Bolitoglossa guaneae, Acosta-Galvis & Gutiérrez-Lamus, 2012

Acosta-Galvis, Andrés R. & Gutiérrez-Lamus, Doris L., 2012, A New Species Of Salamander (Bolitoglossa: Plethodontidae) From The Cordillera Oriental Of The Colombian Andes, Papéis Avulsos de Zoologia (São Paulo) 52 (18), pp. 201-218 : 201-218

Bolitoglossa guaneae sp. nov.

( Fig. 1 )

Holotype: Adult female, ICN-MHN 54440 (field number ARA 4984), collected by Benjamin Tapley and Andrés Acosta, on 8 December 2008, deposited in the Amphibian collection, Institute of Natural Sciences, Universidad Nacional de Colombia.

Type Locality: Colombia, Department de Santander, Municipality of Encino, Cartagena stream, enlargement area of Flora and Fauna Sanctuary Guanentá, Alto Río Fonce , vereda La Chapa, Department of Santand- er, west flank from Cordillera Oriental , Colombia, elevation 1836 m. 06°06’58.2”N and 73°07’34.4”W .

Paratypes: (9) ICN-MHN 34230, UIS-A 2203, UIS-A 2320, UIS-A 2324, UIS-A 2897 adult males from Municipality of Charalá, km 38 Hacienda La Sierra Santuario de Fauna y Flora Guanentá , Alto Río Fonce , vereda Santa Helena, Departamento of Santander Colombia, elevation 2400 m. Collected by Claudia Vélez, Jeannette Nieto, Doris Gutiérrez , Ruth Estupiñán , and Sandy Arroyo ; MUJ 7193- 4 adult males, Boyacá , Municipality of Moniquirá, site El Arizal elevation 2050 m. ICN-MHN 12772 adult female Guillermo river, Bogotacito 55-56 km. along the road from Duitama to Charalá ; ICN-MHN 8557 adult female, from Municipality of Charalá , Virolin , Departamento of Santander .

Etymology: The specific epithet of this new salaman- der is allusive to the native Guanes who lived within the distribution range of the species. They inhabited the area since 600 B.C and in 1586 they almost disappeared after fighting with the Spanish. Nowadays they live in Guane, a town of Santander in a small Indian area.

Referred Specimens: (43) ICN-MHN 12770-1, ICN-MHN 19558, ICN-MHN 4418 (Cleared and stained skeleton) Bogotacito 55-56 km. along on the road from Duitama to Charalá by the Guillermo river , Department of Santander, Colombia elevation 2340 m. ICN-MHN 8555, ICN-MHN 8556 (Cleared and stained skeleton), ICN-MNH 5197 Virolín, Vereda El Reloj, Cañaverales, Municipality of Charalá, Departamento of Santander Colombia, elevation 1750-2200 m. Adult males: UIS-A 1307, UIS-A 2026, UIS-A 2079, UIS-A 2325, UIS-A 2891, UIS-A 2898. Adult females: UIS-A 1369, UIS-A 2078, UIS-A 2082, UIS-A 2179, UIS-A 2316-7, UIS-A 2326, UIS-A 2893, UIS-A 2895, UIS-A 2899. Young males: UIS-A 2070, UIS-A 2184, UIS-A 2321. Juvenile females: UIS-A 1366, UIS-A 1890, UIS-A 1891, UIS-A 2048, UIS-A 2081, UIS-A 2318, UIS-A

2894, UIS-A 2896, UIS-A 2900, ICN-MNH 26325 70 km. along the road from Duitama to Charalá, Departmento of Santander Colombia, elevation 1760 m . All UIS-A were collected at La Sierra farm, Santuario de Fauna y Flora Guanentá, Alto Río Fonce, vereda Santa Helena, Departmento of Santander, Colombia elevation 2400 m .

Diagnosis: A endemic Bolitoglossa recognizable from the other species of the genus by a combination of the following characters: small size of the adult males (SVL = 31.53-41.56 mm) and adult females (SVL = 39.76-46.08 mm); extensively interdigital webbing with free digital tips on the fingers and toes (Type D in South American species, sensu Brame & Wake, 1963, and category D, sensu Wake & Brame, 1969) ( Fig. 2 ); ventral surface of digit tips without terminal flattened tubercles ( Fig. 2 ); snout short and rounded in lateral profile, head length 2.0- 2.92 mm in adult males, 1.88-2.88 mm in adult females; snout rounded in dorsal view; protruding eyes on dorsal view; well-defined postcephalic constriction ( Fig. 3 ). Ventral surfaces (in alcohol) brown with numerous tiny cream guanophores with irregular spots; nasolabial grooves cream on preserved specimens; males have white testis; adult males 1 or 2 premaxillary teeth pierce the lip; copper iris; Snout in profile lack numerous tiny cream guanophores; terminal phalanges of digits not expanded; pelvic girdle without posterolateral projections.

Bolitoglossa guaneae differs from other Colombian salamanders in morphologic and chromatic characteristics. From species with basal webbing with free digital tips on the fingers and toes (Type A in South American species, sensu Brame & Wake, 1963, and type slightly and moderately webbed sensu Wake & Brame, 1969), as B. adspersa , B. hiemalis , B. hypacra , B. ramosi , B. savagei , B. tatamae , B. walkeri and B. vallecula ( Fig. 4 ), by having extensively interdigital webbing with free digital tips on the fingers and toes (Type D in South American species, sensu Brame & Wake, 1963, and category D, sensu Wake & Brame, 1969) ( Fig. 2 ). From species with extensively or completely webbing (sensu Wake & Brame, 1969 and Types C, D. E and F sensu Brame & Wake, 1963), Bolitoglossa guaneae can be differentiated from B. altamazonica in having extensively interdigital webbing with a longer and triangular third finger (completely webbing and tips rounded in B. altamazonica ) ( Fig. 5 ); differs from B. biseriata and B. silverstonei in having a ventral surface brown with white dots (ventral surface cream intensified by numerous small irregular light brown spots in B. biseriata and B. silverstonei ), white testes in males (usually black testes in B. biseriata and B. silverstonei ) and extensively interdigital webbing with free digital tips on the fingers and toes (being completely webbing without free digital tips in B. biseriata and B. silverstonei ); from B. medemi in the absence of digital depressions (present in B. medemi ); B. guaneae can be differentiated from B. lozanoi and B. nicefori in having extensively interdigital webbing with free digital tips on the fingers and toes (being completely webbing without free digital tips in B. lozanoi and B. nicefori ) ( Fig. 5 ) and more protruding eyes (not protruding in B. lozanoi and B. nicefori ) ( Fig. 6 ); from B. phalarosoma in being dorsal and lateral surfaces is dark brown (usually dorsal surfaces of head and body reddish brown with a cream patch which covers the surface of the face in B. phalarosoma ); B. guaneae can be differentiated from B. sima by less extensive webbing (being completely webbing sensu Wake & Brame, 1969 and types F sensu Brame & Wake, 1963); from B. pandi in adult size (SVL = 39.76-46.08 mm in adult females in B. guaneae , versus SVL = 44.71-50.4 mm in adult females in B. pandi ) and less protruding eyes on dorsal view (more protruding eyes on dorsal view in B. pandi ) ( Fig. 6 ); from B. capitana , in adult size (SVL = 31.53-41.56 mm in adult males and SVL = 39.76-46.08 mm in adult females in B. guaneae , versus SVL = 62.78-62.83 in adult males and SVL = 74.19-77.39 mm in adult females of B. capitana ).

Description and Definition: This is tiny species, recognized as one of the smallest species among the Colombian species of Bolitoglossa . Males are 31.53-41.56 (mean = 37.6, SD = 4.53, n = 12) mm and females are 39.76-46.08 (mean = 37.7 mm, SD = 4.5, n = 11) mm of SVL. The snout is short in profile, rounded, sloping ventrolaterally and projects ahead of the upper jaw; canthus rostralis not prominent; postcephalic constriction clearly distinguished; fingertips and toe tips without a tubercle; nearly oval mental gland in males, transversally located and posteriorly prominent, 30-45.9% (mean = 37.7) the of head width in males. In profile, nasolabial grooves short, prominent ( Fig. 3 ); curved towards the labial region being more prominent in males. Diameter of the eye versus head length, 30.80-36.80% (mean = 33.63, n = 12) in males, and 18.04-29.85% (mean = 23.98, n = 11) in females. The upper lip is neither prominent nor wide. The ratio of head width versus head length ranges from 73.34-92.18% (mean = 85.02, n = 12) in adult males, and 72.16-88.18% (mean = 82.77, n = 11) in adult females; postiliac glands absent; gular fold prominent, projected towards ventrolateral cephalic region; the upper lip of males bears two teeth between the nasolabial grooves (= premaxilary teeth piercing lip sensu Ehmcke et al. 2004); tongue enlarged and rounded in its anterior tip, with a notch. Relative length of fingers III > II ≥ IV > I; extensively interdigital webbing with free digital tips on the fingers (Type D in South American species, sensu Brame & Wake, 1963, and category D, sensu Wake & Brame, 1969) ( Fig. 2 ); tips of digits are triangular and short ( Fig. 2 ); hand width versus head length, 26.55-48.71% (mean = 40.23, n = 12) in males, and 28.79-42.09% (mean = 37.08, n = 11) in females.

Hindlimbs short and robust; relative length of digits are III > IV > II > V > I, with extensively interdigital webbing with free digital tips on the toes (Type D in South American species, sensu Brame & Wake, 1963, and category D, sensu Wake & Brame, 1969); foot length versus head length between 38.63-53.84% (mean = 46.91, n = 11) in adult males, and between 35.2-48.0% (mean = 43.19, n = 11) in adult females. Fingers and toes without terminal flattened tubercles.

A very distinct constriction at the base of the tail with a slight lateral compression; tail length versus SVL 67.08-89.96% (mean = 78.50, n = 9) in males, and 60.71-79.53% (mean = 72.04, n = 9) in females. Skin of dorsum with lateral and ventral smooth surfaces.

Color in Life: The dorsal and lateral ground color is dark brown, sometimes mottled or streaked with white, diffuse paled blue and cream guanophores, in some individuals reddish marks are also present. Hindlimbs are dark brown with reddish; Tail is light to dark brown, in occasions mottled with white; ventral surfaces are Brown with dense white cream irregular spots and guanophores blue very faint, sometimes mottled white, tail is ventrally mottled white in some individuals; copper iris without black reticulations ( Fig. 1 ).

Coloration in Alcohol: The ground color of head, trunk, and tail of the type series is dark brown, with palmar and plantar surfaces light brown. Venter dark brown with a few cream patches and cream guanophores.

Variation: The type series presents in general terms, a uniform color in all adult specimens; the dorsal color pattern with inconspicuous paler brown dorsolateral stripe, extending from posterior edge of eyelid dropping towards the base of thighs.

Osteology: The description is based on two specimens (ICN-MNH 8556, ICN-MNH 4418). Skull is oval and well formed, anterior elements are generally well ossified and articulated. Pars dentalis of the premaxilla are short with four to five teeth. The internasal fontanelle is relatively narrow. Nasal bones are protrusive anteriorly and articulate with maxillaries, frontals, and premaxillaries. Prefrontal bones are nearly rectangular, with irregular edges, slightly longer along the posterior rather than the anterior ridge. Frontal bones occupy nearly half the length of the skull, irregular in shape; they overlap each other and the parietal bones. Irregular parietal bones; lateral parietal spur short, spreading to the level of both frontal and orbitosphenoid bones, separating their posterior edges. Parietals overlap each other mesially, and are overlapped by frontal bones. Occipital condyles rectangular, posterolaterally directed.

The right maxilla bears 13-21 teeth and the left one 16-24. Vomers triangular and rounded, bearing 19-28 teeth transversely positioned on the proximal end of the vomer, and wide internal nares. Antorbitalis cartilage is articulated to the posterior edge of the vomer, laterally to the maxilla and posteriorly to the orbitosphenoid bone. A large triangular parasphenoid bone making part of the cranial vault, extended from the vomer interspace to the ear capsules, bearing two plates covered with teeth arranged in short and oblique rows: the right plate bears 13-14 rows with 70-122 teeth, and the left one supports 12-13 rows with 73-138 teeth. Orbitosphenoid rectangular, with articular processes at anterior and posterior rims, and articulating with antorbitalis and pterygoid cartilages anterior and posterior, respectively. A large optic fenestrum; squamosal rectangular, irregular, and thin, investing the palatoquadrate laterally and articulating with the skull roof. The palatoquadrate is small, distally articulating with the articular by cartilaginous process, while the proximal end articulates to the posterior branch of the pterygoid. The right dentary bears 26-42 teeth and the left one 22-46. Fourteen presacral vertebrae, two caudosacral and 24 caudal. The neural arch of the atlas bears a cartilaginous spine. Transverse process forked, perpendicular to the body axis on the posterior vertebrae. Ribs curved; at the distal end of the second vertebra is a V-shaped cartilaginous plate. There are no carpal or tarsal fusions nor reductions. Metacarpi are robust. The phalangeal formula in hands is 1-2-3-2; in feet is 1-2-3-2-2; wide foot and robust metatarsi.

Hyobranchial apparatus cartilaginous; basibranchial cylindrical, with a rectangular posterior end articulating with the visceral arches while its anterior end is laterally extended; ceratobranchial is anteroposteriorly directed from the medial region of the basibranchial, having a wider and more irregular base; the origin of ceratobranchial II is at the posterior end of the basibranchial. The ceratohyal articulates with the distal end of the hypohyal, having narrow plates at the base, and cylindrical ones on the posterior half.

Measurements of Holotype: All measures in mm. SVL: 38.01; HW: 5.86; HL: 6.02; OD: 2.03; IOD: 1.61; END: 1.21; RL: 1.66; RW: 1.61; EW: 1.26; IN: 1.68; SGD: 8.30; TL: 3.55; FEL: 3.56; FL: 4.85 and HDW: 3.02.

Distribution: Bolitoglossa guaneae is a species restricted to the central region of the west flank of the Colombian Cordillera Oriental, in Andean forests; its distribution is based on political geography and is known from the Departamentos of Santander and Boyacá from elevations between 1700-2400 m. Annual rainfall ranges between 1400-2000 mm ( Fig. 7 ).

Habitat and Behavior: Bolitoglossa guaneae is a rare nocturnal species, restricted to near streams in primary or secondary forests with little anthropic intervention. (“low montane humid forest” (bh-mb) sensu Holdridge 1996, see also Gutiérrez-Lamus et al. 2004). This species was found standing on low leaves (0,50-1, 30 m) of Araceae , Melastomataceae , and ferns. The Holotype was collected on a leaf 0.5 m above ground at 22:06 h, with temperature of 21.1°C and 91% relative humidity.

Figures

FIGURE 1: Lateral (A) and dorsal (B) view of Holotype (adult female; ICN-MHN 54440) of Bolitoglossa guaneae. Photos by Andrés Acosta.

FIGURE 2: Ventral surfaces showing extensively interdigital webbing with free digital tips on the fingers and toes (Type D in South American species, sensu Brame & Wake, 1963, and category D, sensu Wake & Brame, 1969), in species of the genus Bolitoglossa in the Cordillera Oriental of Colombia. Bolitoglossa pandi: a) ICN 45500; Bolitoglossa guaneae: b) ICN 8557, c) ICN 12772, d) ICN 19558; Bolitoglossa capitana: e) ICN 9221.

FIGURE 3: Paratype (ICN-MNH 8557) of Bolitoglossa guaneae (A) Head in right lateral view, (B) Head in dorsal view. Photos by Andrés Acosta.

FIGURE 4: Ventral surfaces showing basal webbing with free digital tips on the fingers and toes (Type A in South American species, sensu Brame & Wake, 1963, and type slightly and moderately webbed sensu Wake & Brame, 1969); in species of the genus Bolitoglossa in the Cordillera Oriental of Colombia. Bolitoglossa adspersa: Parque Nacional Natural Chingaza, a) MUJ 267, Fomeque (Cundinamarca), b) MUJ 1907, c) MUJ 720; Bolitoglossa palmata from border Huila-Caquetá Departaments d) ICN 20792, e) ICN 20793, f) ICN 20794.

FIGURE 5: Ventral surfaces extensively or completely webbing (sensu Wake & Brame, 1969 and Types C, D. E and F sensu Brame & Wake, 1963), in species of the genus Bolitoglossa in the Cordillera Oriental of Colombia. Bolitoglossa altamazonica: Villavicencio (Meta), a) MUJ 4096, b) MUJ 4101, c) MUJ 4097, Leticia (Amazonas), d) ICN 46852, Leticia (Amazonas). Bolitoglossa lozanoi: e) MO 589, f) MOM 587, g) MOM 588, h) MOM 590; Bolitoglossa nicefori: Mesa de los Santos (Santander), i) MUJ 2445, j) MUJ 2377, k) ICN 50000, l) ICN 50001.

FIGURE 6: Dorsal view of heads from species of the genus Bolitoglossa from the Eastern Cordillera of Colombia: Bolitoglossa adspersa: a) ICN 37863, Municipality San Antonio de Tena (Cundinamarca), b) ICN 12774, Páramo of Rusia, Municipality of Duitama Boyacá (Cundinamarca), c) ICN 37563, Alto del tigre (Meta); B. altamazonica: Villavicencio (Meta), d) MUJ 4095, e) MUJ 4097, f) MUJ 4096, Leticia (Amazonas), g) ICN 36510, h) ICN 46852; B. capitana: i) ICN 9221; B. lozanoi: j) MO 589, k) MOM 587, l) MOM 588; B. nicefori: Mesa de los Santos (Santander), m) MUJ 2377, Tona, n) ICN 50001; B. palmata: Political limits from Caquetá-Huila, o) ICN 20792, p) ICN 20793, q) ICN 20794; B. pandi: Municipality of Pandi, r) ICN 45500, Municipality of San Francisco (Cundinamarca), s) MUJ 7921; B. guaneae: Santander, t) UIS-A 2203, u) UIS-A 2184, v) UIS-A 2324.

FIGURE 7: Localities of Bolitoglossa guaneae from western slopes of Cordillera Oriental from Colombia, red dot corresponds at type locality.