A New Species of Pseudoeurycea (Amphibia: Caudata)

from the Mountains of Central Veracruz, Mexico

Authors: García-Bañuelos, Paulina, Aguilar-López, José Luis, Kelly-

Hernández, Alfonso, Vásquez-Cruz, Víctor, Pineda, Eduardo, et al.
Source: Journal of Herpetology, 54(2) : 258-267
Published By: Society for the Study of Amphibians and Reptiles
URL: https://doi.org/10.1670/19-052

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 Journal of Herpetology, Vol. 54, No. 2, 258–267, 2020
Copyright 2020 Society for the Study of Amphibians and Reptiles

A New Species of Pseudoeurycea (Amphibia: Caudata) from the Mountains of Central

Veracruz, Mexico

PAULINA GARCÍA-BAÑUELOS,1 JOSÉ LUIS AGUILAR-LÓPEZ,1 ALFONSO KELLY-HERNÁNDEZ,2 VÍCTOR VÁSQUEZ-CRUZ,2 EDUARDO
PINEDA,1 AND SEAN M. ROVITO3,4
1
Red de Biologı́a y Conservación de Vertebrados, Instituto de Ecologı́a, A.C., Xalapa, Veracruz, México
2
Predio o Instalación para el Manejo de la Vida Silvestre, Herpetario Palancoatl, Córdoba, Veracruz, México
3
Unidad de Genómica Avanzada (Langebio), Centro de Investigación y de Estudios Avanzados del Instituto Politécnico Nacional, Irapuato, Guanajuato,
México

ABSTRACT.—We describe a new species of plethodontid salamander of the genus Pseudoeurycea from the mountains of central Veracruz,
Mexico. This species can be distinguished from its congeners by color pattern and morphological characters. The molecular phylogenetic
analysis indicates that Pseudoeurycea granitum sp. nov. is part of the Pseudoeurycea leprosa subclade and is closely related to
Pseudoeurycea nigromaculata, Pseudoeurycea firscheini, and Pseudoeurycea lynchi. Known localities for P. granitum sp. nov. are mostly
located in cloud forest fragments, with one from a shaded coffee plantation and one from an ecotone between cloud forest and pine–oak
forest, all immersed in highly modified landscapes. This suggests the need to implement a conservation strategy in order to preserve the
limited remaining habitat for this species.

RESUMEN.—Describimos una nueva especie de salamandra pletodóntida del género Pseudoeurycea de las montañas del centro de
Veracruz, México. Esta especie puede distinguirse de sus congéneres por su patrón de coloración y sus caracteres morfológicos. El análisis
filogenético molecular indica que Pseudoeurycea granitum sp. nov. es parte del subclado de P. leprosa y está estrechamente relacionada
con P. nigromaculata, P. firscheini y P. lynchi. Las localidades conocidas para Pseudoeurycea granitum sp. nov. están principalmente
ubicadas en fragmentos de bosque de niebla, una en una plantación de cafetal de sombra y otra en un ecotono entre bosque de niebla y
bosque de pino-encino, todos inmersos en paisajes altamente modificados, lo que sugiere la necesidad de implementar una estrategia de
conservación para preservar el limitado hábitat remanente para esta especie.

The genus Pseudoeurycea Taylor 1944, composed of 39 species,
is the third most diverse among plethodontid salamanders and
the most diverse in Mexico. Except for Pseudoeurycea brunnata,
Pseudoeurycea goebeli, and Pseudoeurycea rex, which are distrib-
uted in both Mexico and Guatemala, and Pseudoeurycea
exspectata, which is endemic to Guatemala, all species of
Pseudoeurycea are endemic to Mexico (AmphibiaWeb, 2019).
Most of these species are distributed in the Sierra Madre del Sur,
northern Oaxaca highlands, and central to eastern Trans-
Mexican Volcanic Belt (TMVB; Parra-Olea, 2002; Raffaëlli,
2013). The elevational range of the genus is from the sea level
to over 4,000 m above sea level (a.s.l.), but the greatest species
diversity (28 spp.) is between 1,500 and 2,500 m a.s.l. in pine
and oak forests (Parra-Olea et al., 2010).
The mountains of central Veracruz are at the intersection
between the eastern TMVB, the southern Sierra Madre Oriental
and northern highlands of the Oaxaca Province (CONABIO,
1997). This region, with elevations between 800 and 5,600 m
a.s.l., was historically covered by pine, fir, and oak forests,
grassland, and sizeable areas of cloud forest, and is recognized
for its high diversity of plethodontid salamanders (Wake et al.,
1992; Parra-Olea et al., 2001). To date, there are 28 described
species of plethodontids in this region. However, the diversity
seems to be even higher, because four species have been
described in the last decade: Aquiloeurycea cafetalera, Isthmura
corrugata, Chiropterotriton aureus, and Chiropterotriton nubilus. All
of these species were described from cloud forest, with more
species in collections awaiting description. Of the total
plethodontid species richness of central Veracruz, seven species
(a quarter of the total) belong to the genus Pseudoeurycea
4
Corresponding author. E-mail: sean.rovito@cinvestav.mx
DOI: 10.1670/19-052
(Pseudoeurycea firscheini, Pseudoeurycea gadovii, Pseudoeurycea
leprosa, Pseudoeurycea lineola, Pseudoeurycea lynchi, Pseudoeurycea
melanomolga, and Pseudoeurycea nigromaculata).
During fieldwork carried out from 2015 to 2019 in different
locations in the mountainous region of central Veracruz, we
found 24 salamander specimens with morphological character-
istics that did not coincide with those of any described species of
Pseudoeurycea. Based on morphological and molecular evidence,
we describe a new species of the genus Pseudoeurycea from three
localities in central Veracruz (Fig. 1), closely related to P.
nigromaculata, P. firscheini, and P. lynchi.
MATERIALS AND METHODS
Phylogenetic Analyses.—We extracted DNA from liver tissue of
three individuals from the populations from Tepexilotla and
Cerro Petlalcala, including a salamander that was regurgitated by
a snake (Thamnophis sumichrasti), one individual of P. nigromacu-
lata, and one individual of P. firscheini using a high salt protocol
(Aljanabi and Martinez, 1997). We amplified a fragment of the
16S ribosomal RNA gene (16S) using primers 16Sal and 16Sbr
(Palumbi et al., 1991) and a fragment of the cytochrome b (cytb)
gene using primers MVZ15 and MVZ16 (Moritz et al., 1992). PCR
products were cleaned using 1 lL of 1:5 diluted EXOSAP-IT (USB
Corporation, Cleveland, Ohio), cycle sequenced using BigDye v3
terminator chemistry (Applied Biosystems, Foster City, Califor-
nia) and sequenced on an ABI3730 capillary sequencer. Voucher
information and GenBank numbers for all sequences used in
phylogenetic analyses are given in Table 1. We used Aquiloeurycea
galeanae as an outgroup.
We edited sequences using Geneious v8.1.8 (Biomatters) and
aligned sequencers for each gene using Muscle v.3.8 (Edgar,
2004) with default parameters. Alignments were trimmed to the

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 NEW SPECIES OF PSEUDOEURYCEA FROM MEXICO 259

FIG. 1. Localities where Pseudoeurycea granitum sp. nov., P. nigromaculata, P. firscheini, and P. lynchi have been recorded in central Veracruz and
Puebla, Mexico.

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 260 P. GARCÍA-BAÑUELOS ET AL.

TABLE 1. Voucher information and GenBank numbers for specimens Morphological Analyses.—We compared salamanders from the
included in phylogenetic analysis. new populations from Tepexilotla, Mundo Nuevo, and Cerro
Petlalcala to the other two species of Pseudoeurycea with greenish
Species Voucher number GenBank 16S GenBank cytb
or yellowish-brown coloration from the cloud forests of central
Pseudoeurycea ahuitzotl IBH 30211 MT303858 MT295473 Veracruz: P. lynchi and P. nigromaculata. We measured 45
P. altamontana IBH 22220 KP886861 KP900064 specimens (12 males and 33 females of all three species) for the
P. anitae MVZ 137939 AF451227 — following morphological characteristics using digital calipers:
P. aurantia IBH 20370 KP886844 KP900048 Distance from snout to posterior angle of vent (snout–vent length
P. brunnata MVZ 137947 AF451232 —
P. cochranae IBH 23064 KP886864 KP900067 [SVL]), distance from posterior angle of vent to tip of tail (tail
P. conanti MVZ 146786 AF451241 — length, TL), axilla–groin distance (AX), forelimb length (FLL),
P. exspectata MVZ 160919 AF451234 — hind-limb length (HLL), distance from tip of snout to gular fold
P. firscheini IBH 30995 MT303859 MT295469 (head length, HL), width of head at angle of jaw (HW), head
P. firscheini IBH 23102 KP886865 KP900068
P. gadovii IBH 22982 KP886846 KP900050 depth (HD), interocular distance (IO), distance between external
P. goebeli CRVA1017 MT303860 MT295472 nares (IN), right foot width (RFW), length of longest (third) toe
P. granitum sp. nov. IBH 31829 MT303861 MT295471 (T3), and length of fifth toe (T5). We counted ankylosed maxillary
P. granitum sp. nov. CARIE 1263 MT303862 MT295470 and premaxillary teeth (MT) and vomerine teeth (VT), with left
P. granitum sp. nov. CARIE 1264 MT303863 — and right sides summed. In order to increase the number of adult
P. juarezi IBH 29718 KP886848 KP900052
P. leprosa IBH 22406 KP886866 KP900069 salamanders in our sample, we obtained a subset of measure-
P. lineola IBH 29719 KP886867 KP900070 ments (SVL, AX, TL, FLL, HLL, HL, HW, RFW, MT, VT) for part
P. longicauda IBH 22247 KP886849 KP900053 of the type series of P. lynchi (four males, six females) from David
P. lynchi GP160 AF451225 AF451204 B. Wake (pers. comm.). Salamanders measured for our morpho-
P. melanomolga IBH 22784 KP886868 KP900071
P. mixcoatl IBH 14194 KP886869 KP900072 logical analyses are listed in Appendix 1. We used Köhler (2012)
P. mixteca GP289 AF380829 — to describe color patterns, with capitalized color names and color
P. mystax GP372 AF380795 — numbers in parentheses.
P. nigromaculata MVZ 185977 AF451238 — We separated males and females for all morphological analysis
P. nigromaculata CARIE 1260 MT303864 MT295468 because Neotropical salamanders are typically sexually dimor-
P. obesa MVZ 241574 KP886870 KP900073
P. orchileucos IBH 22562 KP886858 KP900062 phic. We calculated mean, standard deviation, and range for all
P. orchimelas IBH 22999 KP886860 KP900063 morphological measurements and tooth counts and compared
P. papenfussi IBH 14198 KP886850 KP900054 the central Veracruz populations to the other two species.
P. rex MVZ 263590 KP886852 KP900056
P. robertsi IBH 22232 KP886853 KP900057
P. ruficauda IBH 21646 KP886871 KP900074 RESULTS
P. saltator IBH 22895 KP886854 KP900058
P. smithi IBH 29720 KP886855 KP900059 The mtDNA phylogeny (Fig. 2) showed that the populations of
P. tenchalli IBH 29721 KP886856 KP900060 Pseudoeurycea from Tepexilotla and Cerro Petlalcala are closely
P. tlahcuiloh IBH 30233 MT303865 MT295474 related; sequenced individuals from Cerro Petlalcala differ from
P. unguidentis MVZ 117432 MT303866 — those from Tepexilotla by only 0.6% (16S) and 2.2% (cytb). These
P. werleri IBH 22294 KP886872 KP900075
Ixalotriton niger IBH 29715 KP886874 KP900077 populations are part of a strongly supported clade (bootstrap
I. parvus AMA2534 KP886873 KP900076 support [BS] = 82, posterior probability [PP] = 0.95) that also
Aquiloeurycea galeanae IBH 24595 KP886847 KP900051 contains P. nigromaculata, P. lynchi, and P. firscheini. For 16S, mean
pairwise genetic distance between the Petlalcala/Tepexilotla
populations and their closest relatives varies from 0.025 (to P.
point where a majority of sequences had data, resulting in lynchi) to 0.035 (to P. firscheini); for cytb, mean pairwise distance
alignment lengths of 527 base pairs (bp) for 16S and 809 bp for varies from 0.101 (to P. firscheini) to 0.141 (to P. leprosa; Table 2).
cytochrome b. We concatenated the 16S and cytb alignments, Relationships between species within this clade are weakly
partitioned the data by gene and by codon position (for cytb) supported (BS < 50, PP < 50). All four of the species in this clade
and used PartitionFinder v2.1 (Lanfear et al., 2017) to select a are endemic to the mountains of central Veracruz and adjacent
partitioning scheme and nucleotide substitution models for each regions of Puebla. The sister species to this clade is P. leprosa (BS =
partition with the Bayesian Information Criterion (BIC). The 99, PP = 1.0), which ranges more widely throughout the
selected partitioning scheme and substitution models were 16S mountains of central Mexico at higher elevations.
+ cytb codon position 1 (GTR + I + G model), cytb codon The results of our phylogenetic analysis and morphological
position 3 (GTR + G), and cytb codon position 2 (HKY + I); we comparisons (see below) support the distinctiveness of the
used this partitioning scheme for both maximum likelihood Tepexilotla, Mundo Nuevo, and Cerro Petlalcala populations as
(ML) and Bayesian phylogenetic analyses. a separate species.
We used RAxML v.8.2 (Stamatakis, 2014) for ML analysis Pseudoeurycea granitum sp. nov.
with the GTR + G model for all partitions (RAxML does not Suggested English name: Granite-colored salamander
implement less complex models) and 1,000 bootstrap replicates Suggested Spanish name: Salamandra de color granito
to assess nodal support. We conducted a Bayesian phylogenetic (Fig. 3A,D–H)
analysis using MrBayes v3.2 (Ronquist et al., 2012) with four Zoobank ID: urn:lsid:zoobank.org:act:2375F867-3CC2-4345-
chains (three hot, one cold) run for 20,000,000 generations and B982-179BFBF2CD18
sampled every 1,000 generations. The first 5,000 samples were
discarded as burn-in when estimating the consensus tree. Holotype.—CARIE 1274 (Colección de Anfibios y Reptiles del
Finally, we estimated genetic distances between species using Instituto de Ecologı́a A.C.: field number PGB044). An adult
the GTR model in PAUP v4 (Swofford, 1998). female (Fig. 3A,D) from Mundo Nuevo, Chocamán, Veracruz

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 NEW SPECIES OF PSEUDOEURYCEA FROM MEXICO 261

FIG. 2. Phylogeny from maximum-likelihood analysis of 16S + cytb sequence data. Bootstrap values are given above branches, and posterior
probabilities from Bayesian analyses are given below branches. Bootstrap proportions and posterior probabilities below 50 and 0.5, respectively, are
not shown.

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 262 P. GARCÍA-BAÑUELOS ET AL.
TABLE 2. Mean GTR genetic distances between species closely related to P. granitum sp. nov. Values above diagonal are from 16S data, and values
below diagonal are from cytb data.
P. granitum sp. nov.
Tepexilotla
P. granitum sp. nov. Tepexilotla – 0.006
P. granitum sp. nov. Cerro Petlalcala 0.022
P. nigromaculata 0.119
P. lynchi 0.102
P. firscheini 0.099
P. leprosa 0.141
(18857 0 N, 9782 0 W, 1,919 m a.s.l., WGS84 datum), collected on 25
November 2018 by P. Garcı́a-Bañuelos at 1443 h.
Paratypes.—Twelve specimens, all from central Veracruz,
Mexico. Four males: CARIE 1261, 1264, from riparian forest of
the Metlac River near Tepexilotla, Chocamán (18858 0 N, 9784 0 W,
1,574 m a.s.l., WGS84 datum); CARIE 1286, Mundo Nuevo,
Chocamán (18857 0 N, 9782 0 W, 1,922 m a.s.l., WGS84 datum); IBH
31826 (Instituto de Biologı́a, UNAM), Cerro Petlalcala above
village of Petlalcala (18846 0 N, 9786 0 W, 2,043 m a.s.l., WGS84
datum). Eight females: CARIE 1262, 1263 (Fig. 3F), 1265 same
data as CARIE 1261; CARIE 1285, 1287, 1288, IBH 32550, same
data as CARIE 1286; IBH 31827, same data as IBH 31826.
Referred Specimens.— CARIE 1260 (Fig. 3E), 1266, from riparian
forest of the Metlac River near Tepexilotla, Chocamán, Veracruz
(18858 0 N, 9784 0 W, 1,574 m a.s.l., WGS84 datum); CARIE 1273,
from Mundo Nuevo, Chocamán, Veracruz (18857 0 N, 9782 0 W,
1,919 m a.s.l., WGS84 datum); IBH 31828–31830, same data as
IBH 31826.
Diagnosis.—Assigned to the genus Pseudoeurycea because of the
presence of a sublingual fold, relatively reduced foot webbing,
and mtDNA sequence data. Differs from all other species of
Pseudoeurycea by the presence of an interorbital bar (generally
FIG. 3. Dorsal (A) and ventral view (D) of the holotype of
Pseudoeurycea granitum sp. nov. (CARIE 1274) and variation of tail,
lateral surface (E, F), and interorbital bar (G, H) coloration between
individual of this species. Coloration of Pseudoeurycea lynchi from La
Cortadura, Coatepec, Veracruz (B) and Pseudoeurycea nigromaculata from
Cerro Chicahuaxtla, Ixtaczoquitlán, Veracruz (C). Scale bar = 10 mm.
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P. granitum sp. nov.
Cerro Petlalcala P. nigromaculata P. lynchi P. firscheini P. leprosa
0.027 0.022 0.032 0.026
– 0.033 0.028 0.038 0.032
0.130 – 0.025 0.035 0.037
0.112 0.146 – 0.030 0.028
0.103 0.143 0.121 – 0.026
0.141 0.167 0.130 0.111 –
conspicuous, sometimes less distinct) extending between the
outer edges of the eyelids; this bar varies in coloration from white
to brown (Fig. 3) but is always paler than the background dorsal
color. An interorbital bar is present in some species of the genus
Dendrotriton, which occurs only east of the Isthmus of Tehuante-
pec, but species of Dendrotriton differ in many morphological
features from P. granitum sp. nov. including having rugose skin,
smaller size, and the lack of a prefrontal bone (Lynch and Wake,
1975). Differs from most other species of this genus except P.
nigromaculata and P. lynchi by having greenish-yellow to reddish-
brown granite-like coloration with numerous splotches and small
flecks on the tail with black spots on the body. Pseudoeurycea
conanti often has a yellowish-green tail, but this yellowish
coloration is either unbroken or mottled with very dark gray–
brown or black, rather than having lighter splotches and specks;
P. conanti also has much smaller and more webbed feet, with only
the terminal phalanx free of webbing on the third toe. Like P.
conanti, P. kuautli has a yellowish tail but without the granite-like
flecks and splotches seen in P. granitum sp. nov.; P. kuautli also
has more webbed feet and a shorter tail (TL/SVL = 0.9 in
holotype of P. kuautli; Campbell et al., 2013). Pseudoeurycea saltator
has an unbroken and distinct yellowish band on tail, but this
band extends as a distinct dorsal stripe that is lacking in P.
granitum sp. nov.; P. saltator lacks black spots on body, although
some individuals have a few black spots on head.
Two of the species most closely related to P. granitum sp. nov.
based on our mtDNA phylogeny (Fig. 2), P. lynchi and P.
nigromaculata, are generally similar to P. granitum sp. nov. but
differ in multiple morphological characteristics as well as
specific aspects of color pattern, in addition to the previously
mentioned interorbital bar. Pseudoeurycea lynchi tends to have
greenish-yellow coloration with black spots or blotches from
head to tip of tail, while the yellowish coloration in P. granitum
sp. nov. is mostly limited to the tail and posterior part of
dorsum; the tail of P. lynchi also lacks paler blotches or specks
seen in P. granitum sp. nov. (Fig. 3A,C) Pseudoeurycea
nigromaculata also has a tail with paler blotches similar to those
seen in P. granitum sp. nov., but its tail tends to be reddish-
brown rather than yellowish in coloration (Fig. 3A,C). Pseu-
doeurycea granitum sp. nov. is smaller than the other two species
(males: P. granitum sp. nov. SVL 34.3–43.5 mm, P. lynchi 40.6–
48.3 mm, P. nigromaculata 42.5–49.2 mm; females: P. granitum sp.
nov. 37.2–50.2 mm, P. lynchi 37.3–52.1 mm, P. nigromaculata 43.4–
57.7 mm; Tables 3 and 4) and has a narrower head (HW males:
P. granitum sp. nov. 5.3–6.3 mm, P. lynchi 6.5–7.8 mm, P.
nigromaculata 6.5–7.7 mm; females: P. granitum sp. nov. 5.6–7.4
mm, P. lynchi 5.9–8.2 mm, P. nigromaculata 6.9–9.0 mm). Females
of P. granitum sp. nov. have a longer tail relative to body size
(TL/SVL females: P. granitum sp. nov. 1.10–1.49, P. lynchi 0.88–
1.19, P. nigromaculata 0.84–1.15). On average, P. granitum sp.
nov. has fewer maxillary and premaxillary teeth (mean MT
 NEW SPECIES OF PSEUDOEURYCEA FROM MEXICO 263

TABLE 3. Mean, standard deviation, and range of 13 morphological measurements and tooth counts for males.

Pseudoeurycea granitum sp. nov. n = 4 Pseudoeurycea lynchi n = 7 Pseudoeurycea nigromaculata n = 5

SVL 39.3 6 4.68 (34.3–43.5) 44.8 6 2.95 (40.6–48.3) 46.0 6 2.53 (42.5–49.2)
TL 39.5 6 3.40 (35.8–42.5) n = 3 49.2 6 10.04 (39.8–59.8) n = 3 50.6 6 8.29 (40.4–60.5) n = 4
AX 22.1 6 3.83 (17.7–25.9) 23.7 6 1.77 (21.4–26.0) 24.3 6 2.41 (21.1–27.0)
FLL 9.9 6 1.80 (8.3–11.9) 12.0 6 1.34 (9.1–13.1) 11.2 6 1.14 (9.9–12.7)
HLL 10.7 6 1.27 (9.3–12.0) 13.4 6 1.09 (11.9–14.9) 12.8 6 0.92 (11.8–13.9)
HL 9.1 6 0.87 (8.3–10.1) 10.3 6 0.88 (8.9–11.1) 10.7 6 1.10 (9.1–11.7)
HW 5.9 6 0.40 (5.3–6.3) 7.0 6 0.59 (6.5–7.8) 7.0 6 0.48 (6.5–7.7)
HD 2.8 6 0.11 (2.7–2.9) 3.2 6 0.05 (3.2–3.3) n = 3 3.9 6 0.68 (2.9–4.7)
IO 2.1 6 0.14 (1.9–2.2) 2.4 6 0.12 (2.2–2.4) n = 3 2.5 6 0.21 (2.2–2.8)
IN 1.9 6 0.24 (1.6–2.1) 2.3 6 0.33 (1.9–2.5) n = 3 2.3 6 0.23 (1.9–2.5)
RFW 4.5 6 0.74 (3.8–5.2) 4.9 6 0.96 (3.5–5.9) 5.0 6 0.76 (4.1–5.8)
T3 2.2 6 0.33 (1.9–2.6) 2.6 6 0.39 (2.2–3.0) n = 3 2.4 6 0.45 (1.8–3.0)
T5 1.1 6 0.23 (0.9–1.4) 1.1 6 0.38 (0.9–1.6) n = 3 1.2 6 0.31 (0.8–1.6)
MT 65.0 6 6.73 (57–73) 73.9 6 16.73 (43–92) 77.0 6 13.73 (62–91)
VT 21.8 6 4.35 (16–26) 30.1 6 4.88 (25–40) 26.4 6 2.70 (23–29)

males: P. granitum sp. nov. 65.0, P. lynchi 73.9, P. nigromaculata
77.0; females: P. granitum sp. nov. 71.6, P. lynchi 74.9, P.
nigromaculata 82.8) and vomerine teeth (mean VT males: P.
granitum sp. nov. 21.8, P. lynchi 30.1, P. nigromaculata 26.4;
females: P. granitum sp. nov. 24.1, P. lynchi 26.5, P. nigromaculata
28.5). When comparing morphological measurements adjusted
for body size by dividing each individual measurement by SVL,
only the relative tail length in females remains distinct between
P. granitum sp. nov. and the two other species.
The other two species most closely related to Pseudoeurycea
granitum sp. nov., P. firscheini and P. leprosa, differ in numerous
aspects of color pattern and morphology. Pseudoeurycea firscheini
is larger, reaching up to 63.5 mm SVL (Kelly-Hernández et al.,
2019), has a tail that is typically shorter than SVL, a relatively
narrower head (HW/SVL = 0.13–0.14 in P. firscheini vs. 0.14–
0.16 in P. granitum sp. nov.) and is gray–brown in coloration,
lacking black spots on body, with some pale splotches on the tail
similar but no yellow or yellow–green coloration; its tail often
has a reddish tip that is lacking in P. granitum sp. nov., and it
does not have a pale interorbital bar that is typically present in
P. granitum sp. nov. Pseudoeurycea leprosa is also larger
(maximum SVL approximately 58 mm), has a conspicuously
narrow head with a poorly defined neck (HW/SVL = 0.12–
0.15), little webbing on the feet, very short limbs, and a
relatively shorter tail that is approximately 80–100% SVL.
Pseudoeurycea leprosa is variable in coloration but never has
any yellowish color and is typically dark grayish brown with
reddish-brown dorsal coloration and numerous gray specks on
sides of body.
Description of the Holotype.—An adult female, moderately sized
for its genus (SVL 50.2 mm), body fairly slender (shoulder width
= 6.8). Head relatively narrow (HW/SVL = 0.15), only slightly
wider than body, with clearly defined neck. Mouth bluntly
rounded, eyes only slightly protuberant, not visible beyond
margin of jaw when viewed from above. External nares small,
oval-shaped. Tail longer than body (TL/SVL = 1.17), slender, and
tapering gradually along length; roughly rectangular at base,
becoming gradually more rounded towards tip. Limbs relatively
short (FLL + HLL/SVL = 0.55), adpressed limbs separated by
two costal interspaces. Digits long and slender with limited
webbing, reaching only to base of penultimate phalanx on third
toe of foot. First toe very short, fifth toe comparatively long; toe
tips blunt with distinct subterminal pads present. Order of digits
by increasing length: manus I–IV–II–III, pes I–V–II–IV–III.
Phalangeal formulae: manus 1–2–3–2, pes 1–2–3–3–2. Maxillary
teeth relatively small and numerous (71); premaxillary teeth
relatively numerous (8), same size as maxillary teeth; vomerine
teeth arranged in two arcs that curve towards palate at midline.
Nasolabial protuberances relatively weakly developed.
Variation.—All other specimens in the type series are smaller
than the holotype (SVL males 34.3–43.5 mm, females 37.3–45.0
mm). One female (CARIE 1262) has a substantially longer tail
(148% SL) compared to all other specimens, all of whose tails are
<121% SL. Oval-shaped mental gland present in adult males,
externally visible but not especially prominent; premaxillary

TABLE 4. Mean, standard deviation, and range of 13 morphological measurements and tooth counts for females.

Pseudoeurycea granitum sp. nov. n = 9 Pseudoeurycea lynchi n = 20 Pseudoeurycea nigromaculata n = 10

SVL 41.6 6 3.96 (37.2–50.2) 45.4 6 4.12 (37.3–52.1) 50.7 6 4.78 (43.4–57.7)
TL 49.2 6 8.31 (42.0–63.0) n = 8 45.6 6 5.12 (39.3–53.3) n = 11 54.4 6 8.34 (44.5–62.7)
AX 22.2 6 2.79 (19.6–27.9) 24.4 6 2.53 (20.2–30.0) 27.7 6 3.28 (23.0–33.0)
FLL 9.9 6 1.42 (7.8–12.8) 10.8 6 1.45 (8.1–12.9) 12.1 6 0.98 (11.1–13.7)
HLL 11.2 6 1.44 (9.8–14.6) 12.1 6 1.23 (9.5–14.2) 13.1 6 1.13 (11.5–14.8)
HL 9.5 6 0.84 (8.5–10.9) 10.5 6 0.84 (9.2–11.9) 11.4 6 0.70 (10.3–12.4)
HW 6.3 6 0.59 (5.6–7.4) 7.0 6 0.63 (5.9–8.2) 8.0 6 0.69 (6.9–9.0)
HD 3.0 6 0.36 (2.2–3.5) 3.6 6 0.42 (2.8–4.3) n = 14 4.2 6 0.48 (3.3–5.1)
IO 2.2 6 0.19 (1.9–2.6) 2.4 6 0.25 (2.0–2.9) n = 14 2.7 6 0.26 (2.2–3.2)
IN 1.9 6 0.15 (1.7–2.1) 2.0 6 0.17 (1.6–2.3) n = 14 2.2 6 0.16 (2.0–2.6)
RFW 4.3 6 0.69 (3.4–5.3) 4.5 6 0.80 (3.2–6.0) n = 19 4.9 6 0.88 (3.9–6.4)
T3 2.0 6 0.41 (1.5–2.7) 2.3 6 0.43 (1.7–2.9) n = 14 2.5 6 0.16 (2.2–2.8)
T5 1.1 6 0.17 (0.9–1.4) 1.2 6 0.27 (0.7–1.6) n = 14 1.3 6 0.27 (0.7–1.7)
MT 71.6 6 10.22 (59–86) 75.1 6 10.16 (58–96) n = 19 82.8 6 13.31 (58–96)
VT 24.1 6 1.96 (22–28) 26.5 6 3.86 (19–34) n = 19 28.5 6 5.95 (16–39)

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 264 P. GARCÍA-BAÑUELOS ET AL.

teeth fewer (2–4 vs. 5–10 in females) but more prominent in adult
males, nasolabial protuberances blunt but well developed. Adult
males can be substantially smaller than holotype (smallest male
CARIE 1286, SVL 34.2 mm).
Measurements of the Holotype (in mm) and Tooth Counts.—Head
width 7.4; snout to gular fold (head length) 11.0; head depth at
posterior angle of jaw 3.5; eyelid length 2.9; eyelid width 1.8;
anterior rim of orbit to snout 2.8; horizontal orbit diameter 1.9;
interorbital distance 2.2; snout to forelimb 15.2; distance
separating external nares 2.1; snout projection beyond mandible
0.1; snout to posterior angle of vent (SVL) 50.2; snout to anterior
angle of vent 47.3; axilla to groin 27.9; tail length 58.8; tail width
at base 3.5; tail depth at base 3.8; forelimb length 12.8; hind-limb
length 14.6; width of right manus 3.5; width of right pes 5.2;
length of longest toe 2.7; length of shortest (first) toe 0.7; length of
fifth toe 1.4; maximum nostril diameter 0.5; shoulder width 6.8.
Tooth counts: maxillary teeth 71, premaxillary teeth 8, vomerine
teeth 26.
Coloration in Life of the Holotype.—Dorsal surface of head to
forelimbs Vandyke Brown (281) with Jet Black (300) speckles and
white stipples. Dorsal surface from shoulders to base of tail
Citrine (119) with Olive Yellow (117) speckles and black spots.
Dorsal surface of the tail has Olive Yellow (117) background
combined with Light Lime Green (113) and black spots. Sides of
the body from end of jaw to the insertion of hind limbs Jet Black
(300) with lichen shape Light Greenish White (98) blotches.
Dorsal surface of limbs Citrine (119) combined with Olive Yellow
(117), Light Lime Green (113) and black splotches (Fig. 3A). Gular
region Light Neutral Gray (297), venter to cloaca Jet Black (300)
with Light Greenish White (98) speckles and white stipples. The
ventral surface of tail Olive Yellow (117) combined with Light
Lime Green (113) and black splotches. Light Neutral Gray (297)
underside of hands and feet (Fig. 3D). Iris golden with Olive
Brown (278) reticles.
Coloration in Alcohol.—Dorsum medium gray with pale gray
specks, flanks very dark gray with extensive pale gray mottling.
Dorsal surface of tail medium gray with dark gray spots or
granite-like lichenous blotches of pale gray and a few dark gray
specks near tip. Dorsal side of head dark gray with indistinct
medium gray interorbital bar. Upper side of limbs dark gray with
paler gray mottling. Ventral surface of body, limbs, and gular
region dark gray with numerous narrow specks of pale gray.
Underside of tail medium gray, becoming paler and more
yellowish towards tip, with numerous dark gray spots or
blotches.
Color Variation.—Some specimens have an interorbital bar
that varies from Citrine (119) in adults (Fig. 3E) to white color
or Olive Yellow (117) in young specimens (Fig. 3G,H); this
band can vary from being as broad as the eyelid to a barely
distinguishable narrow band (Fig. 3F). The amount of spotting
on the belly and chin varies between individuals. Some
individuals have a continuum of the dorsal Citrine (119) onto
the lateral portion of the body with small Light Greenish White
(98) speckles (Fig. 3F).
Distribution, Habitat, and Natural History.—Pseudoeurycea gran-
itum sp. nov. is known from three localities: Tepexilotla (Fig. 4A),
Cerro Petlalcala (Fig. 4B), and Mundo Nuevo (Fig. 4C), all in
central Veracruz (Fig. 1). This species occurs in cloud forest and
ecotone with pine–oak forest, in an elevational range between
1,550 and 2,250 m a.s.l. All the individuals from Tepexilotla were
found inactive in bromeliads of the genus Pseudalcantarea aff.
macropetala. The two juveniles observed at this locality were
found in fallen bromeliads and the seven adults collected were
FIG. 4. Forest at Tepexilotla (A), Cerro Petlalcala (B), and Mundo
Nuevo (C) localities. Coffee plantation at Tepexilotla locality (D).
Regurgitated individual of Pseudoeurycea granitum sp. nov. by
Thamnophis sumichrasti (E).
found in bromeliads 1–5 m above ground in trees: three of them
were in a forest fragment and four in a contiguous shaded coffee
plantation (Fig. 4D). The landscape in the vicinity of this locality
is dominated by corn and chayote fields, banana and coffee
plantations, and cattle pastures with small fragments of cloud
forest, mainly located on steep slopes. At another site less than 2
km south of Tepexilotla, no salamanders were found after
inspecting about 200 bromeliads with a search effort of 24 person
hours. The specimens of Cerro Petlalcala were found in a
fragment of highly disturbed cloud forest and ecotone between
cloud forest and pine–oak forest surrounded by cornfields in
different terrestrial microhabitats (under rock, log, moss, under
bark of log, and in a cracked dirt wall). Two juveniles were
observed at this locality under rocks in March and June 2015. At
the Mundo Nuevo locality, after inspecting about 300 bromeliads
on November 2018, with a sampling effort of 15 person hours, we
found two individuals (one juvenile [CARIE 1273, Fig. 3H] and
the gravid female holotype) inactive in bromeliads; in July–
August 2019, after inspecting about 400 bromeliads, with a
sampling effort of 40 person hours, we found five adult
salamanders (CARIE 1285–1288, JLAL872 [to be catalogued at
IBH]) inactive in bromeliads. The seven adults collected on
Mundo Nuevo were found in bromeliads 1–6 m above ground in
trees. Mundo Nuevo is surrounded by fields of corn, beans, and
ornamental plants as well as coffee plantations. The individuals
of Pseudoeurycea granitum sp. nov. were collected throughout the
year (February, March, June, July, August, October, and
November).
During fieldwork in October 2015 on Cerro Petlalcala, an
individual of Pseudoeurycea granitum sp. nov. with a low level of
decomposition was regurgitated by an individual of Thamnophis
sumichrasti captured in a fragment of cloud forest (Fig. 4E). In
Tepexilotla during our fieldwork we observed Aquiloeurycea
cafetalera, Parvimolge townsendi and Thorius pennatulus in
sympatry with P. granitum sp. nov. At Mundo Nuevo we
observed Parvimolge townsendi and Thorius pennatulus. At Cerro

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 NEW SPECIES OF PSEUDOEURYCEA FROM MEXICO
Petlalcala we observed Aquiloeurycea cafetalera, Isthmura gigantea,
and Pseudoeurycea firscheini, in addition to P. granitum sp. nov.
(Kelly-Hernández et al., 2019). The female holotype had ovarian
ova at the time of capture in November. This species might lay
eggs in the first months of the year like other species of the
genus, such as Pseudoeurycea juarezi, which lay eggs clutch in
January or P. exspectata, which lay eggs between March and
April (McDiarmid and Worthington, 1970).
Etymology.—The species name is a noun in apposition meaning
‘‘granite’’ in Latin. The name refers to the coloration of this
species, which resembles granite in its numerous speckles and
blotches.
DISCUSSION
The description of Pseudoeurycea granitum sp. nov. increases
the number of species in the genus to 40 and the species richness
of plethodontid salamanders recorded in Mexico to 133
(AmphibiaWeb, 2019). Pseudoeurycea granitum sp. nov. is very
similar to its close relatives (P. lynchi and P. nigromaculata) in
morphology and nearly identical after accounting for size
differences. This is perhaps not surprising, given that the three
species are semiarboreal and occur in cloud forest habitat within
a relatively small region of central Veracruz. Thus, they are
likely subject to similar selection pressures imposed by both
microhabitat use and climatic conditions, resulting in similar
morphology. Color pattern reliably diagnoses the new species
from its close relatives, however, as does its smaller body size.
Although mtDNA data support its distinctiveness as a species,
the addition of nuclear data for P. granitum sp. nov. and its close
relatives would clarify relationships between species in this
clade.
The high topographic relief of central Veracruz, coupled with
the current allopatric distribution of P. granitum sp. nov., P.
lynchi, and P. nigromaculata, suggests that isolation in allopatry
may have led to divergence of the three species. Cloud forest
distribution depends on the input of cloud water and thus cloud
formation, which in turn depends on both topography and
broader climatic conditions in a region. Past climatic fluctua-
tions could have fragmented a more widely distributed ancestor
occurring in cloud forest into distinct populations, promoting
population divergence and eventually speciation (Ramı́rez-
Barahona and Eguiarte, 2013). Nuclear data would be of great
use to determine whether the timing of divergence of the three
species is more likely explained by climatic fluctuations,
geological changes in the region, or a combination of both
factors.
Although P. lynchi occurs somewhat farther to the north of its
close relatives, P. granitum sp. nov. and P. nigromaculata are
found in near sympatry, with the type locality of P. nigromaculata
(Cerro Chicahuaxtla) between known localities for P. granitum
sp. nov. Decades of fieldwork have been done on Cerro
Chicahuaxtla, and it is unlikely that additional species of
salamanders remain to be sampled there, but the Sierra de
Zongolica (of which Cerro Petlalcala is part) is much less well
known herpetologically. It remains possible that the two species
could occur in sympatry within the Sierra de Zongolica or near
the type locality of P. granitum sp. nov. If they do occur in
sympatry, this would further support the case for recognition of
the new species.
Pseudoeurycea granitum sp. nov. seems to be a species with
low abundance at our study localities. On average, we sampled
for 7.6 person hours to find an individual of P. granitum sp. nov.
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265
in Tepexilotla, 7.8 person hours in Mundo Nuevo, and 44.5
person hours on Cerro Petlalcala. With a sampling effort of 21
person hours carried out in Tepexilotla in November 2018, after
inspecting about 80 bromeliads, no individuals were found.
Similarly, after searching for 80 person hours on Cerro Petlalcala
on August 2019, including the inspection of 37 bromeliads, no
salamanders were found. However, it is necessary to carry out
fieldwork at different times of the year and in surrounding areas
to have a better idea about the abundance or density of this
species. The geographical distribution of P. granitum sp. nov.
seems to be very restricted; less than 30 km separate the
localities where the species has been recorded, and all of them
are in an elevational range between 1,550 and 2,250 m a.s.l.
Forest fragments and shaded coffee plantations located between
the three known localities where P. granitum sp. nov. occurs, or
even nearby localities in Puebla and Oaxaca with similar
environmental conditions, should be searched to detect other
populations and improve the knowledge about their distribu-
tion, as has happened with other salamander species in the
mountainous region of central Veracruz (Sandoval-Comte et al.,
2012).
From a conservation perspective, habitat transformation
appears to be the main threat to P. granitum sp. nov., because
the rate of deforestation in the state of Veracruz is one of the
highest in the country (Muñiz-Castro et al., 2015) and in the
central region forest cover continues to decline (Gómez-Dı́az et
al., 2018). The Tepexilotla and Mundo Nuevo localities are
within the Metlac-Rı́o Blanco state reserve, but the landscape
in the reserve is composed mostly of highly transformed
environments, such as agricultural fields and pastures (INEGI,
2017), where P. granitum sp. nov. is unlikely to occur. We have
observed that previously forested areas both inside and
outside the state reserve have been deforested in recent years.
In fact, a portion of the forest at Tepexilotla and Cerro
Petlalcala where we found salamanders no longer exists; the
forest was cut down in 2018 and 2019, respectively. In addition
to P. granitum sp. nov., six salamander species occur in the
Metlac-Rı́o Blanco state reserve (Garcı́a-Bañuelos et al., 2019),
thus it is necessary to urgently protect the forest fragments that
still remain in the reserve and in the region. Additionally,
because P. granitum sp. nov. was found in shaded coffee
plantations, it is possible that this environment could serve to
maintain populations of this salamander species, as in the case
of several amphibian species in the region (Pineda et al., 2005;
Murrieta-Galindo et al., 2013), but this needs to be evaluated in
the future. Because two of its three known localities have been
partially or completely deforested in the last 2 yr, we suggest
that P. granitum sp. nov. be provisionally classified as
endangered (EN) based on IUCN Red List criteria A2c
(inferred population size reduction of >50% over 10 yr based
on a decline in quality of habitat) and B2ab(iii) (area of
occupancy <500 km2, known to exist at no more than five
localities, and continuing decline in area, extent, and quality of
habitat; IUCN, 2012).
The species richness of plethodontid salamanders in the
mountainous region of central Veracruz (29 species, including
Pseudoeurycea granitum sp. nov.) represents 22% of the number
of species known from Mexico. The number of species known
from this region will continue to increase because there are
species of the genus Pseudoeurycea and Chiropterotriton awaiting
description (Cázares-Hernández et al., 2018; Garcı́a-Castillo et
al., 2018), which highlights the mountainous region of central
 266 P. GARCÍA-BAÑUELOS ET AL.
Veracruz as one of the most remarkable areas in diversity of
salamander species in Mexico.
Acknowledgments.—We thank Itzel Cabrera-Hernández, Ma-
rı́a Chanel Juárez-Ramı́rez, Dania Venerozo-Tlazalo, Arı́s-
tides Garcı́a-Vinalay, Luis Garcı́a-Feria, Eder León-López,
Mauricio Sánchez-Eugenio, Felipe Ramos-Garcı́a, Jorge Ama-
dor González-Pelayo, Anthony Missael Hernández-Garcı́a
and Cenobio Pacheco for the valuable help during fieldwork.
Feliza Ramón-Farı́as and Pedro Adrián Aguilar-Rodrı́guez
helped with the vegetation identification. Esther Quintero-
Rivero and Héctor Gadsden made useful suggestions on
early versions of this manuscript. Two anonymous reviewers
made valuable comments that improved this manuscript.
Scientific collection permits were issued by the Secretarı́a de
Medio Ambiente y Recursos Naturales (SGPA/DGVS/
01833/15 and SGPA/DGVS/03444/15). This study was
supported by the Comisión Nacional para el Conocimiento
y Uso de la Biodiversidad (projects HK006 and JF-212). PGB
was awarded graduate studies scholarship by Consejo
Nacional de Ciencia y Tecnologı́a (grant 261709). This article
was written in partial fulfillment of the requirements of
Graduate Studies Program of the Instituto de Ecologı́a, A.C.
for PGB’s doctoral degree.
LITERATURE CITED
ALJANABI, S. M., AND I. MARTINEZ. 1997. Universal and rapid salt-
extraction of high quality genomic DNA for PCR-based techniques.
Nucleic Acids Research 25:4692–4693.
BUMZAHEM, C. B., AND H. M. SMITH. 1955. Additional notes and
descriptions of plethodontid salamanders from Mexico. Herpetolog-
ica 11:73–75.
CAMPBELL, J. A., E. D. BRODIE JR., J. C. BLANCAS-HERNÁNDEZ, AND E. N.
SMITH. 2013. Another new salamander of the genus Pseudoeurycea
from the State of Guerrero, Mexico. South American Journal of
Herpetology 8:198–202.
CÁZARES-HERNÁNDEZ, E., E. MOLOHUA-TZITZIHUA, E. C. MÉNDEZ-QUIAHUA,
E. QUIAHUA-COLOTL, I. TEMOXTLE-MARQUEZ, J. C. RODRIGO-MERINO, AND
R. APALE-PACHECO. 2018. Tlaconetes. Los hijos de la tierra. Zongolica.
Instituto Tecnológico Superior de Zongolica. México.
CONABIO (COMISIÓN NACIONAL PARA EL CONOCIMIENTO Y USO DE LA
BIODIVERSIDAD), MÉXICO. 1997. Provincias Biogeográficas de México,
escala 1:4,000,000.
EDGAR, R. C. 2004. MUSCLE: a multiple sequence alignment method
with reduced time and space complexity. BMC Bioinformatics 5:1–
19.
GARCÍA-BAÑUELOS, P., S. M. ROVITO, AND E. PINEDA. 2019. Representation of
threatened biodiversity in protected areas and identification of
complementary areas for their conservation: plethodontid salaman-
ders in Mexico. Tropical Conservation Science 12:1–15.
GARCÍA-CASTILLO, M. G., A. F. SOTO-POZOS, J. L. AGUILAR-LÓPEZ, E. PINEDA,
AND G. PARRA-OLEA. 2018. Two new species of Chiropterotriton
(Caudata: Plethodontidae) from central Veracruz, Mexico. Amphib-
ian and Reptile Conservation 12:37–54.
GÓMEZ-DÍAZ, J. A., K. BRAST, J. DEGENER, T. KRÖMER, E. ELLIS, F. HEITKAMP,
AND G. GEROLD. 2018. Long-term changes in forest cover in central
Veracruz, Mexico (1993–2014). Tropical Conservation Science 11:1–
12.
INEGI (INSTITUTO NACIONAL DE ESTADÍSTICA Y GEOGRAFÍA), MÉXICO. 2017.
Conjunto de datos vectoriales de la carta de uso del suelo y
vegetación, escala 1:250,000, Serie VI, Continuo Nacional.
IUCN. 2012. IUCN Red List Categories and Criteria: version 3.1. 2nd ed.
International Union for the Conservation of Nature, Gland,
Switzerland.
KELLY-HERNÁNDEZ, A., S. M. ROVITO, AND V. VÁSQUEZ-CRUZ. 2019. A new
population of the Endangered salamander Pseudoeurycea firscheini
(Caudata: Plethodontidae) from Mexico, with notes on natural
history. Phyllomedusa 18:97–100.
Downloaded From: https://bioone.org/journals/Journal-of-Herpetology on 17 Jul 2020
Terms of Use: https://bioone.org/terms-of-use Access provided by Society for the Study of Amphibians and Reptiles
KÖHLER, G. 2012. Color Catalogue for Field Biologists. Herpeton,
Germany.
LANFEAR, R., P. B. FRANDSEN, A. M. WRIGHT, T. SENFELD, AND B. CALCOTT.
2017. PartitionFinder 2: New methods for selecting partitioned
models of evolution for molecular and morphological phylogenetic
analyses. Molecular Biology and Evolution 34(3):772–773.
LYNCH, J. F., AND D. B. WAKE. 1975. Systematics of the Chiropterotriton
bromeliacia group (Amphibia: Caudata), with descriptions of two
new species from Guatemala. Contributions in Science, Natural
History Museum of Los Angeles County 265:1–45.
MCDIARMID, R. W., AND R. D. WORTHINGTON. 1970. Concerning the
reproductive habitats of tropical plethodontid salamanders. Herpe-
tologica 26:57–70.
MORITZ, C., C. J. SCHNEIDER, AND D. B. WAKE. 1992. Evolutionary
relationships within the Ensatina eschscholtzii complex confirm the
ring species interpretation. Systematic Biology 41:273–291.
MUÑIZ-CASTRO, M. A., G. WILLIAMS-LINERA, AND J. BENÍTEZ-MALVIDO. 2015.
Restoring montane cloud forest: Establishment of three Fagaceae
species in the old fields of central Veracruz, Mexico. Restoration
Ecology 23:26–33.
MURRIETA-GALINDO, R., F. LÓPEZ-BARRERA, A. GONZÁLEZ-ROMERO, AND G.
PARRA-OLEA. 2013. Matrix and habitat quality in a montane cloud-
forest landscape: amphibians in coffee plantations in central
Veracruz, Mexico. Wildlife Research 40:25–35.
PALUMBI, S., A. MARTIN, S. ROMANO, W. O. MCMILLAN, L. STICE, AND G.
GRABOWSKI. 1991. The simple fool’s guide to PCR, version 2.0.
University of Hawaii, USA.
PARRA-OLEA, G. 2002. Molecular phylogenetic relationships of Neotrop-
ical salamanders of the genus Pseudoeurycea. Molecular Phyloge-
netics and Evolution 22:234–246.
PARRA-OLEA, G., T. J. PAPENFUSS, AND D. B. WAKE. 2001. New species of
lungless salamanders of the genus Pseudoeurycea (Amphibia:
Caudata: Plethodontidae) from Veracruz, Mexico. Scientific Papers,
Natural History Museum, The University of Kansas 20:1–9.
PARRA-OLEA, G., S. M. ROVITO, L. MÁRQUEZ-VALDELAMAR, G. CRUZ, R.
MURRIETA-GALINDO, AND D. WAKE. 2010. A new species of Pseu-
doeurycea from the cloud forest in Veracruz, México. Zootaxa 2725:
57–68.
PINEDA, E., C. MORENO, F. ESCOBAR, AND G. HALFFTER. 2005. Frog, bat, and
dung beetle diversity in the cloud forest and coffee agroecosystems
of Veracruz, Mexico. Conservation Biology 19:400–410.
RAFFAËLLI, J. 2013. Les Urodèles du Monde (2e édition). Penclen Édition,
France.
RAMÍREZ-BARAHONA, S., AND L. EGUIARTE. 2013. The role of glacial cycles
in promoting genetic diversity in the tropics: the case of cloud
forests during the last glacial maximum. Ecology and Evolution 3:
725–738.
RONQUIST, F., M. TESLENKO, P. VAN DER MARK, D. L. AYRES, A. DARLING, S.
HOHNA, B. LARGET, L. LIU, M. A. SUCHARD, AND J. P. HUELSENBECK.
2012. MrBayes 3.2: Efficient Bayesian phylogenetic inference and
model choice across a large model space. Systematic Biology 61:
539–542.
SANDOVAL-COMTE, A., E. PINEDA, AND J. L. AGUILAR-LÓPEZ. 2012. In search
of critically endangered species: the current situation of two tiny
salamander species in the neotropical mountains of Mexico. PLoS
ONE 7:e34023.
STAMATAKIS, A. 2014. RAxML version 8: a tool for phylogenetic analysis
and post-analysis of large phylogenies. Bioinformatics 30:1312–
1313.
SWOFFORD, D. L. 1998. Paup 4.0 Beta Version for Windows: Phylogenetic
Analysis Using Parsimony. Sinauer Associates, USA.
WAKE, D. B., T. J. PAPENFUSS, AND J. F. LYNCH. 1992. Distribution of
salamanders along elevational transects in Mexico and Guatemala.
Tulane Studies in Zoology and Botany, Supplementary Publication 1:
303–319.
Accepted: 28 September 2019.
Published online: 16 July 2020.
APPENDIX 1
Specimens Examined.—Pseudoeurycea granitum sp. nov.: Mexico:
Veracruz: Chocamán Municipality: Metlac River near Tepexilotla,
CARIE 1261–1265; Mundo Nuevo: CARIE 1274, 1285–1288, IBH
32550; San Andrés Tenejapan Municipality: east slope of Cerro
Petlalcala, IBH 31826, 31827.
 NEW SPECIES OF PSEUDOEURYCEA FROM MEXICO 267

Pseudoeurycea lynchi—Mexico: Veracruz: Acajete Municipality: Pseudoeurycea nigromaculata—Mexico: Veracruz: Ixtaczoquitlán

forest west of La Joya, MVZ 158821, 158824–158825, 178841, 203670, Municipality: Cerro Chicahuaxtla, SMR32549, MVZ 86083, 86084,
203672, 203674; Chiconquiaco Municipality: Loma Alta microwave 106562, 106563, 106565, 106568, 106569, 118958, 129948, 150534,
station, MVZ 230994–230999, 231001, IBH 32544–32548; Coatepec 186707, CARIE 1085, 1087; Huatusco Municipality: Granja Los
Municipality: La Cortadura, CARIE 1099, 1102. Naranjos, CARIE 1063.

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