Corresponding author: Luis Canseco Márquez (
Academic editor: Uwe Fritz
We describe a new species of plethodontid salamander of the genus
Se describe una nueva especie de salamandra pletodóntida del género
Mexico has the world’s seventh highest diversity of amphibian species (418 species) and is second only to the USA in salamander species diversity (158 species;
Following a taxonomic revision of the
With 42 species of plethodontid salamanders (listed by
In a coniferous forest (
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 number allowed by the SEMARNAT collecting permit) and deposited in the Museo de Zoología Alfonso L. Herrera, Facultad de Ciencias, Universidad Nacional Autónoma de México (
The description of the new species follows the format used by
We compared the new species with all currently recognized members of the
Mean, standard deviation, and range of 13 morphological measurements and tooth counts for males of species of the
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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) |
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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) |
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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) |
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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) |
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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) |
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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) |
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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) |
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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) |
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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) |
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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) |
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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) |
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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) |
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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) |
69±3.16 (66–73) | 85.5±13.43 (76–95) | 67±10.68 (49–90) | 31 | 67±20.5 (52–81) | |
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23±4.54 (17–27) | 30.5±7.77(25–36) | 22±2.86 (17–27) | 17 | 26±2.82 (24–28) |
Mean, standard deviation, and range of 13 morphological measurements and tooth counts for females of species of the
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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) |
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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 |
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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) |
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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) |
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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) |
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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) |
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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) |
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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) |
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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) |
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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) |
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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) |
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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) |
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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) |
74±9.89 (67–81) | 92.6±20.25 (67–116) | 73±18.05 (26–96) | — | 73±8.71 (67–83) | |
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22±1.41 (21─23) | 30.3±7.06 (18–37) | 23.2±2.79 (19–26) | — | 22±2.64 (19–23) |
We extracted DNA from liver tissue using a high salt protocol (
We obtained 16S and cyt
Voucher information and GenBank accession numbers for sequences used in phylogenetic analysis.
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CRVA1017 |
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GP160 |
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GP0289 |
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GP372 |
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AMA2534 |
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We used RAxML v8.2 (
Suggested English name: Jaguar Salamander.
Suggested Spanish name: Tlaconete jaguar.
Seven. One male:
Assigned to the genus
Morphologically, we distinguish the new species from the other salamanders that occur in the region and from the others of the genus
Based on mtDNA, this new species is closely related to members of the
The new species is further distinguished from all members of the
Live specimens of all members of the
A relatively large adult male (58.7
Snout to posterior angle of vent (
The type series includes eight specimens, two males and six females. There is marked sexual dimorphism; adult females reach a larger size than males (
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 dorsum and even larger and more continuous on tail. Sides of head brown with yellow speckling, with the same proportion 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 flecks (small flecks combined with larger and elongated flecks) 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 flecks. Iris dark brown with yellow specks around the pupil.
Color pattern of
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.
The color pattern is similar in most adult specimens. 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 become larger along the dorsum and even larger on the tail, but their size varies from specimen to specimen (Fig.
Other species of plethodontids that share habitat with
Antagonistic behavior between individuals in a population has been described in several species of plethodontid salamanders (
The specific epithet
The results of our mtDNA phylogenetic analysis (Fig.
Generalized time-reversible (GTR) distances between species of the
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0.034 | 0.043 | 0.049 | 0.047 | 0.043 | 0.056 |
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0.043 | 0.041 | 0.041 | 0.043 | 0.041 | 0.043 | 0.038 | 0.043 |
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— | 0.036 | 0.045 | 0.052 | 0.043 | 0.047 |
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0.048 | 0.050 | 0.050 | 0.038 | 0.038 | 0.038 | 0.034 | 0.034 |
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0.172 | — | 0.023 | 0.030 | 0.023 | 0.036 |
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0.034 | 0.037 | 0.037 | 0.041 | 0.039 | 0.036 | 0.032 | 0.038 |
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— | — | — | 0.032 | 0.030 | 0.043 |
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0.038 | 0.041 | 0.041 | 0.047 | 0.050 | 0.047 | 0.039 | 0.052 |
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0.201 | 0.137 | — | — | 0.023 | 0.050 |
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0.045 | 0.048 | 0.048 | 0.059 | 0.061 | 0.052 | 0.050 | 0.052 |
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0.199 | 0.100 | — | 0.090 | — | 0.038 |
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0.039 | 0.041 | 0.041 | 0.045 | 0.048 | 0.045 | 0.041 | 0.050 |
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0.180 | 0.191 | — | 0.192 | 0.204 | — |
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0.025 | 0.028 | 0.028 | 0.050 | 0.057 | 0.052 | 0.043 | 0.061 |
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0.150 | 0.175 | — | 0.166 | 0.154 | 0.165 |
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— | 0.006 | 0.006 | 0.050 | 0.050 | 0.047 | 0.036 | 0.047 |
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0.158 | 0.182 | — | 0.159 | 0.160 | 0.146 |
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0.037 | — | 0.000 | 0.043 | 0.050 | 0.045 | 0.034 | 0.045 |
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0.161 | 0.193 | — | 0.166 | 0.171 | 0.162 |
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0.043 | 0.037 | — | 0.043 | 0.050 | 0.045 | 0.034 | 0.045 |
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0.113 | 0.187 | — | 0.204 | 0.188 | 0.171 |
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0.158 | 0.174 | 0.185 | — | 0.032 | 0.036 | 0.027 | 0.038 |
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0.117 | 0.172 | — | 0.201 | 0.199 | 0.180 |
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0.150 | 0.158 | 0.161 | 0.113 | — | 0.045 | 0.030 | 0.038 |
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0.193 | 0.156 | — | 0.153 | 0.167 | 0.173 |
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0.184 | 0.179 | 0.184 | 0.175 | 0.193 | — | 0.030 | 0.034 |
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0.165 | 0.193 | — | 0.169 | 0.186 | 0.135 |
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0.176 | 0.173 | 0.176 | 0.180 | 0.165 | 0.170 | — | 0.021 |
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0.157 | 0.168 | — | 0.166 | 0.162 | 0.161 |
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0.189 | 0.193 | 0.195 | 0.171 | 0.157 | 0.175 | 0.083 | — |
Phylogeny estimated from maximum likelihood analysis of 16S and cyt
Based on mtDNA,
Because of its arboreal habits,
Based on our direct observations and with the best available evidence on salamander communities in the region, we can affirm that
The species diversity of the Sierra de Zongolica is high, and it is important to continue exploring other areas, since several records and new species have recently been discovered (
We thank Rafael Rosales Martínez, Julián Rodríguez Lobato and Mauro Daniel Castro Morales, students of the ITSZ Forest Engineering career, for their collaboration in fieldwork; Ángel Iván Contreras Calvario from Facultad de Biología of the Universidad Veracruzana for his support and collaboration in the fieldwork; Ing. Wenceslao Cosme Reyes, Coordinator of the Zongolica campus of the ITSZ, for his support in relation to field trips and finally especially to the Instituto Tecnológico Superior de Zongolica. Víctor Vásquez Cruz helped with the map. Collecting permits SEMARNAT-08-049, Oficio Núm.SGPA/DGVS/02924/15 were issued to Erasmo Cázares-Hernández and collaborators. Three reviewers provided valuable comments that improved the quality of the manuscript.
Specimens examined:
Institutional abbreviations for museums and collections follow