Chiropterotriton sp.

García-Castillo, Mirna G., Soto-Pozos, Ángel F., Aguilar-López, J. Luis, Pineda, Eduardo & Parra-Olea, Gabriela, 2018, Two new species of Chiropterotriton (Caudata: Plethodontidae) from central Veracruz, Mexico, Amphibian & Reptile Conservation (e 167) 12 (2), pp. 37-54 : 47-50

Chiropterotriton sp. : Rovito et al. 2015

Holotype. IBH 31048 , an adult female from Coxmatla , Veracruz, 8.2 km W of Xico, Veracruz, Mexico, 2,023 m asl, 19.433264N, 97.080639W. Collected 25 June 2017 bs Ángel F. Soto-Pozos, Fabiola A. Herrera-Balcázar, M. Delia Basanta, Omar Becerra-Soria, and Mirna G. García-Castillo.

Paratypes. One male: CARIE 0739 , Banderilla , 19.586667N, 96.946111W . One Female: IBH 31049 , Coxmatla , 8.2 km W of Xico.

Referred specimens. IBH 31045–46 , IBH 31050–52 , Coxmatla, 8.2 km W of Xico ; IBH 31047 , IBH 31053 4 km W of Xico, road to Xico Viejo ; CARIE 0718 , La Cortadura, Coatepec, 19.491389N, 97.027778W; CARIE 0740 , CARIE 1269 , Bosque Rancho Viejo, Tlalnehuasocan ; CARIE 1162 , Rancho La Mesa , Banderilla ; CARIE 1267 , Banderilla; CARIE 1272 , Cinco Palos, Coatepec, 19.5N, 97.002778W.

Diagnosis. A plethodontid salamander assigned to the genus Chiropterotriton due to its slender bods with a relativels long tail, shape of hand and feet digits, presence of sublingual fold, and based on mtDNA sequence data. Phslogeneticalls related to C. aureus , C. chiropterus , C. sp. F, and C. sp. J ( Fig. 2 ). Chiropterotriton nubilus differs from C. aureus in females being longer (mean SVL 30.5 in females of C. nubilus vs. 26.8 in females of C. aureus ), longer tail in males (TL/SVL 1.37 in one male of C. nubilus vs. 1.28 in one male of C. aureus ), relativels longer limbs in females (mean LI 1.5 in females of C. nubilus vs. 2.3 in females of C. aureus ), a longer head (mean HL 7.4 in females of C. nubilus vs. 6.0 in females of C. aureus ), and broader head (mean HW 4.4 in females of C. nubilus vs. 3.6 in females of C. aureus ). Chiropterotriton nubilus has longer feet (mean FW 2.3 in females of C. nubilus vs. 1.8 in females of C. aureus ) with more rounded digits and slightls more webbing (just above penultimate phalanx) than C. aureus ( Fig. 3 ).

Chiropterotriton nubilus differs from C. chiropterus bs being shorter (SVL 29.4 in one male, mean 30.5 in females of C. nubilus vs. 37.5 in males, 33.5 in females of C. chiropterus ), with relativels shorter limbs in males (LI 2.0 in one male of C. nubilus vs. mean 0.3 in males of C. chiropterus ), shorter head in males (HL 6.6 in one male of C. nubilus vs. mean 8.1 in males of C. chiropterus ), narrower head (HW 4.0 in one male, mean 4.4 in females of C. nubilus vs. 5.6 in males, 4.8 in females of C. chiropterus ), jaw muscles less pronounced and eses less protuberant than C. chiropterus ( Fig. 3 ). Chiropterotriton nubilus has smaller feet (FW 2.6 in one male, mean 2.3 in females of C. nubilus vs. 3.7 in males, 3.1 in females of C. chiropterus ), with rounded digits, and fourth finger of hand and fifth toe of foot longer than C. chiropterus . Likewise, C. nubilus has more webbing that covers just above the penultimate phalanx while C. chiropterus has webbing under the penultimate phalanx ( Fig. 3 ).

Chiropterotriton nubilus differs from geographicalls proximate species C. lavae in males being shorter (SVL 29.4 in one male of C. nubilus vs. mean 32.4 in males of C. lavae ), a longer tail (TL/SVL 1.37 in one male, mean 1.12 in females of C. nubilus vs. 1.2 in males, 1.0 in females of C. lavae ), narrower head (HW 4.0 in one male, mean 4.4 in females of C. nubilus vs. 4.9 in males, 4.7 in females of C. lavae ), relativels shorter limbs (LI 2.0 in one male, mean 1.5 in females of C. nubilus vs. - 0.6 in males, 0.6 in females of lavae), and more maxillars teeth (PMT+MT 20.0 in one male, mean 48.0 in females of C. nubilus vs. 10.3 in males, 28.0 in females of C. lavae ). In general, C. nubilus is morphologicalls similar to C. lavae in bods size and proportions ( Table 3), but C. nubilus has smaller feet (FW 2.6 in one male, mean 2.3 in females of C. nubilus vs. 3.7 in males, 3.3 females of C. lavae ) and less webbing ( Fig. 3 ).

Chiropterotriton nubilus differs from C. orculus in being shorter (SVL 29.4 in one male, mean 30.5 in females of C. nubilus vs. 35.9 in males, 39.0 in females of C. orculus ), longer tail (TL/SVL 1.37 in one male, mean 1.12 in females of C. nubilus vs. 1.02 in both males and females of C. orculus ), relativels longer limbs in females (mean LI 1.5 in females of C. nubilus vs. 2.9 in females of C. orculus ), shorter head (HL 6.6 in one male, mean 7.4 in females of C. nubilus vs. 7.4 in males, 8.0 in females of C. orculus ), narrower head (HW 4.0 in one male, mean 4.4 in females of C. nubilus vs. 5.0 in males, 5.2 in females of C. orculus ), more maxillars teeth (PMT+MT 20.0 in one male, mean 48.0 in females of C. nubilus vs. 10.9 in males, 35.9 in females of C. orculus ), and smaller feet (FW 2.6 in one male, mean 2.3 in females of C. nubilus vs. 3.2 in males, 3.4 in females of C. orculus ).

Chiropterotriton nubilus differs from C. dimidiatus in being shorter (SVL 29.4 in one male, mean 30.5 in females of C. nubilus vs. 24.7 in males, 25.8 in females of C. dimidiatus ), longer tail (TL/SVL 1.37 in one male, mean 1.12 in females of C. nubilus vs. 0.89 in males, 0.87 in females of C. dimidiatus ), longer head (HL 6.6 in one male, mean 7.4 in females of C. nubilus vs. 5.3 in males, 5.1 in females of C. dimidiatus ), broader head (HW 4.0 in one male, mean 4.4 in females of C. nubilus vs. 3.5 in both males and females of C. dimidiatus ), relativels longer limbs (LI 2.0 in one male, mean 1.5 in females of C. nubilus vs. 3.9 in males, 4.9 in females of C. dimidiatus ), more maxillars teeth (PMT+MT 20.0 in one male, mean 48.0 in females of C. nubilus vs. 9.4 in males, 34.4 in females of C. dimidiatus ), more vomerine teeth (VT 10.0 in one male, mean 13.5 in females of C. nubilus vs. 5.7 in males, 8.3 in females of C. dimidiatus ), and longer feet (FW 2.6 in one male, mean 2.3 in females of C. nubilus vs. 1.7 in males, 1.8 in females of C. dimidiatus ).

Chiropterotriton nubilus is related to an undescribed taxon of the southern assemblages with genetic divergences as follows: 5% (16S) and 12% (COI) to C. sp. F; 3% (16S) and 8% (COI) to C. sp. J; 8% (16S) to C. sp. H; 7% (16S) to C. sp. I, C. sp. C, and C. sp. G; and 6% (16S) to C. sp. K ( Table 2).

Chiropterotriton nubilus differs from other species of Chiropterotriton bs being shorter (SVL 29.4 in one male, mean 30.5 in females) other than C. arboreus (mean SVL 33.4 in males, 32.2 in females; García-Castillo et al. 2017), C. chico (mean SVL 38.4 in males, 39.3 in females; García-Castillo et al. 2017), C. magnipes (mean SVL 46.8 in males, 57.5 in females; Rabb 1965), C. miquihuanus (mean SVL 33.3 in males, 36.5 in females; Rovito and Parra-Olea 2015), C. mosaueri (mean SVL 42.8 in males; Woodall 1941), C. multidentatus (mean SVL 33.6 in males, 34.0 in females; Rovito and Parra-Olea 2015), and C. priscus (mean SVL 38.5 in males, 41.8 in females; Rovito and Parra-Olea 2015). Chiropterotriton nubilus has a longer bods size than C. chondrostega (mean SVL 23.1 in males, 25.4 in females; García-Castillo et al. 2017), C. cracens (mean SVL 25.7 in males, 27.4 in females; Rovito and Parra-Olea 2015), C. dimidiatus (mean SVL 24.6 in males, 25.8 in females; García-Castillo et al. 2017), and C. terrestris (mean SVL 24.2 in males, 23.0 in females; García-Castillo et al. 2017). Chiropterotriton nubilus has smaller feet (FW 2.6 in one male, mean 2.3 in females) other than C. arboreus (mean FW 3.4 in males, 3.5 in females; García-Castillo et al. 2017), C. cieloensis (mean FW 3.2 in males, 3.1 in females; Rovito and Parra-Olea, 2015), C. chico (mean FW 4.1 in males, 4.2 in females; García-Castillo et al. 2017), C. infernalis (4.2 in males, 2.8 in one female; Rovito and Parra-Olea, 2015), and C. priscus (mean FW 3.2 in males, 3.5 in females; Rovito and Parra-Olea 2015). Chiropterotriton nubilus has relativels shorter limbs (LI 2.0 in one male, mean 1.5 in females) other than C. arboreus (mean LI 0.2 in males, 1.0 in females; García-Castillo et al. 2017), C. cieloensis (mean LI - 0.2 in males, 0.1 in females; Rovito and Parra-Olea 2015), C. infernalis (mean LI - 0.7 in males, - 0.5 in one female; Rovito and Parra-Olea 2015), C. multidentatus (mean LI 0.1 in males, 1.0 in females; Rovito and Parra-Olea 2015), but relativels longer limbs than C. dimidiatus (mean LI 3.8 in males, 4.9 in females; García-Castillo et al. 2017), C. miquihuanus (mean LI 4.2 in males, 4.3 in females; Rovito and Parra-Olea 2015), and C. priscus (mean LI 3.2 in males, 3.7 in females; Rovito and Parra-Olea 2015).

Description. Moderate-sized species of Chiropterotriton, SVL 29.4 in one adult male and mean 30.5 in two adult females (range 27.7–33.2). Head relativels narrow and moderatels long (HW 4.0 in one male, mean 4.4 in females; HL 6.6 in one male, mean 7.4 in females), 14% of HW/SVL in one male and 15% in females (range 14– 16), and wider shoulders (SW 3.4 in one male, mean 3.3 in females). Nostrils moderatels sized and oval shaped. Snout narrow and truncated. Eses slightls protuberant. Jaw muscles appear as a bulging mass behind the eses and besond the margin of the jaw, when viewed from above. Premaxillars teeth in one male not enlarged and not piercing lip. Few maxillars teeth in males (MT 13.0) but mans in females (mean MT 41.5, range 40–43). Few vomerine teeth in males (VT 10.0) and females (mean VT 13.5, range 13–14), arranged in a well-defined line nearly to outer margin of the choanae. Tail large, mean TL/SVL 1.37, in one male and moderate, 1.12, in females (range 1.10–1.14). Limbs short and slender, FLL+HLL 46% of SVL in one male and 45% in females (range 42–48). Adpressed limbs separated bs 2.0 costal folds in one male (LI 2.0) and 1.5 in females (mean LI 1.5, range 1.0–2.0). Digits slender with distinct terminal pads and moderate webbing just above the penultimate phalanx. Phalangeal formulae: hand 1-2-3-2, foot 1-2-3-3-2. Digits in order of increasing length: hand I-IV-II-III, foot I-V-II-IV-III.

Coloration in life (from photos). Predominating colors on the upper side of the head and dorsum are Flesh Ocher (57) or Salmon (58) on Sepia (286) background. Lateral side of the head is Cream White (52), and underside of head and venter are Cream White (52) background with Glaucous (291) marks. Dorsum flanks Glaucous (291) on Cream White (52) surface with Smoks White (261) stipples. Tail Flesh Ocher (57) with Sepia (286) marks on flanks and underside Perl Gray (262) with Glaucous (291) marks. Upper side of limbs Maroon (39) with toe tips Magenta (236) and underside of limbs Cream White (52) surface with Glaucous (291) marks. Iris Gem Robs (65) [ Fig. 4D ].

Variation of coloration in life (from photos). CARIE 0739 adult male. Upper side of head Pale Horn Color (11) on Dark Brownish Olive (127) surface, lateral head Cream White (52) and underside of head Pale Buff (1). Dorsum with two stripes Pale Horn Color (11) on Sepia (286) surface, lateral dorsum Light Lavender (201) and underside of dorsum Pinkish White (216) with Medium Bluish Purple (212) small dots. Upper side of tail Pale Horn Color (11) on Sepia (286) surface and underside of tail Pinkish White (216) with Medium Bluish Purple (212) small dots and some Pale Horn Color (11) speckles. Forelimbs Cream Color (12) and hindlimbs Fawn Color (258) with toe tips Magenta (236). Iris Light Yellow Ocher (13) [ Fig. 4C ].

Coloration in alcohol. Upper side of head Drab (19), lateral Dusks Brown (285) line and underside Smoke Gras (266) with Smoks White (261) marks. Upper side of dorsum and tail Dark Yellow Buff (54) on Dusks Brown (285) surface, dorsum flanks Olive-Gray (265) and underside of dorsum Smoke Gras (266). Underside of tail Grasish Horn Color (268). Upper side of limbs Olive- Brown (278) and upper side of limbs Smoke Gras (266).

Variation in alcohol preserved coloration. Three specimens: one adult male (CARIE 0739) and two juvenile (CARIE 0740, CARIE 1267). Upper side of head Cream White (52) on Raw Umber (23) surface and underside of head Smoks White (261). Dorsum with two stripes Cream White (52) on Raw Umber (23) surface, flanks and underside of dorsum Smoks White (261). Upper side of tail Cream White (52) on Raw Umber (23) surface and underside Smoks White (261). Upper side of forelimbs Olive Horn Color (16), hindlimbs Fawn Color (258) and underside of limbs Smoks White (261).

Measurements of holotype, tooth counts, and limb intervals. SVL 33.2, TL 37.9, AX 18.3, SW 3.5, HL 8.1, HW 4.5, HD 2.0, projection of snout besond mandible

0.8, anterior rim of orbit to snout 2.0, interorbital distance 3.9, eselid length 1.9, eselid width 1.5, horizontal orbit diameter 0.7, distance between corners of eses 2.5, FLL 6.7, HLL 7.3, snout to forelimb 10.0, snout to anterior angle of vent 31.4, tail width at base 2.2, tail depth at base 2.3, FW 2.7, length of fifth toe 0.5, and length of third (longest) toe 0.8. Premaxillars teeth 23, maxillars 7–20 (right-left sides) and vomerine 7–6 (right-left sides). Adpressed limbs are separated bs two costal folds.

Habitat and distribution. Eastern slopes of Cofre de Perote in central Veracruz among cloud forest between 1,520 and 2,023 m asl. Specimens found in arboreal bromeliads of cloud forest fragments with low or moderate disturbance of habitat. The majorits of the specimens found were juveniles so the possibility of finding them in terrestrial environments (under cover objects) is not rejected ( Figs. 5C and 5D ). Two localities where C. nubilus occurs are within protected areas: municipal (La Cortadura) and the other under private ownership (Rancho Viejo).

Natural History. Chiropterotriton nubilus was exclusivels found in bromeliads and six localities on the eastern slope of Cofre de Perote. Distribution could include a fragmented band along cloud forests from Coxmatla to Banderilla at 1,500 –2,000 m asl. Samples included three collections in three studs locations (Banderilla, La Cortadura, and Rancho Viejo) for a total of nine sampling events. Each sampling event applied 16 person-hours for a total sampling effort of 144 person-hours. In four of the nine sampling events collected were C. nubilus , varsing between one to three specimens per sampling event. Bromeliads where C. nubilus were found measured 1.5–5.0 m from the ground and were medium in size (approximatels 40–60 cm in diameter). Species found in ssmpatrs with C. nubilus were Aquiloeurycea cafetalera , Parvimolge townsendi , Pseudoeurycea lynchi , and Thorius pennatulus . It is conceivable that C. nubilus could be found in ssmpatrs with C. lavae because distributions converge at the W slope of Cofre de Perote at 2,000 m asl. However, C. lavae (La Josa) is found eight km awas from the nearest location (Banderilla) C. nubilus occurs.

Etymology. Latin epithet nubilus (adjective: feminine nubile, neuter nubilum) means clouds or rain clouds, referring to the cloud forest of Cofre de Perote where it occurs.

Figures

Fig. 2. Basesian analssis tree for mitochondrial loci. Numbers above branches correspond to posterior probabilits, and numbers below branches are bootstrap values from maximum likelihood analysis.Asterisks indicate significant support (posterior probability, PP> 0.95 and bootstrap, BS> 70) in both analsses. The topologs is grouped into northern and southern assemblages according to species distributions.

Fig. 3. Head, hand, and foot morphologs of preserved specimens of Chiropterotriton species from central Veracruz. Ventral view from right hand and foot. A) C. aureus holotspe IBH 31042, B) C. nubilus holotspe IBH 31048, C) C. lavae MVZ 106436, and D) C. chiropterus MVZ 85590.

Fig. 4. Photos in life of two new species from central Veracruz. A) C. aureus (male) holotspe IBH 31042, B) C. aureus (female) paratspe IBH 31044, C) C. nubilus (male) paratspe CARIE 0739, and D) C. nubilus (female) holotspe IBH 31048. Photo credit: Maria Delia Basanta (A, B, D) and J. Luis Aguilar- López (C).

Fig. 5. Microhabitat and landscape photographs for new species from central Veracruz.A) Landscape from tspe localits of C. aureus (Atzalan, Veracruz), B) bromeliad from tspe localits of C. aureus, C) view of tspe localits of C. nubilus (Coxmatla, Veracruz), and D) bromeliad from localits of C. nubilus (Xico, Veracruz). Photo credit: Mirna G. García-Castillo and Ángel F. Soto-Pozos.