Bolitoglossa tamaense, Acevedo, Aldemar A., Wake, David B., Márquez, Roberto, Silva, Karen, Franco, Rosmery & Amézquita, Adolfo, 2013

Acevedo, Aldemar A., Wake, David B., Márquez, Roberto, Silva, Karen, Franco, Rosmery & Amézquita, Adolfo, 2013, Two New Species of Salamanders, Genus Bolitoglossa (Amphibia: Plethodontidae), from the Eastern Colombian Andes, Zootaxa 3609 (1), pp. 69-84 : 75-79

Bolitoglossa tamaense , sp. nov.

Tama salamander, Salamandra del Tama ( Figure 3 ).

Holotype. MCNUP 50, an adult male from La Asiria de Belén, Parque Nacional Natural Tamá (PNNT) (7.319278, -72.374778) at 2,700 m elevation, Departamento de Norte de Santander, Colombia. The specimen was collected by Aldemar A. Acevedo, Karen Silva and Rosmery Franco on August 2010.

Paratypes. MCNUP 51–53, adult males with same data as holotype, MCNUP 54–55, adult females with same data as holotype. MCNUP 56–57, adult males and MCNUP 58–61, adult females, collected 2 km away from Los Remansos, Provincia Toledo, (7.330888,-72.475472), at 2,000 m elevation, Departamento de Norte de Santander, Colombia. All collected by Aldemar A. Acevedo, Karen Silva and Rosmery Franco on October 2010.

Diagnosis. A member of the genus Bolitoglossa because of the absence of a sublingual fold and presence of extensive digital webbing and 13 costal grooves between the limbs. A species from the high Andean forests of the eastern flank of the Eastern Colombian Andes (Cordillera Oriental). Distinguished from all other members of South American Bolitoglossa by a combination of coloration, morphological and molecular characters ( Fig. 2 ). Adult individuals exhibit highly distinctive coloration: dorsal and caudal region with orange shades in females and yellow to brown in males. Moderate size (maximum SVL 52.7) with a short snout, short limbs and moderate to extensively webbed hands and feet with longest digits broadly triangular at tip and bluntly pointed ( Fig. 4 ). Males with rounded hedonic mental gland, not prominent. This species is apparently sexually dimorphic in size, mean SVL= 37.2 mm in males and 47 mm in females. Bolitoglossa tamaense can be further distinguished from other South American species of Bolitoglossa (Eladinea) as follows. It is much smaller than B. capitana , which also has more extensively webbed feet and is uniformly black. It is smaller and more extensively webbed than B. adspersa , B. guaramacalensis , B. hypacra , B. tatamae and B. vallecula . It is larger and more extensively webbed than B. orestes and B. spongai . It is larger and has less extensively webbed feet than B. altamazonica , B. biseriata , B. chica , B. digitigrada , B. paraensis , B. peruviana . B. medemi . B. hiemalis and B. walker i. It is a little larger (females) than B. guaneae , has less webbed feet with longest digits more rounded and less pointed at tips, and a tail longer rather than shorter than SVL. It is smaller and has less webbed feet than B. borburata , B. ramosi , B. pandi , B. nicefori and B. lozanoi . Lack of dermal subterminal pads on the ventral surface of the digits differentiates B. tamaense from the highland species B. adspersa , B. hiemalis , B. hypacra , B. savagei and B. vallecula . It has more maxillary teeth (males mean=35; females mean=40) and more vomerine teeth (males mean =18; females mean=21) than B. hiemalis , B. orestes , B. spongai , B. palmata , B. pandi , B. altamazonica , B. sima , B. peruviana , B. chica , B. lozanoi , and B. leandrae sp. nov. It has fewer maxillary teeth and fewer vomerine teeth that B. vallecula , B. guaramacalensis , B. savage i, B. tatamae , B. ramosi , B. capitana , B. nicefori , B. borburata , B. medemi , B. silverstonei and B. biseriata . While B. asdpersa , B. walkeri and B. equatoriana have fewer maxillary teeth (>35), they have more vomerine teeth (<18).

Description. A medium-sized species, SVL ranges from 39.2 to 52.7 mm (mean= 47 mm) for six females, and from 36.2 to 40.3 mm (mean= 37.2 mm) for six males. Maxillary teeth of moderate size, range from 38 to 39 (mean=39) in males, and from 39 to 42 (mean=40) in females. Vomerine teeth range from 17 to 19 (mean=18) in males, and from 19 to 23 (mean=21) in females. The trunk is relatively long, ranging from 23.8 to 34.7 mm (mean= 29.3 mm) in females, and from 22.1 to 24.5 mm (mean= 23.5 mm) in males. Distance across the shoulders is 6.0 to 7.5 mm (mean= 6.6 mm) in females, and 4.4 to 5.7 mm (mean= 5.1 mm) in males. Tails are long and generally slender, usually exceeding standard length (from 1.0 to 1.1 mm, mean= 1.1 mm, in males and from 0.9 to 1.1 mm, mean=1.0 mm, in females. The head is moderately broad, from 7.0 to 7.7 mm (mean= 7.6 mm) in females, and 6.9 to 7.2 mm (mean= 7.1 mm) in males. Hind limbs are relatively long (8.4–9.8 mm, mean= 9.1 mm in males; 9.5–11.2 mm, mean= 10.1 mm in females). Moderately webbed feet bearing subterminal pads on digits 2–3–4. Fingers, in order of decreasing length, are 3–4–2–1; toes are 3–4–5–2–1 ( Fig. 4 ).

Measurements and morphology of holotype. Male (MCNUP 50) SVL 40.3 mm; head width 7.5 mm; snout to gular fold 10.5 mm; eyelid width 1.6 mm; eyelid length 3.4 mm; anterior rim of orbit to tip of snout 5.6 mm; horizontal orbit diameter 2.4 mm; interorbital distance between angle of eyes 3.1 mm; distance between nuchal groove and gular fold 3.9 mm; snout to forelimb 16.2 mm; distance separating external nares 2.1 mm; distance separating internal nares 2.0 mm; hedonic mental gland 1.9 mm; axilla to groin 25.7 mm; shoulder width 2.1 mm; tail length 43.2 mm; tail width at base 4.6 mm; forelimb length 7.0 mm; hind limb length 10.4 mm; appressed limbs are separated by 3 1/2 costal folds; hand width 3.2 mm; foot width 3.8 mm; 32 maxillary teeth; 20 vomerine teeth. Tail and limbs yellow color, dorsum dark brown with yellow spots, gray abdomen, and head darker than the rest of the body ( Fig. 3 ).

Coloration in life. We examined 35 adults of Bolitoglossa tamaense , which showed eight patterns of variation in color: A) males with tail and limbs yellow, dorsum dark brown with small yellow spots; B) females with the tail, dorsum and limbs orange, with a dark brown head; C) males and females dark brown with orange feet; D) females orange from the tail to the head, with limbs dark brown. E) Males with the tail, head and feet slightly reddish; F) males with dorsum partially orange with dark spots, with dark brown limbs; G) Males with the tail orange, with yellow feet, head slightly stained orange; H) Males and females with a cream-colored dorsum. All juveniles exhibit dark gray skin coloration. In all individuals, regardless of dorsal color patterns, ventral surface is gray with small brown spots ( Fig. 3 ).

Coloration in preservative. All yellow individuals and individuals with gray and brown retain the original color. The orange individual lost all color and turned gray.

Variation. The type series consists of 12 specimens (six males and six females), with a SVL range from 36.2- 52.7 mm. Size is sexually dimorphic: mean SVL of six males= 37.2 mm, six females= 47 mm. The largest individual was a gravid female (SVL= 52.7 mm), and the smallest male had a SVL= 36.2 mm. The number of maxillary teeth also varies between the sexes (male mean=35; females mean=40), as well as the vomerine teeth (males mean=18; females mean=21).

Etymology. The specific epithet refers to the Tama National Natural Park; B. tamaense is the first species of salamander recorded from this part of Colombia.

Geographic distribution and natural history. Bolitoglossa tamaense is known only from high Andean forests in the eastern flank of the Cordillera Oriental, municipality of Toledo, Departamento de Norte de Santander, Colombia ( Fig. 5 ). Only two localities have been recorded for this species, both in small forest patches, one at 2,000 m elevation and the other population at 2,700 m elevation. Field trips in the area, between 1,800 to 3,300 m elevation, revealed no additional localities. Individuals were active during nighttime, mostly between 18:00 and 19:00 hours, when they are found perched on ferns, bromeliads and low vegetation. During daytime they were found under leaf litter. The population of B. tamaense from Remansos (2,000 m) inhabits a small patch associated with a stream. Whereas several individuals were found in riparian vegetation at Remansos, the salamanders at Asiria inhabit a forest patch without flowing water. After dissecting three females from Remansos we found five pairs of oviductal eggs in each of them. In both localities B. tamaense shares the habitat with at least two species of frogs of the genus Pristimantis .

Figures

FIGURE 2. Bayesian tree of Bolitoglossa (Eladinea) based on analysis of the 16 S rRNA gene; the South american clade of Eladinea is indicated by heavier lines. The values on the internodes indicate support values for each node in the following way: Bayesian posterior probability / ML bootstrap support. Support values for nodes with prior probabilities lower than 0.95 and bootstrap support below 70 are not shown. B. mexicana and B. zapoteca were used as outgroups.

FIGURE 3. Adults of Bolitoglossa tamaense sp. nov. Coloration in life: A. Males with tail and limbs yellow, dorsum dark brown with small yellow spots. B. Females with the tail, dorsum and limbs orange, with a dark brown head. C. Males and females dark brown with orange feet. D. Females orange from the tail to the head, with limbs dark brown. E. Males with the tail, head and feet slightly stained red. F. Males with dorsum partially orange with dark spots, with dark brown limbs. G. Males with the tail orange, with yellow feet, head slightly stained orange. H. Males and females with a cream-colored dorsum.

FIGURE 4. Bolitoglossa tamaense, holotype. A. Dorsum. B. Head, lateral view. C. Hand. D. Leg. E. Head, dorsal view. F. Head, ventral view.

FIGURE 5. Localities of new species of Bolitoglossa. Tama National Natural Park, Departamento de Norte de Santander, Colombia.