–7.6)
7.4±0.99
(6.7–8.1)
8.0±0.52
(7.4–8.9)
HW 3.6±0.10
(3.5–3.7)
4.8±0.21
(4.5–5.0)
3.5±0.25
(3.2–4.0)
4.7±0.30
(4.1–5.0)
4.4±0.14
(4.3–4.5)
5.2±0.29
(4.7–5.6)
HD 1.8±0.02
(1.8–1.8)
2.5±0.14
(2.3–2.6)
2.0±0.20
(1.7–2.2)
2.3±0.18
(2.1–2.7)
2.0±0.07
(1.9–2.0)
2.6±0.32
(2.3–3.4)
SW 3.1±0.17
(3.0–3.3)
3.6±0.38
(3.3–4.1)
3.1±0.26
(2.8–3.5)
3.3±0.33
(2.8–3.8)
3.3±0.28
(3.1–3.5)
3.9±0.46
(3.4–4.8)
IN 1.1±0.06
(1.0–1.1)
1.7±0.38
(1.4–2.1)
1.3±0.15
(1.1–1.7)
1.8±0.13
(1.6–2.0)
1.2±0.02
(1.2–1.2)
2.1±0.25
(1.7–2.5)
FW 1.8±0.21
(1.6–2.0)
3.1±0.37
(2.6–3.5)
1.8±0.26
(1.3–2.2)
3.3±0.27
(3.0–3.7)
2.3±0.57
(1.9–2.7)
3.4±0.37
(2.6–3.9)
LI 2.3±0.58
(2.0–3.0)
2.0±0.41
(1.5–2.5)
4.9±0.26
(4.0–5.0)
0.6±0.73
(0.0–2.0)
1.5±0.71
(1.0–2.0)
2.9±0.32
(2.0–3.0)
PMT+MT 44.7±2.08
(43.0–47.0)
54.3±8.08
(47.0–63.0)
34.4±4.12
(27.0–41.0)
28.0±8.19
(17.0–45.0)
48.0±2.83
(46.0–50.0)
35.9±4.46
(29.0–43.0)
VT 12.3±1.53
(11.0–14.0)
12.5±2.38
(10.0–15.0)
8.3±1.35
(6.0–11.0)
11.4±2.30
(8.0–15.0)
13.5±0.71
(13.0–14.0)
12.0±1.94
(9.0–15.0)
Table 3 (continued). Mean ± standard deviation (above) and range (below) of morphometric variables from males and females of
C. aureus, C. chiropterus, C. dimidiatus, C. lavae, C. nubilus, and C. orculus. Measurements are given in millimeters (mm), except
TL/SLV (proportional value), LI (limb interval), and tooth counts. NOTE: Data taken from García-Castillo et al. 2017.
less webbing (just onto the penultimate phalanx) than C.
nubilus (Fig. 3).
Chiropterotriton aureus differs from C. chiropterus
in being shorter (SVL 28.5 in one male, mean 26.8 in
females of C. aureus vs. 37.5 in males, 33.5 in females
of C. chiropterus), relatively shorter limbs in males (LI
2.0 in one male of C. aureus vs. 0.3 in males of C. chi-
ropterus), shorter head (HL 6.4 in one male, mean 6.0 in
females of C. aureus vs. 8.1 in males, 7.3 in females of
C. chiropterus), narrower head (HW 4.0 in one male, 3.6
in females of C. aureus vs. 5.6 in males, 4.8 in females
of C. chiropterus), and smaller feet (FW 2.4 in one male,
mean 1.8 in females of C. aureus vs. 3.7 in males, 3.1 in
females of C. chiropterus). Chiropterotriton aureus has
narrower digits at the tip and smaller feet and hands than
C. chiropterus (Fig. 3).
Chiropterotriton aureus differs from its geographi-
cally close species C. lavae by being shorter (SVL 28.5
in one male, mean 26.8 in females of C. aureus vs. 32.4
in males, 31.6 in females of C. lavae), shorter head (HL
6.4 in one male, mean 6.0 in females of C. aureus vs. 7.5
in males, 7.0 in females of C. lavae), narrower head (HW
4.0 in one male, 3.6 in females of C. aureus vs. 4.9 in
males, 4.7 in females of C. lavae), shorter limbs (LI 2.0
in one male, mean 2.3 in females of C. aureus vs. -0.6 in
males, 0.6 in females of C. lavae), and smaller feet (FW
2.4 in one male, mean 1.8 in females of C. aureus vs. 3.7
in males, 3.3 in females of C. lavae) with less webbing in
C. aureus than in C. lavae (Fig. 3).
Chiropterotriton aureus differs from C. orculus by be-
ing shorter (SVL 28.5 in one male, mean 26.8 in females
of C. aureus vs. 35.9 in males, 39.0 in females of C. orcu-
lus), longer tail (TL/SVL 1.28 in one male, mean 1.16 in
females of C. aureus vs. 1.02 in both males and females
of C. orculus), relatively larger limbs in females (mean
LI 2.3 in females of C. aureus vs. 2.9 in females of C.
orculus), shorter head (HL 6.4 in one male, mean 6.0 in
females of C. aureus vs. 7.4 in males, 8.0 in females of
C. orculus), narrower head (HW 4.0 in one male, 3.6 in
females of C. aureus vs. 5.0 in males, 5.2 in females of C.
orculus), and smaller feet (FW 2.4 in one male, mean 1.8
in females of C. aureus vs. 3.2 in males, 3.4 in females
of C. orculus).
Chiropterotriton aureus differs from C. dimidiatus in
being longer (SVL 28.5 in one male, mean 26.8 in fe-
males of C. aureus vs. 24.7 in males, 25.8 in females
of C. dimidiatus), longer tail (TL/SVL 1.28 in one male,
mean 1.16 in females of C. aureus vs. 0.89 in males, 0.87
in females of C. dimidiatus), longer head (HL 6.4 in one
male, mean 6.0 in females of C. aureus vs. 5.3 in males,
5.1 in females of C. dimidiatus), longer limbs (LI 2.0 in
one male, mean 2.3 in females of C. aureus vs. 3.9 in
males, 4.9 in females of C. dimidiatus), and more max-
illary teeth (PMT+MT 14.0 in one male, mean 44.7 in
females of C. aureus vs. 9.4 in males, 34.4 in females of
C. dimidiatus).
Chiropterotriton aureus is phylogenetically related
to members of the southern assemblages (Fig. 2), which
includes seven undescribed taxa previously suggested
by allozyme data (Darda 1994) and mtDNA (Parra-Olea
2003). Chiropterotriton aureus differs genetically from
its close relatives as follows: 6% (16S) and 11% (COI) to
C. sp. F; 5% (16S) and 8% (COI) to C. sp. J; 9% (16S) to
C. sp. H, C. sp. I, and C. sp. C; 6% (16S) to C. sp. G; and
5% (16S) to C. sp. K (Table 2).
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Chiropterotriton aureus differs from other members
of Chiropterotriton by its smaller body size (SVL 28.5
in one male, mean 26.8 in females), while C. arboreus
(mean SVL 33.4 in males, 32.2 in females; García-
Castillo et al. 2017), C. cieloensis (mean SVL 32.6 in
males, 31.1 in females; Rovito and Parra-Olea 2015), C.
chico (mean SVL 38.4 in males, 39.3 in females; Gar-
cía-Castillo et al. 2017), C. infernalis (mean SVL 36.4
in males, 29.7 in one female; Rovito and Parra-Olea
2015), C. magnipes (mean SVL 46.8 in males, 57.5 in
females; Rabb 1965), C. miquihuanus (mean SVL 33.3 in
males, 36.5 in females; Rovito and Parra-Olea 2015), C.
mosaueri (mean SVL 42.8 in males; Woodall 1941), C.
multidentatus (mean SVL 33.6 in males, 34.0 in females;
Rovito and Parra-Olea 2015), and C. priscus (mean SVL
38.5 in males, 41.8 in females; Rovito and Parra-Olea
2015). However, this species is longer than C. chondro-
stega (mean SVL 23.1 in males, 25.4 in females; Gar-
cía-Castillo et al. 2017), C. cracens (mean SVL 25.7 in
males, 27.4 in females; Rovito and Parra-Olea 2015), and
C. terrestris (mean SVL 24.2 in males, 23.0 in females;
García-Castillo et al. 2017). Chiropterotriton aureus has
smaller feet (FW 2.4 in one male, mean 1.8 in females)
than C. arboreus (mean FW 3.4 in males, 3.5 in females;
García-Castillo et al. 2017), C. cieloensis (mean FW 3.2
in males, 3.1 in females; Rovito and Parra-Olea 2015),
C. chico (mean FW 4.1 in males, 4.2 in females; García-
Castillo et al. 2017), C. infernalis (mean FW 4.2 in males,
2.8 in one female; Rovito and Parra-Olea 2015), C. mul-
tidentatus (mean FW 3.6 in males, 3.5 in females; Ro-
vito and Parra-Olea 2015), and C. priscus (mean FW 3.2
in males, 3.5 in females; Rovito and Parra-Olea 2015).
Chiropterotriton aureus has shorter limbs (LI 2.0 in one
male, mean 2.3 in females) than C. arboreus (mean LI
0.2 in males, 1.0 in females; García-Castillo et al. 2017),
C. cieloensis (mean LI -0.2 in males, 0.1 in females; Ro-
vito and Parra-Olea 2015), C. infernalis (mean LI -0.7 in
males, -0.5 in one female; Rovito and Parra-Olea 2015),
C. multidentatus (mean LI 0.1 in males, 1.0 in females;
Rovito and Parra-Olea 2015), but it has longer limbs than
C. dimidiatus (mean LI 3.8 in males, 4.9 in females; Gar-
cía-Castillo et al. 2017), C. miquihuanus (mean LI 4.2 in
males, 4.3 in females; Rovito and Parra-Olea 2015), and
C. priscus (mean LI 3.2 in males, 3.7 in females; Rovito
and Parra-Olea 2015).
Description. A small species of Chiropterotriton, mean
SVL 28.5 in one adult male (with mental gland) and 26.8
in three adult females (range 26.0–27.7). Head narrow
and moderately long (HW 4.0 in one male, mean 3.6 in
females; HL 6.4 in one male, mean 6.0 in females), HW/
SVL=14% in one male a mean of 13% in females (range
13–14), and is wider than the shoulders (SW 3.4 in one
male, mean 3.1 in females). Nostril oval shaped. Men-
tal gland in one male small and almost circular shaped.
Snout narrow and squared shaped. Eyes slightly protu-
berant. Jaw muscles are visible as grooves in the “V” be-
hind the eyes. Few maxillary teeth in one male (mean MT
10.0) but a moderately large number in females (mean
MT 38.3, range 37–40). Premaxillary teeth in one male
are not enlarged and not piercing the lip. Few vomerine
teeth in one male (VT 15.0) and females (mean VT 12.3,
range 11–14), and arranged in a well-dened line nearly
to outer margin of the choanae. Tail is longer than SVL,
TL/SVL 1.28 in one male and 1.16 in females. Limbs are
short and slender, FLL+HLL 47% of SVL in one male
and 45% in females (range 43–46). Adpressed limbs
separated by 2.0 costal folds in one male (LI 2.0) and 2.3
in females (mean LI 2.3, range 2.0–3.0). Digits slender
and narrower at the tip with moderate webbing just onto
the penultimate phalanx. Subterminal pads present. Pha-
langeal formulae: hand 1-2-3-2, foot 1-2-3-3-2. Digits in
order of increasing length: hand I-IV~II-III, foot I-V-II-
IV-III.
Coloration in life (from photos). Upper side of head
Buff (5) or Yellow Ocher (14) on Dark Carmine (61) sur-
face, Cream Yellow (82) on the tip of head and part of the
eyelids, and lateral and gular region Pale Buff (1). Dor-
sum Buff (5), Yellow Ocher (14) or Olive Horn (16) on
Pale Buff (1) surface, venter and costal sides Pale Buff
(1). Upper side of tail with progressively darker Dark
Carmine (61) with Buff (5) and Light Pratt’s Rufous (71)
Fig. 3. Head, hand, and foot morphology of preserved
specimens of Chiropterotriton species from central Veracruz.
Ventral view from right hand and foot. A) C. aureus holotype
IBH 31042, B) C. nubilus holotype IBH 31048, C) C. lavae
MVZ 106436, and D) C. chiropterus MVZ 85590.
García-Castillo et al.
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Amphib. Reptile Conserv. December 2018 | Volume 12 | Number 2 | e167
speckles, or uniform Yellow Ocher (14), or Olive Horn
(16) with Peach Red (70) speckles. Underside of tail
Pale Buff (1). Forelimbs Chamois (84), and hands nearly
translucent. Hindlimbs Buff (5), feet nearly translucent.
Underside of limbs Pale Buff (1). Iris Orange-Rufous
(56).
Coloration in alcohol. Upper side of head and dorsum
Drab (19) and underside of head Pale Horn Color (11).
Venter Pale Pinkish Buff (3) and costal region Cream
Color (12) or Cinnamon-Drab (50). Upper side of tail
Dark Drab (45), Cinnamon (225) or Hair Brown (277),
and underside of tail Buff (5) or Drab (19). Upper side
of limbs Drab (19) and underside of limbs Cream Color
(12).
Measurements of the holotype, tooth counts, and limb
intervals. SVL 28.5, TL 36.5, AX 15.5, SW 3.4, HL 6.4,
HW 4.0, HD 1.8, projection of snout beyond mandible
0.7, anterior rim of orbit to snout 1.8, interorbital dis-
tance 1.9, eyelid length 1.7, eyelid width 1.3, horizontal
orbit diameter 0.8, distance between corners of eyes 3.6,
FLL 5.9, HLL 7.5, snout to forelimb 9.2, snout to anterior
angle of vent 26.7, tail width at base 2.0, tail depth at
base 1.9, FW 2.4, length of fth toe 0.5, length of third
(longest) toe 0.9, mental gland length 1.2, and mental
gland width 1.0. Premaxillary teeth four, maxillary 4-6
(right-left sides) and vomerine 8-7 (right-left sides). Ad-
pressed limbs separated by two costal folds.
Habitat and distribution. Western side of Sierra de Chi-
conquiaco, part of the Sierra Madre Oriental in central
Veracruz. Specimens found in a cloud forest with exten-
sive deforestation (near crops and paddocks), exclusively
in arboreal bromeliads over oaks at 1,249 m asl (Figs. 5A
and 5B).
Natural History. Chiropterotriton aureus was found
exclusively in bromeliads in cloud forest around 1,200
m asl. Examined were approximately 40 bromeliads and
found only ve specimens, including four adults (one
male and three females). Sampled bromeliads were at
1.5–3.0 m from the ground and small (approximately 20–
40 cm in diameter). Sampling site was disturbed and de-
forested, but adjacent zones with similar environmental
conditions could be explored to delimit the distributional
range of this species. Species possibly sympatric with C.
aureus may be Aquiloeurycea cafetalera, Bolitoglossa
platydactyla, Isthmura gigantea, Pseudoeurycea lynchi,
and Thorius minydemus.
Etymology. Latin epithet aureus (feminine aurea, neuter
aureus) is derived from “aurum” gold + derivational suf-
x “-eus,” meaning made of gold or gold in color, which
is the featured characteristic color of the species.
Chiropterotriton nubilus sp. nov.
urn:lsid:zoobank.org:act:F785D7FD-301F-4D53-BD7F-AC680A0BB875
Cloud Forest Salamander from Cofre de Perote
Salamandra del Bosque de Niebla del Cofre de Perote
(Figs. 3B, 4C, and 4D)
Chiropterotriton sp.: Rovito et al. 2015
Holotype. IBH 31048, an adult female from Coxmatla,
Veracruz, 8.2 km W of Xico, Veracruz, Mexico, 2,023 m
asl, 19.433264N, 97.080639W. Collected 25 June 2017
by Ángel F. Soto-Pozos, Fabiola A. Herrera-Balcázar,
M. Delia Basanta, Omar Becerra-Soria, and Mirna G.
García-Castillo.
Paratypes. One male: CARIE 0739, Banderilla,
19.586667N, 96.946111W. One Female: IBH 31049,
Coxmatla, 8.2 km W of Xico.
Referred specimens. IBH 31045–46, IBH 31050–52,
Coxmatla, 8.2 km W of Xico; IBH 31047, IBH 31053 4
km W of Xico, road to Xico Viejo; CARIE 0718, La Cor-
tadura, Coatepec, 19.491389N, 97.027778W; CARIE
0740, CARIE 1269, Bosque Rancho Viejo, Tlalnehuayo-
can; CARIE 1162, Rancho La Mesa, Banderilla; CARIE
Fig. 4. Photos in life of two new species from central Veracruz.
A) C. aureus (male) holotype IBH 31042, B) C. aureus (female)
paratype IBH 31044, C) C. nubilus (male) paratype CARIE
0739, and D) C. nubilus (female) holotype IBH 31048. Photo
credit: Maria Delia Basanta (A, B, D) and J. Luis Aguilar-
López (C).
Two new Chiropterotriton from central Veracruz, Mexico
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Amphib. Reptile Conserv. December 2018 | Volume 12 | Number 2 | e167
1267, Banderilla; CARIE 1272, Cinco Palos, Coatepec,
19.5N, 97.002778W.
Diagnosis. A plethodontid salamander assigned to the
genus Chiropterotriton due to its slender body with a rel-
atively long tail, shape of hand and feet digits, presence
of sublingual fold, and based on mtDNA sequence data.
Phylogenetically related to C. aureus, C. chiropterus, C.
sp. F, and C. sp. J (Fig. 2). Chiropterotriton nubilus dif-
fers from C. aureus in females being longer (mean SVL
30.5 in females of C. nubilus vs. 26.8 in females of C.
aureus), longer tail in males (TL/SVL 1.37 in one male
of C. nubilus vs. 1.28 in one male of C. aureus), rela-
tively longer limbs in females (mean LI 1.5 in females
of C. nubilus vs. 2.3 in females of C. aureus), a longer
head (mean HL 7.4 in females of C. nubilus vs. 6.0 in
females of C. aureus), and broader head (mean HW 4.4
in females of C. nubilus vs. 3.6 in females of C. aureus).
Chiropterotriton nubilus has longer feet (mean FW 2.3
in females of C. nubilus vs. 1.8 in females of C. aureus)
with more rounded digits and slightly more webbing
(just above penultimate phalanx) than C. aureus (Fig. 3).
Chiropterotriton nubilus differs from C. chiropterus
by being shorter (SVL 29.4 in one male, mean 30.5 in
females of C. nubilus vs. 37.5 in males, 33.5 in females
of C. chiropterus), with relatively shorter limbs in males
(LI 2.0 in one male of C. nubilus vs. mean 0.3 in males
of C. chiropterus), shorter head in males (HL 6.6 in one
male of C. nubilus vs. mean 8.1 in males of C. chiropter-
us), narrower head (HW 4.0 in one male, mean 4.4 in fe-
males of C. nubilus vs. 5.6 in males, 4.8 in females of C.
chiropterus), jaw muscles less pronounced and eyes less
protuberant than C. chiropterus (Fig. 3). Chiropterotriton
nubilus has smaller feet (FW 2.6 in one male, mean 2.3
in females of C. nubilus vs. 3.7 in males, 3.1 in females
of C. chiropterus), with rounded digits, and fourth nger
of hand and fth toe of foot longer than C. chiropterus.
Likewise, C. nubilus has more webbing that covers just
above the penultimate phalanx while C. chiropterus has
webbing under the penultimate phalanx (Fig. 3).
Chiropterotriton nubilus differs from geographically
proximate species C. lavae in males being shorter (SVL
29.4 in one male of C. nubilus vs. mean 32.4 in males of
C. lavae), a longer tail (TL/SVL 1.37 in one male, mean
1.12 in females of C. nubilus vs. 1.2 in males, 1.0 in fe-
males of C. lavae), narrower head (HW 4.0 in one male,
mean 4.4 in females of C. nubilus vs. 4.9 in males, 4.7 in
females of C. lavae), relatively shorter limbs (LI 2.0 in
one male, mean 1.5 in females of C. nubilus vs. -0.6 in
males, 0.6 in females of lavae), and more maxillary teeth
Fig. 5. Microhabitat and landscape photographs for new species from central Veracruz. A) Landscape from type locality of C. aureus
(Atzalan, Veracruz), B) bromeliad from type locality of C. aureus, C) view of type locality of C. nubilus (Coxmatla, Veracruz), and
D) bromeliad from locality of C. nubilus (Xico, Veracruz). Photo credit: Mirna G. García-Castillo and Ángel F. Soto-Pozos.
García-Castillo et al.
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Amphib. Reptile Conserv. December 2018 | Volume 12 | Number 2 | e167
(PMT+MT 20.0 in one male, mean 48.0 in females of C.
nubilus vs. 10.3 in males, 28.0 in females of C. lavae). In
general, C. nubilus is morphologically similar to C. lavae
in body size and proportions (Table 3), but C. nubilus has
smaller feet (FW 2.6 in one male, mean 2.3 in females of
C. nubilus vs. 3.7 in males, 3.3 females of C. lavae) and
less webbing (Fig. 3).
Chiropterotriton nubilus differs from C. orculus in
being shorter (SVL 29.4 in one male, mean 30.5 in fe-
males of C. nubilus vs. 35.9 in males, 39.0 in females of
C. orculus), longer tail (TL/SVL 1.37 in one male, mean
1.12 in females of C. nubilus vs. 1.02 in both males and
females of C. orculus), relatively longer limbs in females
(mean LI 1.5 in females of C. nubilus vs. 2.9 in females
of C. orculus), shorter head (HL 6.6 in one male, mean
7.4 in females of C. nubilus vs. 7.4 in males, 8.0 in fe-
males of C. orculus), narrower head (HW 4.0 in one male,
mean 4.4 in females of C. nubilus vs. 5.0 in males, 5.2 in
females of C. orculus), more maxillary teeth (PMT+MT
20.0 in one male, mean 48.0 in females of C. nubilus vs.
10.9 in males, 35.9 in females of C. orculus), and smaller
feet (FW 2.6 in one male, mean 2.3 in females of C. nubi-
lus vs. 3.2 in males, 3.4 in females of C. orculus).
Chiropterotriton nubilus differs from C. dimidiatus
in being shorter (SVL 29.4 in one male, mean 30.5 in
females of C. nubilus vs. 24.7 in males, 25.8 in females
of C. dimidiatus), longer tail (TL/SVL 1.37 in one male,
mean 1.12 in females of C. nubilus vs. 0.89 in males, 0.87
in females of C. dimidiatus), longer head (HL 6.6 in one
male, mean 7.4 in females of C. nubilus vs. 5.3 in males,
5.1 in females of C. dimidiatus), broader head (HW 4.0
in one male, mean 4.4 in females of C. nubilus vs. 3.5 in
both males and females of C. dimidiatus), relatively lon-
ger limbs (LI 2.0 in one male, mean 1.5 in females of C.
nubilus vs. 3.9 in males, 4.9 in females of C. dimidiatus),
more maxillary teeth (PMT+MT 20.0 in one male, mean
48.0 in females of C. nubilus vs. 9.4 in males, 34.4 in
females of C. dimidiatus), more vomerine teeth (VT 10.0
in one male, mean 13.5 in females of C. nubilus vs. 5.7 in
males, 8.3 in females of C. dimidiatus), and longer feet
(FW 2.6 in one male, mean 2.3 in females of C. nubilus
vs. 1.7 in males, 1.8 in females of C. dimidiatus).
Chiropterotriton nubilus is related to an undescribed
taxon of the southern assemblages with genetic diver-
gences as follows: 5% (16S) and 12% (COI) to C. sp. F;
3% (16S) and 8% (COI) to C. sp. J; 8% (16S) to C. sp. H;
7% (16S) to C. sp. I, C. sp. C, and C. sp. G; and 6% (16S)
to C. sp. K (Table 2).
Chiropterotriton nubilus differs from other species of
Chiropterotriton by being shorter (SVL 29.4 in one male,
mean 30.5 in females) other than C. arboreus (mean
SVL 33.4 in males, 32.2 in females; García-Castillo et
al. 2017), C. chico (mean SVL 38.4 in males, 39.3 in fe-
males; García-Castillo et al. 2017), C. magnipes (mean
SVL 46.8 in males, 57.5 in females; Rabb 1965), C.
miquihuanus (mean SVL 33.3 in males, 36.5 in females;
Rovito and Parra-Olea 2015), C. mosaueri (mean SVL
42.8 in males; Woodall 1941), C. multidentatus (mean
SVL 33.6 in males, 34.0 in females; Rovito and Parra-
Olea 2015), and C. priscus (mean SVL 38.5 in males,
41.8 in females; Rovito and Parra-Olea 2015). Chirop-
terotriton nubilus has a longer body size than C. chon-
drostega (mean SVL 23.1 in males, 25.4 in females;
García-Castillo et al. 2017), C. cracens (mean SVL 25.7
in males, 27.4 in females; Rovito and Parra-Olea 2015),
C. dimidiatus (mean SVL 24.6 in males, 25.8 in females;
García-Castillo et al. 2017), and C. terrestris (mean SVL
24.2 in males, 23.0 in females; García-Castillo et al.
2017). Chiropterotriton nubilus has smaller feet (FW 2.6
in one male, mean 2.3 in females) other than C. arboreus
(mean FW 3.4 in males, 3.5 in females; García-Castillo et
al. 2017), C. cieloensis (mean FW 3.2 in males, 3.1 in fe-
males; Rovito and Parra-Olea, 2015), C. chico (mean FW
4.1 in males, 4.2 in females; García-Castillo et al. 2017),
C. infernalis (4.2 in males, 2.8 in one female; Rovito
and Parra-Olea, 2015), and C. priscus (mean FW 3.2 in
males, 3.5 in females; Rovito and Parra-Olea 2015). Chi-
ropterotriton nubilus has relatively shorter limbs (LI 2.0
in one male, mean 1.5 in females) other than C. arboreus
(mean LI 0.2 in males, 1.0 in females; García-Castillo et
al. 2017), C. cieloensis (mean LI -0.2 in males, 0.1 in fe-
males; Rovito and Parra-Olea 2015), C. infernalis (mean
LI -0.7 in males, -0.5 in one female; Rovito and Parra-
Olea 2015), C. multidentatus (mean LI 0.1 in males, 1.0
in females; Rovito and Parra-Olea 2015), but relatively
longer limbs than C. dimidiatus (mean LI 3.8 in males,
4.9 in females; García-Castillo et al. 2017), C. miquihua-
nus (mean LI 4.2 in males, 4.3 in females; Rovito and
Parra-Olea 2015), and C. priscus (mean LI 3.2 in males,
3.7 in females; Rovito and Parra-Olea 2015).
Description. Moderate-sized species of Chiropterotri-
ton, SVL 29.4 in one adult male and mean 30.5 in two
adult females (range 27.7–33.2). Head relatively narrow
and moderately long (HW 4.0 in one male, mean 4.4 in
females; HL 6.6 in one male, mean 7.4 in females), 14%
of HW/SVL in one male and 15% in females (range 14–
16), and wider shoulders (SW 3.4 in one male, mean 3.3
in females). Nostrils moderately sized and oval shaped.
Snout narrow and truncated. Eyes slightly protuberant.
Jaw muscles appear as a bulging mass behind the eyes
and beyond the margin of the jaw, when viewed from
above. Premaxillary teeth in one male not enlarged and
not piercing lip. Few maxillary teeth in males (MT 13.0)
but many in females (mean MT 41.5, range 40–43). Few
vomerine teeth in males (VT 10.0) and females (mean VT
13.5, range 13–14), arranged in a well-dened line nearly
to outer margin of the choanae. Tail large, mean TL/SVL
1.37, in one male and moderate, 1.12, in females (range
1.10–1.14). Limbs short and slender, FLL+HLL 46% of
SVL in one male and 45% in females (range 42–48). Ad-
pressed limbs separated by 2.0 costal folds in one male
(LI 2.0) and 1.5 in females (mean LI 1.5, range 1.0–2.0).
Digits slender with distinct terminal pads and moderate
Two new Chiropterotriton from central Veracruz, Mexico
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Amphib. Reptile Conserv. December 2018 | Volume 12 | Number 2 | e167
webbing just above the penultimate phalanx. Phalangeal
formulae: hand 1-2-3-2, foot 1-2-3-3-2. Digits in order
of increasing length: hand I-IV-II-III, foot I-V-II-IV-III.
Coloration in life (from photos). Predominating colors
on the upper side of the head and dorsum are Flesh Ocher
(57) or Salmon (58) on Sepia (286) background. Lateral
side of the head is Cream White (52), and underside of
head and venter are Cream White (52) background with
Glaucous (291) marks. Dorsum anks Glaucous (291)
on Cream White (52) surface with Smoky White (261)
stipples. Tail Flesh Ocher (57) with Sepia (286) marks
on anks and underside Perl Gray (262) with Glaucous
(291) marks. Upper side of limbs Maroon (39) with toe
tips Magenta (236) and underside of limbs Cream White
(52) surface with Glaucous (291) marks. Iris Gem Roby
(65) [Fig. 4D].
Variation of coloration in life (from photos). CARIE
0739 adult male. Upper side of head Pale Horn Color
(11) on Dark Brownish Olive (127) surface, lateral head
Cream White (52) and underside of head Pale Buff (1).
Dorsum with two stripes Pale Horn Color (11) on Sepia
(286) surface, lateral dorsum Light Lavender (201) and
underside of dorsum Pinkish White (216) with Medium
Bluish Purple (212) small dots. Upper side of tail Pale
Horn Color (11) on Sepia (286) surface and underside
of tail Pinkish White (216) with Medium Bluish Purple
(212) small dots and some Pale Horn Color (11) speck-
les. Forelimbs Cream Color (12) and hindlimbs Fawn
Color (258) with toe tips Magenta (236). Iris Light Yel-
low Ocher (13) [Fig. 4C].
Coloration in alcohol. Upper side of head Drab (19), lat-
eral Dusky Brown (285) line and underside Smoke Gray
(266) with Smoky White (261) marks. Upper side of
dorsum and tail Dark Yellow Buff (54) on Dusky Brown
(285) surface, dorsum anks Olive-Gray (265) and un-
derside of dorsum Smoke Gray (266). Underside of tail
Grayish Horn Color (268). Upper side of limbs Olive-
Brown (278) and upper side of limbs Smoke Gray (266).
Variation in alcohol preserved coloration. Three speci-
mens: one adult male (CARIE 0739) and two juvenile
(CARIE 0740, CARIE 1267). Upper side of head Cream
White (52) on Raw Umber (23) surface and underside
of head Smoky White (261). Dorsum with two stripes
Cream White (52) on Raw Umber (23) surface, anks
and underside of dorsum Smoky White (261). Upper side
of tail Cream White (52) on Raw Umber (23) surface and
underside Smoky White (261). Upper side of forelimbs
Olive Horn Color (16), hindlimbs Fawn Color (258) and
underside of limbs Smoky White (261).
Measurements of holotype, tooth counts, and limb in-
tervals. SVL 33.2, TL 37.9, AX 18.3, SW 3.5, HL 8.1,
HW 4.5, HD 2.0, projection of snout beyond mandible
0.8, anterior rim of orbit to snout 2.0, interorbital dis-
tance 3.9, eyelid length 1.9, eyelid width 1.5, horizontal
orbit diameter 0.7, distance between corners of eyes 2.5,
FLL 6.7, HLL 7.3, snout to forelimb 10.0, snout to ante-
rior angle of vent 31.4, tail width at base 2.2, tail depth
at base 2.3, FW 2.7, length of fth toe 0.5, and length
of third (longest) toe 0.8. Premaxillary teeth 23, maxil-
lary 7–20 (right-left sides) and vomerine 7–6 (right-left
sides). Adpressed limbs are separated by two costal folds.
Habitat and distribution. Eastern slopes of Cofre de Pe-
rote in central Veracruz among cloud forest between 1,520
and 2,023 m asl. Specimens found in arboreal bromeli-
ads of cloud forest fragments with low or moderate dis-
turbance of habitat. The majority of the specimens found
were juveniles so the possibility of nding them in ter-
restrial environments (under cover objects) is not rejected
(Figs. 5C and 5D). Two localities where C. nubilus occurs
are within protected areas: municipal (La Cortadura) and
the other under private ownership (Rancho Viejo).
Natural History. Chiropterotriton nubilus was exclu-
sively found in bromeliads and six localities on the east-
ern slope of Cofre de Perote. Distribution could include
a fragmented band along cloud forests from Coxmatla to
Banderilla at 1,500–2,000 m asl. Samples included three
collections in three study locations (Banderilla, La Cor-
tadura, and Rancho Viejo) for a total of nine sampling
events. Each sampling event applied 16 person-hours for
a total sampling effort of 144 person-hours. In four of the
nine sampling events collected were C. nubilus, varying
between one to three specimens per sampling event. Bro-
meliads where C. nubilus were found measured 1.5–5.0
m from the ground and were medium in size (approxi-
mately 40–60 cm in diameter). Species found in sym-
patry with C. nubilus were Aquiloeurycea cafetalera,
Parvimolge townsendi, Pseudoeurycea lynchi, and Tho-
rius pennatulus. It is conceivable that C. nubilus could
be found in sympatry with C. lavae because distributions
converge at the W slope of Cofre de Perote at 2,000 m
asl. However, C. lavae (La Joya) is found eight km away
from the nearest location (Banderilla) C. nubilus occurs.
Etymology. Latin epithet nubilus (adjective: feminine
nubile, neuter nubilum) means cloudy or rain clouds, re-
ferring to the cloud forest of Cofre de Perote where it
occurs.
Discussion
Due to recent systematic reviews, expeditions to poorly
explored areas, and recurrent eld samplings in relatively
well-studied regions, the number of described species of
bolitoglossine salamanders has increased at a slow but
steady pace in recent years (e.g., Rovito et al. 2015;
Kubicki and Arias 2016; Parra-Olea et al. 2016; García-
Castillo et al. 2017; Arias and Kubicki 2018). The Cofre
García-Castillo et al.
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Amphib. Reptile Conserv. December 2018 | Volume 12 | Number 2 | e167
de Perote area has been well studied and is notable for
its salamander richness, which now includes 20 species
representing 16% of Mexican bolitoglossines (Wake et
al. 1992; Parra-Olea et al. 2001). The description of these
two new species increases the salamander diversity in the
state of Veracruz from 37 to 39 (Parra-Olea et al. 2014),
including the recently described Isthmura corrugata
(Sandoval-Comte et al. 2017).
The number of species in the genus Chiropterotriton
has increased by approximately 50% in the last four years
(Campbell et al. 2014; Rovito and Parra-Olea 2015; Gar-
cía-Castillo et al. 2017), but phylogenetic relationships
are not fully resolved. Although the phylogeny exhibited
in this study includes a greater number of well-supported
clades (PP > 0.95, BS > 70), some relationships still lack
strong support. However, previous studies (Parra-Olea
2003; Rovito and Parra-Olea 2015; García-Castillo et
al. 2017) and results here show a well-supported clade
with species from central and southern Mexico, in which
C. dimidiatus is sister to the group. This group also in-
cludes three more described species (C. chiropterus, C.
lavae, and C. orculus) plus seven previously proposed
candidate species (Figs. 1 and 2). Within the southern as-
semblages, Chiropterotriton species form two subclades
with three sister taxa groups. The rst group includes two
terrestrial forms, sister taxa C. orculus + C. sp. G and C.
sp. K, for which only juveniles are known. The second
group includes C. lavae + C. sp. H and C. sp. I + C. sp.
C. The rst sister pair occur in geographical proximity to
Cofre de Perote but in different elevation ranges and dif-
ferent environmental conditions (one terrestrial and one
arboreal): Chiropterotriton lavae at 2,000 m asl in cloud
forest and C. sp. H at 3,000 m asl in pine forest. The
second sister pair (C. sp. C + C. sp. I) occur near Pico de
Orizaba, with C. sp. C at 2,400 m asl in cloud forest and
C. sp. I at 3,000 m asl in pine forest, again one species
being arboreal and the other terrestrial. In contrast to the
previous two groups, the third group is formed by ve
arboreal species (C. sp. F, C. aureus, C. nubilus, C. chi-
ropterus, and C. sp. J), all distributed in similar elevation
ranges (1,200 to 2,000 m asl), and similar environmental
conditions along the cloud forest from Sierra Madre Ori-
ental to Sierra de Juárez, Oaxaca. This continuous cloud
forest belt may have promoted a progressive coloniza-
tion process enabling species formation through time and
isolation and could very well explain the phylogenetic
link between the species of Veracruz and Oaxaca, a pat-
tern also seen in other bolitoglossine groups like Thorius
(Rovito et al. 2013) and Isthmura (Sandoval-Comte et
al. 2017).
The two new species of Chiropterotriton have not
been previously reported, although a sequence of C. nu-
bilus (GenBank number KP886894) was used as a rep-
resentative of Chiropterotriton in a bolitoglossine study
(Rovito et al. 2015). The discovery of these specimens
in a relatively well-studied area is reason to continue ex-
plorations, especially if localities are progressively being
deforested (Williams-Linera 2007). Likewise, salaman-
der diversity numbers are likely underestimated for cen-
tral Veracruz (C. sp. C and C. sp. H), Puebla (C. sp. F, C.
sp. G and C. sp. I.) and Oaxaca (C. sp. J and C. sp. K)
and investigations should therefore continue as species
knowledge is more completely appreciated.
Tropical salamanders are at high risk of extinction
(Rovito et al. 2009), including the genus Chiropterotri-
ton. It is imperative, now more than ever, to make the
best use of available bioresources by biobanking ge-
netic material and living tissue for current and future
uses (Hassapakis and Clark 2017; Zimkus et al. 2018).
Cryobanked genetic material has been essential for sys-
tematic and evolutionary studies of tropical salamanders,
and allowed the description of taxa thought to be extinct
(i.e., Isthmura naucampatepetl), make taxonomic rear-
rangements (Wake et al. 2012), discover cryptic taxa
(Parra-Olea et al. 2016), and propose large genus level
phylogenies (Parra-Olea et al 2004; Rovito et al. 2013)
but may now benet and contribute to best practices in
species conservation (Zimkus et al. 2018). Biobanking
for amphibian conservation may enable us to mitigate or
prevent the complete loss of species already at high risk
(e.g., Bolitoglossa jacksoni, Cryptotriton alvarezdelto-
roi) and archive these bioresources (e.g., cryopreserved
sperm, cell cultures, somatic tissue) and make them
available for present and future conservation technolo-
gies (i.e., Assisted Reproductive Technologies [ART];
Kouba et al. 2012; Kouba and Vance 2013) and method-
ologies. Those with access to specimens (e.g., eld biolo-
gists, zoo and aquarium personnel, et al.) should consider
in their research activities and grant proposals to allow
resources and time for biobanking and preserving am-
phibian genetic resources and living tissues to enhance
species conservation efforts (Zimkus et al. 2018). Final-
ly, the existence of the Genome Resource Banks (GRBs)
can alleviate other issues not related to the biology of
species but rather to pressing political (difculties of ob-
taining eld sampling due to safety) issues and economic
troubles faced by many countries worldwide.
Acknowledgements.—MGGC thanks the Posgrado
en Ciencias Biológicas program, Universidad Nacional
Autónoma de México, and CONACyT for a scholarship
grant (CVU/Becario: 413761/262662). A supporting
grant for our research was kindly provided by Programa
de Apoyo a Proyectos de Investigación e Innnovación
Tecnológica (PAPIIT-UNAM) IN203617. We also would
like to thank Laura Márquez-Valdemar and Andrea
Jimenez for assisting us with laboratory work. Special
thanks to Aldo López-Velázquez for the excellent pho-
tography instruction and the use of his equipment. We
thank Jean H. Raffaëlli, David Darda, and an anonymous
reviewer who all made valuable comments to improve
our manuscript. Finally, we thank Omar Becerra-Soria,
Fabiola A. Herrera-Balcázar, Adriana Sandoval-Comte,
and Flor Vázquez for eld assistance and the Soto-Po-
Two new Chiropterotriton from central Veracruz, Mexico
52
Amphib. Reptile Conserv. December 2018 | Volume 12 | Number 2 | e167
zos family for providing lodging in Teocelo, Veracruz.
Collection permits were provided by the Secretaría del
Medio Ambiente y Recursos Naturales (SEMARNAT):
SGPA/DGVS/00947/16, SGPA/DGVS/03038/17 and
FAUT-0303.
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Appendix 1. Specimens examined for morphological comparisons.
Chiropterotriton aureus: Mexico, Veracruz: IBH 31041–44, 6.5 km N of Atzalan, ejido de desarrollo urbano Quetzal-
cóatl.
Chiropterotriton chiropterus: Mexico, Veracruz: MVZ 85588–92, 85594, 85597–99, 85605, 85613, 85632, 1.4 miles
(mi) SW (by road), SW edge of Huatusco de Chicuellar.
Chiropterotriton lavae: Mexico, Veracruz: MVZ 106537, 106548, W edge of La Joya along Highway (Hwy). 140; MVZ
163912–13, 163915, 171873–74, 171876, 171881, 171885, 171901, 173394–95, 173398, 192788–89, 197788, La
Joya; 178685, La Joya, Mexico Hwy. 140; MVZ 200638, forest W of La Joya.
Chiropterotriton nubilus: Mexico, Veracruz: IBH 31048–49, Coxmatla, 8.2 km W of Xico. CARIE 0739, Banderilla.
Chiropterotriton orculus: Mexico, Estado de México: MVZ 76161, 138686, 138688, 138694, 138696–97, 138700,
138776–79, 138781, 138783–84, 138793, 138796–97, 138804, 200629–30, Ridge between Volcanoes Popocatepetl
and Iztaccihuatl along Mexico Hwy. 196, 16.2 km E (by road) of hwy. 115.
Two new Chiropterotriton from central Veracruz, Mexico
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Mirna Grisel García-Castillo is a Mexican Ph.D. student from Universidad Nacional Autónoma de
México in Mexico City. She began her study of salamanders in Dr. Parra´s Lab in 2011 where she
received a Master’s degree focused on molecular phylogeny of the genus Chiropterotriton. Since that
time, she’s worked on molecular systematics, taxonomy, morphology, biogeography, and evolution of
this group as part of her doctoral thesis she is currently nishing.
Angel Fernando Soto-Pozos received a Master’s degree in Biological Sciences in 2018 from the
Instituto de Biología, Universidad Nacional Autónoma de México. His research interests include edge
effect in communities of amphibians and reptiles, diversity, conservation of endangered species of
salamanders in fragmented forest from Veracruz State, and the historic role of diseases in salamander
declines.
José Luis Aguilar López received his Master´s degree in 2010 from the Instituto de Ecología A.C.
and is currently a doctoral student at the same institution. His research interests include diversity
patterns, taxonomy, and conservation of amphibians and reptiles in tropical forests.
Eduardo Pineda is a titular researcher at the Instituto de Ecología, A.C. in Xalapa, Mexico. His
research is focused on understanding the relationship between the transformation of tropical forest
and biodiversity at different spatial scales, recognizing the importance of conserved areas and
modied habitats to maintain amphibian diversity, and assess the current situation, through eldwork,
of amphibian species in imminent danger of extinction. Currently he has several graduate students
addressing topics in ecology and/or conservation of amphibians in Mexico and Latin America.
Gabriela Parra Olea is a titular researcher at the Instituto de Biología, UNAM, Mexico. Her research
is focused on molecular systematics and conservation of Mexican amphibians. Her laboratory is formed
by students and postdocs from different countries as Mexico, Guatemala, Costa Rica, Colombia, and
Argentina, all working on research projects in systematics and taxonomy, conservation genetics, and
the impact of infectious diseases, specically chytridiomycosis, on the conservation of amphibians.
García-Castillo et al.
