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937
A new arboreal Pseudoeurycea (Caudata: Plethodonti dae)
from the Sierra de Zongolica, Veracruz, Mexico
Erasmo Cázares-Hernández1, H. David Jimeno-Sevilla1, Sean M. Rovito2, Marco Antonio López-
Luna3, Luis Canseco-Márquez4
1 Instituto Tecnológico Superior de Zongolica, Colección Cientíca del ITSZ y Herbario ZON. Km 4 Carretera a la Compañía S/N, Tepetitlanapa
CP 95005, Zongolica, Veracruz, México
2 Unidad de Genómica Avanzada (LANGEBIO), CINVESTAV, Irapuato, Guanajuato, México
3 División Académica de Ciencias Biológicas, Universidad Juárez Autónoma de Tabasco, Carretera Villahermosa-Cárdenas km 0.5
Villahermosa, Tabasco CP 86039, México
4 Laboratorio de Herpetología, Facultad de Ciencias, Universidad Nacional Autónoma de México, Distrito Federal 04510, México
http://zoobank.org/E47F65F6-92D2-4C02-AD2B-0C4B0881C519
Corresponding author: Luis Canseco Márquez (lcanseco@gmail.com)
Academic editor Uwe Fritz | Received 2 June 2022 | Accepted 6 October 2022 | Published 20 October 2022
Citation: Cázares-Hernández E, Jimeno-Sevilla HD, Rovito SM, López-Luna MA, Canseco-Márquez L (2022) A new arboreal Pseudoeurycea
(Caudata: Plethodontidae) from the Sierra de Zongolica, Veracruz, Mexico. Vertebrate Zoology 72 937–950. https://doi.org/10.3897/vz.72.e87275
Abstract
We describe a new species of plethodontid salamander of the genus Pseudoeurycea from the Sierra de Zongolica, Veracruz, Mexico.
The new species is distinguished from all other species in the genus by morphological and genetic features and by coloration. Based
on a mtDNA phylogeny, the new species belongs to the Pseudoeurycea juarezi group and is most closely related to P. rucauda from
the Sierra Mazateca in northern Oaxaca. The newly described salamander increases the number of species of plethodontid salaman-
ders from Veracruz to 43 and those recognized from Mexico to 140.
Resumen
Se describe una nueva especie de salamandra pletodóntida del género Pseudoeurycea de la Sierra de Zongolica en el Estado de
Veracruz. La nueva especie se distingue de todas las especies del género por características morfológicas y genéticas y patrón de
coloración. Con base en la logenia de ADN mitocondrial, la nueva especie pertenece al grupo P. juarezi y se encuentra más cercana-
mente relacionada con P. rucauda de la Sierra Mazateca en el norte de Oaxaca. La descripción de esta nueva salamandra, incrementa
el número de especies de salamandras pletodóntidas de Veracruz a 43 y 140 reconocidas para Mexico.
Keywords
Bolitoglossini, juarezi group, phylogeny, Pseudoeurycea jaguar sp. nov., salamander
Introduction
Mexico has the world’s seventh highest diversity of am-
phibian species (418 species) and is second only to the
USA in salamander species diversity (158 species; Am-
phibiaWeb 2022). The salamanders of the family Pletho-
dontidae are the most diverse group of amphibians in
Mexico with 139 species (AmphibiaWeb 2022), repre-
senting 33.2% of the amphibians present in the country
and including 118 species endemic to Mexico. In Mexico,
16 genera of plethodontids are present, of which 13 (and
all but three species) belong to the Neotropical tribe Boli-
Vertebrate Zoology 72, 2022, 937–950 | DOI 10.3897/vz.72.e87275
Copyright Erasmo Cázares-Hernández et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted
use, distribution, and reproduction in any medium, provided the original author and source are credited.
Cázares-Hernández et al.: New species of Pseudoeurycea from Mexico
938
toglossini (Wake 2012). Bolitoglossines have undergone
an adaptive radiation in Mesoamerica, occupying a wide
array of habitats, including arboreal habitats in cloud for-
ests (Wake 1987).
Following a taxonomic revision of the Bolitoglossini
(Rovito et al. 2015), the genus Pseudoeurycea continues
to have the highest species diversity of the 16 plethodon-
tid salamander genera present in Mexico, with 40 species
distributed from the Trans-Mexican Volcanic Belt south
through the highlands of Guerrero and Oaxaca, Mexico
to Guatemala (AmphibiaWeb 2022). Of the 40 species in
the genus, 36 are endemic to Mexico, three are distributed
in Chiapas and Guatemala, and only one (P. exspectata) is
endemic to Guatemala. Salamanders of the genus Pseu-
doeurycea are small to moderately sized with a gener-
alized morphology, with the exception of three slender,
elongate species formerly assigned to the genus Lineatri-
ton (Rovito et al. 2015).
With 42 species of plethodontid salamanders (listed
by Torres-Hernández et al. 2021), the state of Veracruz
ranks second in terms of salamander diversity after Oax-
aca with 47 species, surpassing Chiapas and Puebla (24
and 23 species, respectively). Despite having been the fo-
cus of herpetological research for decades, new species
of salamanders continue to be described from Veracruz
(García-Castillo et al. 2018; Sandoval-Comte et al. 2017;
García-Bañuelos et al. 2020; Parra-Olea et al. 2020).
In a coniferous forest (Cupressus) in the Sierra de Zon-
golica, we discovered a population of brightly colored
salamanders that diered markedly in color pattern and
external morphology from all species known from the re-
gion. Based on its morphological and molecular distinc-
tiveness, we describe this population as a new species. We
consider it to be distinct because it is diagnosable from
closely related species based on dierences in external
morphology and tooth counts, has a distinct color pattern,
and represents an independent evolutionary lineage based
on phylogenetic analysis that has a similar or greater de-
gree of genetic divergence from described species of the
juarezi and leprosa species groups compared to distances
between described species.
Materials and methods
Field work and study site
From September 2015 to July 2016 we conducted 10
sampling sessions (one three-day session per month) in
a 1.5 ha portion at the study site. Forty-eight individuals
were found, of which 40 were measured and released and
eight were collected and preserved (the maximum num-
ber allowed by the SEMARNAT collecting permit) and
deposited in the Museo de Zoología Alfonso L. Herrera,
Facultad de Ciencias, Universidad Nacional Autónoma
de México (MZFC). We used 10 arboreal traps (cylin-
drical funnel traps) placed in trees with epiphytic plants,
at a height between 2 and 4 m, with a total time of 36
trapping days. We also made random quadrant searches
for hand-captures. The study site is a coniferous forest
situated in the locality El Mirador, municipality of Tex-
huacán, Veracruz, at 2,370 m elevation.
Morphological Analysis
The description of the new species follows the format
used by Lynch and Wake (1989) for other species in the
genus Pseudoeurycea and includes the same basic charac-
ters and measurements. Larger measurements were taken
using dial calipers (to the nearest 0.1 mm); measurements
of feet, toes, and some head dimensions (e.g., additional
measurements of the holotype), as well as tooth counts,
were taken under a stereoscopic microscope. All mea-
surements are in mm and abbreviations are as follows:
Distance from snout to posterior angle of vent (snout-vent
length, SVL), distance from posterior angle of vent to tip
of tail (TL), axilla-groin distance (AX), forelimb length
(FLL), hind limb length (HLL), distance from tip of snout
to gular fold (head length, HL), width of head at angle of
jaw (HW), head depth (HD), interocular distance (IO),
distance between external nares (IN), right foot width
(RFW), length of longest (third) toe (T3) and length of
fth toe (T5). We also counted maxillary teeth (MT) and
premaxillary teeth (PMT) and vomerine teeth (VT) and
both sides were summed. Salamanders measured for our
morphological analyses are listed in Appendix 1. Color
notes are based on living specimens.
We compared the new species with all currently rec-
ognized members of the Pseudoeurycea juarezi group
(i.e., P. aurantia, P. juarezi, P. rucauda and P. saltator).
Because Neotropical salamanders typically are sexually
dimorphic, we separated males and females in the mor-
phological analyses (Tables 1, 2). We calculated mean,
standard deviation, and range for all morphological mea-
surements and tooth counts and compared the new species
to all species of the Pseudoeurycea juarezi group. Teeth
were counted by opening jaws with forceps and count-
ing ankylosed teeth (not including spaces where teeth are
missing) under a dissecting microscope. For comparisons
to other members of the juarezi group, tooth counts for
maxillary and premaxillary teeth were summed (MT+P-
MT) because the counts were not available separately for
several species.
DNA extraction and sequencing
We extracted DNA from liver tissue using a high salt pro-
tocol (Aljanabi and Martinez 1997). We amplied a frag-
ment of the large subunit ribosomal RNA (16S) gene us-
ing primers 16Sal and 16Sbr (Palumbi et al. 1991) and a
fragment of the cytochrome b (cyt b) gene using primers
MVZ15 and MVZ16 (Moritz et al. 1992). For both loci,
PCR conditions consisted of an initial denaturation step at
94°C for 2 min, followed by 35 cycles of denaturation at
94°C for 30 s, annealing at 48°C for 1 min, and extension
at 72°C for 1 min, followed by a nal extension step at
Vertebrate Zoology 72, 2022, 937–950 939
72°C for 7 min. Amplied fragments were cleaned us-
ing 1:5 diluted ExoSAP-IT (USB Corp, Cleveland, OH),
cycle sequenced using BigDye v3 terminator chemistry
(Applied Biosystems, Foster City, CA) and sequenced on
an ABI 3730 capillary sequencer.
Phylogenetic analyses
We obtained 16S and cyt b sequences for other species
of Pseudoeurycea from GenBank; voucher information
and GenBank accession numbers for all sequences used
in phylogenetic analyses are given in Table 3. Aquiloeury-
cea galeanae was used as the outgroup for all phylogenet-
ic analyses. Sequences for each gene were edited in Gene-
ious V8 (Biomatters, Aukland, NZ), aligned using Muscle
v3.8 (Edgar 2004) and concatenated. The nal alignment
consisted of 1335 bp (526 bp of 16S and 809 bp of cyt
b). Partition Finder 2.0 (Lanfear et al. 2017) was used to
select a partitioning scheme and nucleotide substitution
models using the Bayesian Information Criterion (BIC),
with data partitioned by gene and cyt b data partitioned by
codon position. The following partition strategy and sub-
stitution models were selected by Partition Finder: 16S +
cyt b codon position 1—GTR+I+G, cyt b codon position
2—HKY+I, and cyt b codon position 3—GTR+G.
We used RAxML v8.2 (Stamatakis 2014) to perform
maximum likelihood phylogenetic analysis with the par-
titioning strategy given above and a GTR+G model for all
partitions, with 1000 bootstrap replicates to assess nodal
support. We used MrBayes 3.2 (Ronquist et al. 2012) to
perform Bayesian phylogenetic analysis with three hot
and one cold chains run for 20,000,000 generations, sam-
pled every 1000 generations, with the rst 5,000 samples
discarded as burn-in. Finally, we estimated GTR genetic
distances between sequences for each gene using PAUP*
Table 1. Mean, standard deviation, and range of 13 morphological measurements and tooth counts for males of species of the Pseu-
doeurycea juarezi group. Abbreviations are dened in Materials and methods.
P. aurantia (n=4) P. jaguar sp. nov. (n=2) P. juarezi (n=11) P. rucauda (n=1) P. saltator (n=2)
SVL 42.5±1.96 (40.2–45.0) 58.3±0.49 (58.0–58.7) 45.5±2.84 (38.5–49.2) 24.2 41.1±1.68 (40.0–42.3)
TL 41.3±4.47 (36.6–47.0) 54.5±18.58 (41.4–67.6) 46.1±6.62 (29.3–52.6) 21.0 43.9±2.87 (41.9–46.0)
AX 21.3±1.44 (19.8–23.2) 29.5±0.11 (29.4–29.6) 22.2±1.57 (19.4–24.6) 11.2 21.0±0.42 (20.7–21.3)
FLL 11.4±1.14 (10.0–12.7) 15.3±4.03 (12.5–18.2) 12.6±0.81 (11.2–13.5) 7.0 11.7±0.26 (11.5–11.9)
HLL 13.2±0.86 (12.4–14.3) 16.6±3.39 (14.2–19) 14.0±0.91 (13.1–15.1) 6.7 13.2±0 (13.2)
HL 10.4±0.57 (9.9–11.2) 14.4±0.98 (13.7–15.1) 11.4±0.79 (9.4–12.5) 6.7 9.9±0.50 (9.5–10.3)
HW 6.6±0.16 (6.5–6.8) 10.0±–0.36 (9.8–10.3) 6.9±0.45 (5.9–7.8) 4.4 6.5±0.24 (6.3–6.7)
HD 3.7±0.18 (3.5–3.9) 4.5±0.28 (4.3–4.7) 3.7±0.45 (3.1–4.6) — 3.1±0.07 (3.0–3.9)
IO 2.4±0.20(2.2–2.7) 2.9±0 (2.9) 2.4 ±0.22 (2.1–2.7) — 2.4±0 (2.4)
IN 2.1±0.24 (1.8–2.4) 3.2±0.21 (3.1–3.4) 2.5±0.8 (1.9–2.9) 1.1 2.3±0.21 (2.2–2.5)
RFW 4.7±0.40 (4.2–5.2) 7.8±0.21 (7.7–8.0) 5.1±0.45 (4.3–5.6) 2.1 4.3±0.65 (3.9–4.8)
T3 2.3±0.43 (1.8–2.8) 2.9±0.70 (2.4–3.4) 2.5±0.28 (2.0–2.9) 1.1 2.5±0.23 (2.3–2.6)
T5 1.0±0.12 (0.8–1.1) 1.0±0.07 (1.0–1.1) 1.2±0.20 (0.8–1.5) 0.5 1.1±0.11 (1.1–1.2)
MT + PMT 69±3.16 (66–73) 85.5±13.43 (76–95) 67±10.68 (49–90) 31 67±20.5 (52–81)
VT 23±4.54 (17–27) 30.5±7.77(25–36) 22±2.86 (17–27) 17 26±2.82 (24–28)
Table 2. Mean, standard deviation, and range of 13 morphological measurements and tooth counts for females of species of the
Pseudoeurycea juarezi group. Abbreviations are denied in Materials and methods.
P. aurantia (n=2) P. jaguar sp. nov. (n=6) P. juarezi (n=11) P. rucauda (n=1) P. saltator (n=3)
SVL 42.8±1.84 (41.5─44.1) 59.0±11.85 (42.4–71.0) 47.5 ± 2.53 (44.0─51.3) 38.2 37.6±3.95 (33.4─41.2)
TL 38.0±0.45 (37.7─38.3) 66.1±14.22 (46.4–87.9) 43.9 ± 5.13 (33.1─52.1) 38.2 32.3±2.10 (30.8─33.8) n=2
AX 21.7±1.78 (20.5─23.0) 30.8±6.77 (22.0–39.6) 24.1 ± 1.05 (22.3─25.6) 20.7 20.6±3.06 (17.3–23.3)
FLL 11.1±0.53 (10.7─11.5) 15.1±4.66 (10.1–22.4) 12.0 ± 0.59 (11.5─12.9) 9.4 9.6±1.53 (8.5–11.4)
HLL 12.4±1.08 (11.6─13.2) 15.3±4.68 (9.9–22.3) 13.6 ± 0.71 (12.4─14.7) 9.6 10.3±1.50 (9.4–12.0)
HL 10.5±0.28 (10.3─10.7) 14.6±2.70 (11.0–17.6) 11.4 ± 0.95 (10.0─12.8) 8.5 6.9±2.77 (3.8–9.1)
HW 6.4±0.43 (6.1─6.7) 10.2±1.74 (7.8–12.0) 7.1 ±0.47 (6.2–7.9) 6.1 5.9±0.72 (5.4–6.7)
HD 3.8±0.18 (3.7─3.9) 4.5±0.76 (3.6–5.3) 4.0±0.25 (3.6–4.4) — 2.7±0.52 (2.3–3.3)
IO 2.0±0 (2.08─2.09) 3.4±0.80 (2.4–4.3) 2.4±0.38 (1.7–3.0) — 2.0±0.38 (1.8–2.5)
IN 2.0±0.18 (1.9─2.2) 3.2±0.51 (2.7–3.8) 2.1±0.13 (1.9–2.4) 1.2 1.7±0.39 (1.4–2.1)
RFW 4.2±0.18 (4.1─4.3) 7.5±2.10 (4–9.3) 5.0±0.60 (4.0–6.1) 3.9 3.8±0.58 (3.2–4.4)
T3 2.4±0.31 (2.2─2.6) 3.1±0.42 (2.7–3.9) 2.5±0.29 (2.0–3.0) 2 1.8±0.28 (1.6–2.1)
T5 1.0±0.33 (0.8─1.2) 1.0±0.30 (0.7–1.6) 1.1±0.20 (0.9–1.4) 1.3 0.8±0.15 (0.7–1)
MT+PMT 74±9.89 (67–81) 92.6±20.25 (67–116) 73±18.05 (26–96) — 73±8.71 (67–83)
VT 22±1.41 (21─23) 30.3±7.06 (18–37) 23.2±2.79 (19–26) — 22±2.64 (19–23)
Cázares-Hernández et al.: New species of Pseudoeurycea from Mexico
940
v4 (Swoord 2002). All analyses were run on the Cipres
Science Gateway (Miller et al. 2010).
Results
Systematics
Pseudoeurycea jaguar sp. nov.
http://zoobank.org/4A693A70-A098-4E24-B02E-
B602B6B10321
Figs 1, 2, 3, 4
Chresonymy. Pseudoeurycea sp. – Cázares-Hernández
et al. 2021.
Suggested English name: Jaguar Salamander.
Suggested Spanish name: Tlaconete jaguar.
Holotype. MZFC-HE 28694. An adult male from El
Mirador, Texhuacan Municipality, Veracruz, Mexico
(18°38′04.8″N, 97°03′35.0″W, 2,367 m elevation, WGS84
datum), collected by Erasmo Cázares Hernández on
5 September 2015.
Paratypes. Seven. One male: MZFC-HE 35855 (10 Sep-
tember 2015); six females: MZFC-HE 35859 (10 Sep-
tember 2015), MZFC-HE 28686 (11 September 2015),
MZFC-HE 35856–57 (14 September 2015), MZFC-HE
35858, 28685 (10 October 2015). Same locality as the
holotype.
Table 3. Voucher information and GenBank accession numbers for sequences used in phylogenetic analysis.
Species Voucher number GenBank 16S GenBank cyt b
P. ahuitzotl IBH 30211 MT303858 MT295473
P. altamontana IBH 22220 KP886861 KP900064
P. anitae MVZ 137939 AF451227 —
P. aurantia IBH 20370 KP886844 KP900048
P. brunnata MVZ 137947 AF451232 —
P. cochranae IBH 23064 KP886864 KP900067
P. conanti MVZ 146786 AF451241 —
P. exspectata MVZ 160919 AF451234 —
P. rscheini IBH 30995 MT303859 MT295469
P. jaguar sp. nov. MZFC-HE 35855 OP605487 OP617200
P. gadovii IBH 22982 KP886846 KP900050
P. goebeli CRVA1017 MT303860 MT295472
P. juarezi IBH 29718 KP886848 KP900052
P. leprosa IBH 22406 KP886866 KP900069
P. lineola IBH 29719 KP886867 KP900070
P. longicauda IBH 22247 KP886849 KP900053
P. lynchi GP160 AF451225 AF451204
P. melanomolga IBH 22784 KP886868 KP900071
P. mixcoatl IBH 14194 KP886869 KP900072
P. mixteca GP0289 AF380829 AF380790
P. mystax GP372 AF380795 AF380756
P. nigromaculata MVZ 185977 AF451238 —
P. obesa MVZ 241574 KP886870 KP900073
P. orchileucos IBH 22562 KP886858 KP900062
P. orchimelas IBH 22999 KP886860 KP900063
P. papenfussi IBH 14198 KP886850 KP900054
P. rex MVZ 263590 KP886852 KP900056
P. robertsi IBH 22232 KP886853 KP900057
P. rucauda IBH 21646 KP886871 KP900074
P. saltator IBH 22895 KP886854 KP900058
P. smithi IBH 29720 KP886855 KP900059
P. tenchalli IBH 29721 KP886856 KP900060
P. tlahcuiloh IBH 30233 MT303865 MT295474
P. unguidentis MVZ 117432 MT303866 —
P. werleri IBH 22294 KP886872 KP900075
Ixalotriton niger IBH 29715 KP886874 KP900077
I. parvus AMA2534 KP886873 KP900076
Aquiloeurycea galeanae IBH 24595 KP886847 KP900051
Vertebrate Zoology 72, 2022, 937–950 941
Diagnosis. Assigned to the genus Pseudoeurycea based
on the presence of a sublingual fold, comparatively short
fth toe compared to the fourth, limited foot webbing,
relatively large size, and mitochondrial DNA sequences.
Morphologically, we distinguish the new species from
the other salamanders that occur in the region and from
the others of the genus Pseudoeurycea based on size of
the body and tail, limb length, digit shape, shape and size
of the head, and especially by external coloration (dor-
sal and ventral coloration of head, body and tail). Pseu-
doeurycea jaguar is easily distinguished from the other
species of the genus Pseudoeurycea by its unique color
pattern (Figs 1, 4).
Based on mtDNA, this new species is closely related to
members of the P. juarezi group (sensu Canseco-Márquez
and Parra-Olea 2003; Parra-Olea et al. 2004). It is distin-
guished from species of the P. juarezi group by its larger
body size (SVL males: P. aurantia SVL 40.2–45.0, P. jag-
uar sp. nov. 58.0–58.7 mm, P. juarezi 44.0–51.3 mm, P.
rucauda 24.2 mm, P. saltator 40.0–42.3 mm; females:
P. aurantia 41.5–44.1 mm, P. jaguar sp. nov. 42.4–71.0
mm, P. juarezi 38.5–48.0 mm, P. rucauda 38.2 mm, P.
saltator 33.4–41.2 mm; Tables 1 and 2) and wider head
(HW males: P. aurantia 9.9–11.2 mm, P. jaguar sp. nov.
13.7–15.1 mm, P. juarezi 10.0–12.6 mm, P. rucauda
6.7 mm, P. saltator 9.5–10.3 mm; females: P. auran-
Figure 1. Live specimens of all members of the P. juarezi group. A P. aurantia (Peña Verde, Oaxaca), Photo by Sean Rovito; B P.
saltator (Sierra de Juárez, Oaxaca), Photo by Sean Rovito; C, D P. juarezi (Cerro Pelón, Sierra de Juárez, Oaxaca), Photos by Sean
Rovito and Luis Canseco, respectively; E P. rucauda (near Plan de Guadalupe, Oaxaca), Photo by Sean Rovito; F Holotype of P.
jaguar sp. nov. from the type locality, Photo by Erasmo Cázares.
Cázares-Hernández et al.: New species of Pseudoeurycea from Mexico
942
tia 10.3–10.7 mm, P. jaguar sp. nov. 11.0–17.6 mm, P.
juarezi 9.4–12.5 mm, P. rucauda 8.5 mm, P. saltator
3.8–9.1 mm. Females have a relatively longer tail (TL/
SVL females: P. aurantia 0.86–0.90, P. jaguar sp. nov.
0.93–1.25, P. juarezi 0.70–1.12, P. rucauda 0.99, P.
saltator 0.88–0.92). Pseudoeurycea jaguar sp. nov. has
more maxillary and premaxillary teeth (mean MT + PMT
males: P. aurantia 69, P. jaguar sp. nov. 85.5, P. juarezi
73, P. rucauda 31, P. saltator 67; females; P. aurantia
74, P. jaguar sp. nov. 92.6, P. juarezi 67, P. saltator 73)
and vomerine teeth (mean VT males: P. aurantia 25, P.
jaguar sp. nov. 30.5, P. juarezi 23.3, P. rucauda 17, P.
saltator 26; females; P. aurantia 22, P. jaguar sp. nov.
30.3, P. juarezi 22, P. saltator 22). The head of P. jaguar
is longer and wider than that of other species (HL males:
P. aurantia 9.9–11.2, P. jaguar sp. nov. 13.7–15.1 mm,
P. juarezi 10.0–12.6 mm, P. rucauda 6.7 mm, P. salta-
tor 9.5–10.3 mm; females: P. aurantia 10.3–10.7 mm, P.
jaguar sp. nov. 11.0–17.6 mm, P. juarezi 9.4–12.5 mm,
P. rucauda 8.5 mm, P. saltator 3.8–9.1 mm; HW males:
P. aurantia 6.3–6.8, P. jaguar sp. nov. 9.8–10.3 mm,
P. juarezi 6.2–7.9 mm, P. rucauda 4.4 mm, P. saltator
6.3–6.7 mm; females: P. aurantia 6.1–6.7 mm, P. jaguar
sp. nov. 7.8–12.0 mm, P. juarezi 5.9–7.8 mm, P. rucau-
da 6.1 mm, P. saltator 5.4–6.7 mm). No other species of
Pseudoeurycea from Veracruz or Puebla has such large,
extensively webbed feet and long limbs.
The new species is further distinguished from all mem-
bers of the juarezi group, as well as from all other species
of Pseudoeurycea and all salamander species from cen-
tral Veracruz, by color pattern. Pseudoeurycea jaguar has
irregular yellow mottling on the dorsum on a brown or
nearly black background. In P. aurantia the ground color
is reddish brown with bright orange blotches or mottling
present on the dorsum; these blotches coalesce on the tail
(Fig. 1A), and some individuals have small dark spots on
the tail and dorsum. Pseudoeurycea saltator has a uni-
formly dark gray-brown dorsal ground color that is in-
variably overlain by a paler yellow or golden mid-dorsal
stripe (Fig. 1B). Pseudoeurycea juarezi typically has yel-
low-brown dorsal coloration on the head and forming a
dorsal band to the tip of the tail, with scattered black spots
on the dorsum and tail (Fig. 1C); the color pattern in this
species can be variable, some specimens have small dark
spots on a yellowish background (Fig. 1D) while others
are darker brown or reddish-brown dorsally with yellow
or golden mottling on the tail. Dorsal coloration in P. ru-
cauda is orange-tan with coppery-gold highlights that
Figure 2. A Dorsal and B ventral view of the holotype of Pseudoeurycea jaguar sp. nov (MZFC-HE 28694), scale bar = 10 mm. C
Ventral view of right hand and D right foot. Scale bar = 3 mm.
Vertebrate Zoology 72, 2022, 937–950 943
are mixed with black (Fig. 1E). The ventral part of the
tail of P. jaguar is paler than the dorsal part (lead gray),
darkening towards the tip, with very small and clear
ecks evenly distributed along the median portion of the
venter. Both P. melanomolga and P. gadovii have some
version of yellowish spots on a dark background, but the
spots in these species are arranged in regular rows unlike
the mottling seen in P. jaguar. Pseudoeurycea lynchi has
greenish, rather than yellow, blotches and mottling and
tends to show less background color compared to P. jag-
uar. Pseudoeurycea nigromaculata also typically shows
less background color and often is primarily yellowish on
the tail, and P. granitum has irregular blotches and a pale
interorbital bar that are lacking in P. jaguar.
Description of the holotype. A relatively large adult
male (58.7 SVL), body slender, head relatively long and
broad (HW/SVL = 0.17), wider than body, neck region
well dened (Fig. 2A, B). Snout slightly truncate in dorsal
view and rounded in lateral view, eyes moderate in size,
slightly protruding, not exceeding margin of jaw in dor-
sal view. Nostrils small, oval. Nearly round and relatively
prominent mental gland (2.4 mm wide) (Fig. 2B). Costal
folds 13, counting one each in axilla and groin. Tail lon-
ger than body (TL/SVL = 1.15), tapering gradually along
length, slender posteriorly ending in a point. Limbs long,
overlap by 2 costal folds when appressed to side of body
(Fig. 3D). Hands and feet broad, digits long and relatively
slender, blunt with distinct subterminal pads. Hands and
feet highly webbed compared to most other members of
the genus (although only moderately webbed compared
with species of Bolitoglossa or some Chiropterotriton),
with webbing extending to the middle of the penultimate
phalanx on third toe of foot. First toe short, fth toe short
compared to fourth long, 2.1 times larger than the rst
nger. Digits in order of decreasing length: III–II–IV–I
on hands; III–IV–II–V–I on feet. Phalangeal formulae
1–2–3–2 for hands and 1–2–3–3–2 for feet (Fig. 2C, D).
Teeth numerous: maxillary teeth 92 (45/47); premaxillary
teeth 3, enlarged compared to maxillary teeth; vomerine
teeth 36, arranged in two arcs (18/18) extending beyond
the choanae. Nasolabial protuberances well developed.
Measurements of the holotype (in mm). Snout to pos-
terior angle of vent (SVL) 58.7; head width 10.3; head
length 14.8; head depth at angle of jaw 4.7; eyelid length
3.9; eyelid width 3.0; anterior rim of orbit to snout 4.9;
eye diameter 4.4; interorbital distance 3.5; snout to fore-
limb 21.0; internarial distance 3.1; intercanthal distance
4.1; nostril diameter 0.4; snout projection beyond man-
dible 1.1; snout to anterior angle of vent 55.4; axilla to
groin 29.4; tail length 67.6; tail width at base 4.3; tail
depth at base 4.7; forelimb length 18.2; hind limb length
19; hand width 5.1; foot width 8.0; length of the longest
(third) toe 2.5; length of fth toe 0.8; mental gland width
2.4; mental gland length 2.3. Tooth counts: premaxillary
3; maxillary 45/47; vomerine 18/18.
Variation and sexual dimorphism. The type series in-
cludes eight specimens, two males and six females. There
Figure 3. A Ventral view of male paratype of Pseudoeurycea jaguar sp. nov. (MZFC-HE 35855) showing the mental gland. B Ven -
tral view of right hand and C right foot of a female paratype (MZFC-HE 28685). D Male paratype male showing overlapping digits
when appressed to the side of the body (MZFC-HE 35855). Scale bar = 2 mm.
Cázares-Hernández et al.: New species of Pseudoeurycea from Mexico
944
Figure 4. Color pattern of Pseudoeurycea jaguar sp. nov. Adults: A Male holotype (MZFC-HE 28694); B, C, D Female paratypes
(MZFC-HE 35856-57, 28685, respectively; and released male (E) and female (F). Juveniles (all released). G A specimen < 22 mm
SVL; H a specimen measuring 30 mm SVL; I, J specimens measuring 33 mm SVL. Photos by Erasmo Cázares.
Vertebrate Zoology 72, 2022, 937–950 945
is marked sexual dimorphism; adult females reach a larger
size than males (SVL 42.4–71.0 mm in females vs. 58.0–
58.7 mm in males), head relatively broad (9.8–12.0 mm
in females and 9.8–10.3 mm in males) and have a more
robust body compared to males (shoulder width 7.6–10.8
in females vs 7.2–7.3 mm in males). Adult males have a
well-developed, nearly round mental gland (width 2.8 mm)
(Fig. 3A) and few premaxillary teeth (3–4 vs.13–20 in fe-
males). Hands and feet are broader in females (foot width
7.7–8 mm in males and 8.3–9.3 in females) (Fig. 3B, C).
Coloration of the holotype in life (Fig. 4A). Dorsum and
dorsal surface of head solid dark chocolate brown with
extensive yellow speckling or mottling; yellow specks
small on head, becoming larger and mottled on the dor-
sum and even larger and more continuous on tail. Sides of
head brown with yellow speckling, with the same propor-
tion of yellow toward back of the head, mouth and dorsal
surface of the head. Dorsal surface of tail same color as
dorsum, with the yellow mottling more continuous, but
reduced at tip. Sides of body dark brown above midline,
with yellow ecks (small ecks combined with larger and
elongated ecks) and slightly paler brown with limited
yellow mottling below midline. Dorsal surface of limbs
brown chocolate (same color as dorsal surface of head,
body and tail) with yellow specks, which are larger and
elongated on the hind limbs; dorsal surface of feet brown
with small yellow specks. Ventral surface of body, limbs,
gular region and tail pale brown with small yellow ecks.
Iris dark brown with yellow specks around the pupil.
Coloration of the holotype in preservative (Fig. 2A,
B). Dorsum nearly uniformly dark gray, including head
and tail, hands, and feet. All irregular spots on body and
specks on head cream. Ventral surface of body, limbs and
gular region pale gray with numerous cream specks.
Color variation in adult and juvenile specimens
(Fig. 4). The color pattern is similar in most adult speci-
mens. Irregular blotches on the body can vary in size and
shape and can be yellow or orange, forming elongated or
rounded patterns; they are smaller on the head and be-
come larger along the dorsum and even larger on the tail,
but their size varies from specimen to specimen (Fig. 4).
Regardless of the size of the blotches on the dorsum, most
specimens have a mottled pattern. There were two adult
specimens, one female (MZFC-HE 35857, Fig. 4C) and
one male, that were almost completely dark with small
yellow blotches. There appears to be ontogenetic varia-
tion in color pattern. In juveniles, the dorsum is almost
entirely dark brown or black, without the yellow mottling
or blotches as in adults. Some of the smallest specimens
(22 mm SVL) are almost totally black, with little or no
yellow dorsal coloration (Fig. 4G, H). Slightly larger ju-
veniles (30–40 mm SVL), show more yellow dorsal col-
oration (Fig. 4I, J).
Distribution and natural history. Pseudoeurycea jag-
uar sp. nov. is known only from Sierra de Zongolica
(Fig. 5A). It is found in a mature coniferous forest (Fig.
5B, C) at 2,360–2,367 m, which is dominated by Cupres-
sus benthamii, Pinus patula, Alnus acuminata, and Sau-
rauria leucocarpa with an understory of Fuchsia micro-
phyla, Rubus sp., Licianthes synanthera; and herbaceus
plants including Didymaea alsinoides, Chusquea mulleri
and Begonia oaxacana. There is an abundant presence of
epiphytes, including Tillandsia imperialis and Elapho-
glossum paleaceum, with extensive growth of mosses.
Pseudoeurycea jaguar sp. nov. is mainly an arboreal
species with nocturnal habits. The species was observed
active at night on trees, shrubs, rocks, herbaceous plants,
and moss and was also observed moving on the ground.
Most of the trees where P. jaguar was observed con-
tained layers of moss and bromeliads. By day, we found
P. jaguar sp. nov. hidden behind or within the layers of
moss that cover the trunks of the trees, in particular two
species of moss (Ptychomitrium sp. and Anacolia men-
ziesii).
Other species of plethodontids that share habitat
with P. jaguar sp. nov. in the study site are Aquiloeury-
cea cafetalera, Chiropterotriton sp., Isthmura gigantea,
Parvimolge townsendi, Thorius sp. and T. troglodytes.
Intraspecic antagonistic behavior. Antagonistic be-
havior between individuals in a population has been
described in several species of plethodontid salaman-
ders (Jaeger and Forester 1993; Staub 1993; Lynn et al.
2019). However, this behavior has been little studied or
observed in Mexican plethodontids. During our popu-
lation study, we only found one individual of P. jaguar
in each trap or tree studied. Initially, we collected some
individuals, transported them to the laboratory, and took
morphological measurements for mark-recapture stud-
ies. On the rst sampling trip, we kept three containers
with more than one individual, considering that each
container represented a particular section of the study
area. In the rst one, we kept ve immature individuals
with a large adult, in the second container a pair of adult
individuals, and in the third container two large adult in-
dividuals with a young adult. In all the containers there
were signs of serious aggression. In the rst, three of the
ve juvenile individuals disappeared; in the second, the
female and male adults mutilated each other, leaving the
male blind and the female with serious injuries to the
body; in the third container, the youngest individual was
killed. Some individuals of the population studied have
regenerated or regenerating tails, indicating some level
of predation or possibly agonistic interaction between
them.
Etymology. The specic epithet jaguar is a noun in appo-
sition and refers to the similarity between the dorsal color
pattern of the salamander and that of the jaguar (Panthera
onca). In the last three years the presence of this endan-
gered feline has been recorded in some places in the Si-
erra de Zongolica and it seems appropriate to honor this
emblematic species in the region.
Cázares-Hernández et al.: New species of Pseudoeurycea from Mexico
946
Molecular systematics
The results of our mtDNA phylogenetic analysis (Fig. 6)
showed that Pseudoeurycea jaguar sp. nov. belongs to
the Pseudoeurycea juarezi group and is the sister taxon of
Pseudoeurycea rucauda with strong support (Bootstrap
proportion [BS] = 74, Posterior Probability [PP] = 0.98).
These two species were part of a strongly supported clade
(BS = 83, PP = 1.0) with three additional species from
northern Oaxaca: P. aurantia, P. juarezi, and P. saltator.
GTR genetic distances between P. jaguar and P. rucau-
da were 0.030 (16S) and 0.096 (cyt b). These distances
were comparable to those between sister species in the
leprosa group and larger than those between other spe-
cies in the juarezi group ( Table 4). The 16S GTR distance
between P. juarezi and P. saltator is 0.0, and the distance
of P. aurantia to both of these species is 0.006; the larg-
est distance for 16S between sister species in these two
groups is 0.034 between P. conanti and P. mystax. For
cyt b, GTR distances range from a minimum of 0.037 be-
tween P. juarezi and P. saltator to a maximum of 0.113
between P. obesa and P. werleri.
Figure 5. A Geographic distribution of members of the P. juarezi group. B, C Habitat at the type locality of Pseudoeurycea jaguar
sp. nov. Cupressus forest, roots, trunk and branches of the trees have moss of the genera Ptychomitrium sp. and Anacolia.
Vertebrate Zoology 72, 2022, 937–950 947
Figure 6. Phylogeny estimated from maximum likelihood analysis of 16S and cyt b mtDNA sequence data. Numbers above branch-
es are bootstrap proportions from RAxML analysis and numbers below branches are posterior probabilities from Bayesian analysis.
Bootstrap proportions below 50 and posterior probabilities below 50 are not shown. The P. juarezi group is indicated in red.
Cázares-Hernández et al.: New species of Pseudoeurycea from Mexico
948
Discussion
Based on mtDNA, Pseudoeurycea jaguar sp. nov.
belongs to the P. juarezi group. Except for the new
species, which is endemic to the Sierra de Zongolica
of Veracruz, all the species belonging to the P. juarezi
group are distributed in cloud forest in the highlands
of northern Oaxaca, in the Sierra de Juárez, Sierra
Mazateca and Sierra Mixe. There is geographical
congruence in this clade (Figs 5, 6). Pseudoeurycea
jaguar sp. nov. has the northernmost distribution of
this subclade; it is most closely related to another
arboreal species P. rucauda from the cloud forest
from Sierra Mazateca and Sierra de Juárez in north-
ern Oaxaca, to which it is closest geographically.
These two species are the sister group of the other
subclade, which is composed of the terrestrial sal-
amanders P. aurantia, from cloud forest of Peña
Verde, Oaxaca (south of the Sierra Mazateca), and P.
juarezi, from cloud forest of the Sierra de Juárez and
Sierra Mixe, and the arboreal P. saltator from Sierra
Juárez (south of Peña Verde).
Because of its arboreal habits, P. jaguar is like-
ly vulnerable to logging and most of the trees it oc-
cupies are species of economic value. Logging is
occurring in the Sierra de Zongolica, including the
forest where P. jaguar sp. nov. occurs. Logging in
the forests of the Sierra de Zongolica is not properly
regulated and in many places it is carried out clan-
destinely. There is no follow-up to timber harvesting
programs intended to mitigate the impacts on the
ora and fauna found in the forests under manage-
ment. Further studies are required to determine the
distribution and population status of P. jaguar sp.
nov. This would allow us to obtain more information
about its natural history, as well as its relationship
with the forest environments it inhabits. Based on
the circumstances of the habitat, population ecolo-
gy and observed behavior, we apply the evaluation
criteria of the IUCN Red List (IUCN 2012), and the
Risk Assessment Manual (MER for its acronym in
Spanish) of the Secretaría de Medio Ambiente y Re-
cursos Naturales (Sánchez-Salas et al. 2013), and we
propose that P. jaguar sp. nov. should be considered
as a species at risk.
Based on our direct observations and with the best
available evidence on salamander communities in
the region, we can arm that P. jaguar is a vulnera-
ble taxon and that it may face risk of extinction in the
wild. The species is currently known from only one
locality, within a potential distribution range of less
than 20,000 km2 (following MER criteria, Sánchez
et al. 2007) and restricted to a single known type of
vegetation. These populations appear to be strongly
fragmented and we infer that because of the decrease
in habitat quality due to changes in land use and log-
ging of forests, there is a continuous long-term reduc-
tion (more than 10 years) in population size (estimat-
ed in fewer than 10,000 adult individuals currently).
Table 4. Generalized time-reversible (GTR) distances between species of the juarezi and leprosa groups of Pseudoeurycea. 16S distances are given above the diagonal and cyt b distances below the
diagonal. Distances between P. jaguar and other species are shown in bold.
P. mystax P. lynchi P. nigro-
maculata
P. rscheini P. leprosa P. ru
cauda
P. jaguar P. aurantia P. juarezi P. saltator P. obesa P. werleri P. lineola P. orchi
leucos
P. orchi
melas
P. conanti 0.034 0.043 0.049 0.047 0.043 0.056 0.047 0.043 0.041 0.041 0.043 0.041 0.043 0.038 0.043
P. mystax — 0.036 0.045 0.052 0.043 0.047 0.047 0.048 0.050 0.050 0.038 0.038 0.038 0.034 0.034
P. lynchi 0.172 — 0.023 0.030 0.023 0.036 0.028 0.034 0.037 0.037 0.041 0.039 0.036 0.032 0.038
P. nigromaculata — — — 0.032 0.030 0.043 0.036 0.038 0.041 0.041 0.047 0.050 0.047 0.039 0.052
P. rscheini 0.201 0.137 — — 0.023 0.050 0.043 0.045 0.048 0.048 0.059 0.061 0.052 0.050 0.052
P. leprosa 0.199 0.100 — 0.090 — 0.038 0.041 0.039 0.041 0.041 0.045 0.048 0.045 0.041 0.050
P. rucauda 0.180 0.191 — 0.192 0.204 — 0.030 0.025 0.028 0.028 0.050 0.057 0.052 0.043 0.061
P. jaguar 0.161 0.158 — 0.177 0.177 0.096 — 0.023 0.026 0.026 0.040 0.048 0.045 0.025 0.045
P. aurantia 0.150 0.175 — 0.166 0.154 0.165 0.164 — 0.006 0.006 0.050 0.050 0.047 0.036 0.047
P. juarezi 0.158 0.182 — 0.159 0.160 0.146 0.161 0.037 — 0.000 0.043 0.050 0.045 0.034 0.045
P. saltator 0.161 0.193 — 0.166 0.171 0.162 0.168 0.043 0.037 — 0.043 0.050 0.045 0.034 0.045
P. obesa 0.113 0.187 — 0.204 0.188 0.171 0.174 0.158 0.174 0.185 — 0.032 0.036 0.027 0.038
P. werleri 0.117 0.172 — 0.201 0.199 0.180 0.161 0.150 0.158 0.161 0.113 — 0.045 0.030 0.038
P. lineola 0.193 0.156 — 0.153 0.167 0.173 0.174 0.184 0.179 0.184 0.175 0.193 — 0.030 0.034
P. orchileucos 0.165 0.193 — 0.169 0.186 0.135 0.172 0.176 0.173 0.176 0.180 0.165 0.170 — 0.021
P. orchimelas 0.157 0.168 — 0.166 0.162 0.161 0.172 0.189 0.193 0.195 0.171 0.157 0.175 0.083 —
Vertebrate Zoology 72, 2022, 937–950 949
These changes may not be reversible in the long term due
to the eects of human activity, climate change aecting
its habitat, and/or chance events (for example, disease
outbreaks) that could put the species at greater risk. With
this information, we suggest a provisional assessment of
P. jaguar as “Vulnerable” using the Red List criteria (VU
4a, c; B1ab (iii); C1; D2) (IUCN 2012) and Threatened
(A) by the MER (Sánchez et al. 2007).
The species diversity of the Sierra de Zongolica is
high, and it is important to continue exploring other ar-
eas, since several records and new species have recently
been discovered (Canseco-Márquez et al. 2016; De La
Torre-Loranca et al. 2020; García-Vázquez et al. 2022).
This region is topographically complex, which generates
a diversity of microclimates and a great variety of eco-
systems, including Evergreen Tropical Forest, Cloud For-
est, Coniferous Forest and Pine-Oak forests (Rzedowski
2006; Miranda and Hernández 1963). The northern and
southwestern parts of the Sierra contain pine-oak forest,
coniferous forest, cloud forest; the central and south-
east portions have elements of cloud forest, tropical for-
est-cloud forest ecotones, evergreen tropical forest and
tropical lowland vegetation; the elevations generally go
from 100 to 2500 m. The Sierra de Zongolica is very im-
portant for its high number of plethodontid salamanders
(14 species; Cázares-Hernández et al. 2021): Bolitoglos-
sa platydactyla (in cloud forest and evergreen tropical
forest), B. rufescens (in evergreen tropical forest), Aqui-
loeurycea cafetalera, Isthmura gigantea (in cloud forest
and pine-oak forest), Parvimolge townsendi, Pseudoeury-
cea rscheini, P. granitum, P. nigromaculata, P. lineola,
P. jaguar, P. werleri, Thorius pennatulus, T. troglodytes
and Chiropterotriton sp. (in cloud forest, evergreen trop-
ical forest and Cupressus forest). Additional eldwork in
the Sierra de Zongolica and adjacent regions of Puebla
could reveal additional salamander species or expand the
known distribution of P. jaguar, highlighting the need for
continued herpetological exploration in the region.
Acknowledgments
We thank Rafael Rosales Martínez, Julián Rodríguez Lobato and Mauro
Daniel Castro Morales, students of the ITSZ Forest Engineering career,
for their collaboration in eldwork; Ángel Iván Contreras Calvario from
Facultad de Biología of the Universidad Veracruzana for his support and
collaboration in the eldwork; Ing. Wenceslao Cosme Reyes, Coordina-
tor of the Zongolica campus of the ITSZ, for his support in relation to
eld trips and nally especially to the Instituto Tecnológico Superior de
Zongolica. Víctor Vásquez Cruz helped with the map. Collecting permits
SEMARNAT-08-049, Ocio Núm.SGPA/DGVS/02924/15 were issued
to Erasmo Cázares-Hernández and collaborators. Three reviewers pro-
vided valuable comments that improved the quality of the manuscript.
References
Aljanabi SM, Martinez I (1997) Universal and rapid salt extraction of
high quality genomic DNA for PCR-based techniques. Nucleic Ac-
ids Research 25: 4692 –4693. https://doi.org/10.1093/nar/25.22.4692
AmphibiaWeb (2022) https://amphibiaweb.org University of Califor-
nia, Berkeley, CA, USA. Accessed 5 Apr 2022.
Canseco-Márquez L, Parra-Olea G (2003) A new species of Pseudo-
eurycea (Caudata: Plethodontidae) from northern Oaxaca, México.
Herpetological Journal. 13: 21–26.
Canseco-Márquez L, Pavón-Vázquez CJ, López-Luna MA, Nieto-Mon-
tes de Oca A (2016) A new species of earth snake (Dipsadidae, Geo-
phis) from Mexico. Zookeys 610: 131–45. https://doi.org/10.3897/
zookeys.610.8605
Cázares-Hernández E, Méndez Quiahua EC, Molohua Tzitzihua E, Vás-
quez-Cruz V (2021) Tlaconetes: Los Hijos de la Tierra. Zongo lica,
Veracruz, Mexico: Instituto Tecnológico Superior de Zongolica. 96 pp.
De La Torre-Loranca, MA, Martínez-Fuentes RG, Canseco-Márquez L,
García-Vázquez UO (2020) New records of amphibians and reptiles
from Sierra de Zongolica, Veracruz and Puebla. Herpetological Re-
view 51: 550–553.
Edgar RC (2004) MUSCLE: a multiple sequence alignment method
with reduced time and space complexity. BMC Bioinformatics 5:
1–19. https://doi.org/10.1186/1471-2105-5-113
García-Bañuelos P, Aguilar-López JL, Kelly-Hernández A, Vásquez-
Cruz V, Pineda E, Rovito SM (2020) A new species of Pseudoeu-
rycea (Amphibia: Caudata) from the mountains of central Vera-
cruz, Mexico. Journal of Herpetology 54: 258–267. https://doi.
org/10.1670/19-052
García-Castillo MG, Soto-Pozos AF, Aguilar-López JL, Pineda E, Par-
ra-Olea G (2018) Two new species of Chiropterotriton (Caudata:
Plethodontidae) from central Veracruz, Mexico. Amphibian & Rep-
tile Conservation 12: 37–54.
García-Vázquez UO, Clause AG, Gutiérrez-Rodríguez J, Caza-
res-Hernández E, De La Torre-Loranca MA (2022) A new species
of Abronia (Squamata:Anguidae) from the Sierra de Zongolica of
Veracruz, Mexico. Ichthyology and Herpetology 110: 33–49. https://
doi.org/10.1643/h2021051
Jaeger RG, Forester DC (1993) Social behavior of plethodontid sala-
manders. Herpetologica 49: 163–175. https://www.jstor.org/stable/
38 92793
IUCN (2012) IUCN Red List Categories and Criteria: Version 3.1. Sec-
ond edition. Gland, Switzerland and Cambridge, UK: iv + 32pp.
Lanfear R, Frandsen PB, Wright AM, Senfeld T, Calcott B (2017) Par-
titionFinder 2: new methods for selecting partitioned models of
evolution for molecular and morphological phylogenetic analyses.
Molecular Biology and Evolution 34: 772–773.
Lynch JF, Wake DB (1989) Two new species of Pseudoeurycea (Am-
phibia: Caudata) from Oaxaca, Mexico. Contributions in Science,
Los Angeles County Museum of Natural History 411: 11–22.
Lynn CS, Dalton B, Mathis A (2019) Territorial behavior in South-
ern Red-Backed and Ozark Zigzag salamanders: eects of sex,
species, and ownership. Behaviour 156: 1017–1037. https://doi.
org/10.1163/1568539X-00003554
Miller MA, Pfeier W, Schwartz T (2010) Creating the CIPRES Sci-
ence Gateway for inference of large phylogenetic trees. Proceedings
of the Gateway Computing Environments Workshop (GCE) 2010:
1–8. https://doi.org/10.1109/GCE.2010.5676129
Miranda F, Hernández XE (1963) Los tipos de vegetación de México
y su clasicación. Boletín de la Sociedad Botánica de México 28:
29–179.
Moritz C, Schneider CJ, Wake DB (1992) Evolutionary relationships
within the Ensatina eschscholtzii complex conrm the ring species
interpretation. Systematic Biology 41: 273–291. https://doi.org/
10.1093/sysbio/41.3.273
Cázares-Hernández et al.: New species of Pseudoeurycea from Mexico
950
Appendix
Specimens examined:
Institutional abbreviations for museums and collections follow Sabaj
(2022). Museum of Vertebrate Zoology, UC Berkeley (MVZ), Instituto
de Biología, UNAM (IBH), Museo de Zoología, Facultad de Ciencias,
UNAM (MZFC) and Escuela de Biología, Benemérita Universidad
Autónoma de Puebla (EBUAP).
Pseudoeurycea aurantia (n=5). MEXICO: OAXACA: 11.8 km E
(by air) Concepción Pápalo, Sierra de Juárez (MVZ 270127─28,
270132, 270134–35). 4 km W Peña Verde (EBUAP 2051, Holo-
type).
Pseudoeurycea jaguar sp nov. (n=9) MEXICO: VERACRUZ: Texhua-
can Municipality: El Mirador (MZFC-HE 28694 holotype, MZFC-
HE 28685–86, 35855–59, paratypes, MZFC-HE 35860)
Pseudoeurycea juarezi. (n=21) MEXICO: OAXACA: 52 km W
Guelatao along Mexico Hwy. 175 (MVZ 112192, 147186, 162121);
N slope of Cerro Pelón along Mexico Hwy. 175, 52–54 km N (by
road) Guelatao (MVZ 131031, 131037, 131039); 0.7 mi E (by road)
Cerro Pelón from point where road crosses top (MVZ 86053); 0.7
mi E (by road) or NE Cerro Pelón (MVZ 86072); 1.1 km N (by rd)
of Cerro Pelón mirador on MX Hwy 175 (IBH 32420); Cerro Pelón,
near Mexico Hwy. 175, 108.9 km N (by road) Oaxaca de Juárez
from junction Pan American Hwy. (MVZ 114386–87); 52 km W
Guelatao along Mexico Hwy. 175 (MVZ 112196, 147198); Mexico
Hwy. 175, 57 km N (by road) Guelatao (MVZ 131044); 1.5 km NE
(by road) Cuajimoloyas, Distrito Ixtlán (MVZ 131097); 18.7 km E
(by road) Cuajimoloyas along road to Cajones (MVZ 194309); 5.8
km NW (by rd) of junction with road to Zacatepec on MX Hwy 179
to Totontepec Villa de Morelos (IBH 32365); 11.1 mi SW (by road)
Totontepec [=Totontepec Villa de Morelos] (MVZ 163769); 19 km
SW (by road) Totontepec [=Totontepec Villa de Morelos] (MVZ
164662–63, 164665); Cerro Zempoaltepec (MVZ 163874);
Pseudoeurycea rucauda. (n=2) MEXICO: OAXACA: 4.0 km NE (by
rd) of Peña Verde on road to Tlalixtac Viejo, Sierra de Juárez (IBH
22610); 2 km NW Puerto de Soledad (MVZ 236762).
Pseudoeurycea saltator. (n=5) MEXICO: OAXACA: San Pedro Yolox
Municipality: La Galera, 11.0 km SW (by rd) of La Esperanza on
MX Hwy 175 (IBH 32193, 32402); 65 km NE of Guelatao, by Mex-
ico Hwy. 175 (MVZ 112230, 147263); forest along Mexico Hwy.
175, 129.9 km N (by road) Oaxaca de Juárez Ctr., junction Panam
Hwy (MVZ 114398).
Palumbi SR, Martin AP, Romano S, McMillan WO, Stice L, Grabowski
G (1991) The Simple Fool’s Guide to PCR. Department of Zoology,
University of Hawaii, Honolulu, Hawaii, U.S.A. 45 pp.
Parra-Olea G, García-París M, Hanken J, Wake DB (2004) A new spe-
cies of arboreal salamander (Caudata: Plethodontidae: Pseudoeu-
rycea) from the mountains of Oaxaca, Mexico. Journal of Natural
History. London 38: 2119–2131. https://doi.org/10.1080/00222930
310001617724
Parra-Olea G, García-Castillo MG, Rovito SM, Maisano JA, Hanken J,
Wake DB (2020) Descriptions of ve new species of the salaman-
der genus Chiropterotriton (Caudata: Plethodontidae) from eastern
Mexico and the status of three currently recognized taxa. PeerJ 8:
e8800. https://doi.org/10.7717/peerj.8800
Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Hohna S,
Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2:
Ecient Bayesian phylogenetic inference and model choice across
a large model space. Systematic Biology 61: 539–542. https://doi.
org/10.1093/sysbio/sys029
Rovito SM, Parra-Olea G, Recuero E, Wake DB (2015) Diversica-
tion and biogeographical history of Neotropical plethodontid sala-
manders. Zoological Journal of the Linnean Society 175: 167–188.
https://doi.org/10.1111/zoj.12271
Rzedowski J (2006) Vegetación de México.1ra Edición digital, Comis-
ión Nacional para el Conocimiento y Uso de la Biodiversidad, Méx-
ico. 504 pp.
Sabaj MH (2022) Codes for Natural History Collections in Ichthyolo-
gy and Herpetology (online supplement). Version 9.0 (14 February
2022). Electronically accessible at https://asih.org, American Soci-
ety of Ichthyologists and Herpetologists, Washington, DC.
Sánchez-Salas J, Muro G, Estrada-Castillón E, Alba-Ávila JA (2013) El
MER: un instrumento para evaluar el riesgo de extinción de especies
en México. Revista Chapingo Serie Zonas Áridas 12: 30–35.
Sandoval-Comte A, Pineda-Arredondo E, Rovito SM, Luría-Manza-
no R (2017) A new species of Isthmura (Caudata: Plethodontidae)
from the montane cloud forest of central Veracruz, Mexico. Zootaxa
4277: 573–582. https://doi.org/10.11646/zootaxa.4277.4.7
Stamatakis A (2014) RAxML version 8: A tool for phylogenetic analysis
and post-analysis of large phylogenies. Bioinformatics 30: 1312–
1313. https://doi.org/10.1093/bioinformatics/btu033
Staub NL (1993) Intraspecic agonistic behavior of the salamander
Aneides avipunctatus (Amphibia: Plethodontidae) with compar-
isons to other plethodontid species. Herpetologica 49: 271–282.
https://www.jstor.org/stable/3892804
Swoord D (2002) PAUP*: phylogenetic analysis using parsimony
(* and other methods), version 4. Sinauer Associates, Sunderland,
Massachusetts, U.S.A.
Torres-Hernández LA, Ramírez-Bautista A, Cruz-Elizalde R, Hernán-
dez-Salinas U, Berriozabal-Islas C, DeSantis DL, Johnson JD,
Rocha A, García-Padilla E, Mata-Silva V, Fucsko LA, Wilson LD
(2021) The herpetofauna of Veracruz, Mexico: composition, distri-
bution, and conservation status. Amphibian & Reptile Conservation
15: 72–155 (e285). http://amphibian-reptile-conservation.org/pdfs/
Volume/Vol_15_no_2/ARC_15_2_[General_Section]_72-155_
e285.pdf
Wake DB (1987) Adaptive radiation of salamanders in Middle Amer-
ican cloud forests. Annals of the Missouri Botanical Garden 74:
242–264.
Wake DB (2012) Taxonomy of salamanders of the family Plethodonti-
dae (Amphibia: Caudata). Zootaxa 3484: 75–82.
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