Bolitoglossa muisca, López-Perilla & Fernández-Roldán & Meza-Joya & Medina-Rangel, 2023

Lopez-Perilla, Yeny Rocio, Fernandez-Roldan, Juan David, Meza-Joya, Fabio Leonardo & Medina-Rangel, Guido Fabian, 2023, A new Bolitoglossa (Amphibia, Caudata, Plethodontidae) from the Cordillera Oriental of Colombia, ZooKeys 1158, pp. 27-48 : 27

Bolitoglossa muisca sp. nov.

Figs 2 , 3 , 4 , 5 , 6 , 7

Type material.

Holotype. IAvH-Am-17413, an adult female from Finca La Esmeralda, vereda Roble Hueco, Bojacá municipality, Cundinamarca department, Colombia (4.6963, -74.3624, 2630 m a.s.l.), collected by Y.R. López-Perilla on 4 February 2021 (Fig. 1 ).

Paratypes (n = 8: 3 females, 5 males). IAvH-Am-17417 (adult male), IAvH-Am-17419 (adult female), IAvH-Am-17421 (adult male) and IAvH-Am-17422 (adult male) from Finca La Esmeralda, vereda Roble Hueco, Bojacá municipality, Cundinamarca department, Colombia (4.6963, -74.3624, 2630 m a.s.l.), collected by Y.R. López-Perilla in February 2021 (Fig. 5 ). IAvH-Am-17423 (adult male), IAvH-Am-17425 (adult male) and IAvH-Am-17428 (adult female) from Finca Peñas Blancas, vereda Roble Hueco, Bojacá municipality, Cundinamarca department, Colombia (4.6916, -74.3581, 2390 m a.s.l.), collected by Y.R. López-Perilla in February 2021. IAvH-Am-17429 (adult female) from vereda Cascajal, Soacha municipality, Cundinamarca department, Colombia (4.5954, -74.2922, 2700 m a.s.l.), collected by G.F. Medina-Rangel in September 2022.

Referred specimens

(n = 36: 7 females, 29 juveniles and subadults). ICN 3544-48, five adult females from Hacienda 'El Soche’, Granada municipality, Cundinamarca department, Colombia (4.5153, -74.3240, 2600 m a.s.l.) obtained by Rurithza Velandia in December 1977; specimen ICN 3545 is cleared and stained; MVZ 167997, a juvenile obtained by Pere Alberch on 17 December 1978. ICN 58245-58268, a batch of 23 juvenile and subadult specimens from Hacienda 'La Tribuna’, vereda Noruega Alta, Silvania municipality, Cundinamarca department, Colombia (4.4836, -74.3203, 2700 m a.s.l.) obtained by Cesar Monguí in September 2008. ICN 60319-20, two juveniles from vereda Roquemonte, San Antonio del Tequendama municipality, Cundinamarca department, Colombia (4.6056, -74.3008, 2600 m a.s.l.) obtained by J.D. Fernández in August 2013. ICN 60321-22, a juvenile and an adult female (respectively) also from vereda Roquemonte, San Antonio del Tequendama municipality, Cundinamarca department, Colombia (4.6056, -74.3008, 2600 m a.s.l.) obtained by J.D. Fernández in March 2017. ICN 60323-25, two juveniles and an adult female (respectively) from vereda Roquemonte, San Antonio del Tequendama municipality, Cundinamarca department, Colombia (4.6056, -74.3008, 2600 m a.s.l.), also obtained by J.D. Fernández in January 2018.

Diagnosis.

Bolitoglossa muisca is a member the subgenus Bolitoglossa Eladinea and of the Bolitoglossa adspersa species group. The new species is characterized by the following morphological characters: a large-size body; a broad head; a rounded snout in dorsal and ventral views; a very thick postocular fold; a moderate subgular fold; smooth skin texture; moderately long limbs; moderate webbing on third finger and toe; and a short, robust tail.

Even though Bolitoglossa adspersa and B. muisca share their moderate webbing on hands and feet, we regard the former as having less webbing than the latter. Moreover, the tips of the fingers and toes are separated from the distal margin of the webbing, exposing their subcircular-shaped digits; unlike those of B. muisca (Fig. 4 ). The new species is slightly larger on average than B. adspersa (mean SVL 52.8 ± 3.4 mm; range 33.0-72.1 mm; n = 22 vs. 45.0 ± 3.4 mm; range 30.7-66.3 mm; n = 26), additionally, the tail of B. adspersa is thin and long in relation to the trunk but thick and short in relation to the trunk in B. muisca (Table 1 ). Bolitoglossa muisca differs from B. capitana by having moderately webbed hands and feet (vs. almost fully webbed hands and feet in B. capitana ), by being overall smaller in size (mean SVL 52.8 ± 3.4 mm; range 33.0-72.1 mm; n = 22 vs. 75.2 ± 10.9 mm; range 59.2-85.5 mm; n = 5), by having fewer maxillary teeth (mean MT 24.4 ± 3.0; range 14-36; n = 17 vs. 30.4 ± 5.1; range 22-37; n = 5) by having fewer vomerine teeth (mean VT 35.1 ± 4.0; range 24-53; n = 17 vs. 66.4 ± 14.8; range 46-87; n = 5), and because the new species bears a faint, very small gular fold that is large, thick and notable in B. capitana ( Cruz-Rodríguez et al. 2021: figs 13, 14). Bolitoglossa muisca differs from B. pandi because the tips of the third digit and toe of the latter are triangular and pointed in outline (vs. third digits and toes oval and webbed to a higher degree in B. muisca ), by being a slightly larger species (mean SVL 52.8 ± 3.4 mm; range 33.0-72.1 mm; n = 22 vs. 44.0 ± 2.9 mm; range 35.9-52 mm; n = 12), and by having mostly smooth skin (vs. coarse skin in B. pandi ); moreover, the tail of B. pandi tapers gradually and symmetrically from broad to slender antero-posteriorly, unlike that of the new species, which is slightly rectangular in outline, becoming abruptly wider than the base, and ending in a rounded tip (Table 1 ).

Description of the holotype.

An adult female (SVL = 61.3 mm) with a broad head (HW/SVL = 0.16); head longer than wide (HW/HL = 0.90); neck with a small, faint gular fold; snout short and truncated in profile, and dorsal view, but less so in ventral view (SNL = 3.1 mm); large eyes that do not extend beyond the outline of the head in dorsal view and smaller than interorbital distance (EYD = 3.18 mm, IOD = 3.37 mm); with a thick post ocular fold that extends past the posterior commissure of the eye onto the anterior margin of the gular fold; canthus rostralis subtle, small, rounded in outline; 33 maxillary teeth, 16 to the right and 17 to the left; vomerine teeth 25, these are not arranged in a single row but grouped towards the margins of the parasphenoid bone; with three premaxillary teeth that pierce the upper lip in males; nasolabial grooves well developed; moderate interdigital webbing on hands but third finger extends slightly further than the other fingers; toes with less interdigital webbing than fingers, toes II-V with less membrane than toe I; with subterminal pads on digits, digits in order of increasing length I<II<IV>III; toes I<II<III<IV>V; longest digits of hand and feet are subcircular (L3T and L3F), limbs relatively long (FL/SVL = 0.23, HLL/SVL = 0.23); tail not exceeding standard length (TL/SVL 0.93), narrower than the body at the base (posterior to the vent), slightly rectangular in outline, becoming abruptly wider than the base and ending in a rounded tip, but this condition is artefactual because the tip of the tail is missing; a long trunk (52.5 mm); with 13 costal grooves (Fig. 3 ). See Table 2 for meristic data of all type specimens.

Coloration of the holotype in life.

The color pattern of the holotype is described based on a photograph taken the day after capture. The dorsal surfaces of the head, the body and the tail are Raw Umber (280), strongly speckled with Dark Salmon (59); white stipples on the lateral surface of the head; the flanks, dorsum, legs, and tail have an irregular thin white stripe; the iris is Light Sky Blue (191) with Pratt’s Rufous (72) reticulations. The throat and ventral surfaces are white with Raw Umber (280) speckles and reticulations; the ventral surfaces of the limbs and tails with some Light Orange Yellow (7) vermiculated; underside of hands and feet are Olive-Brown (278).

Coloration of the holotype in preservative.

The color pattern of the holotype was recorded after approximately five months stored in 70% ethanol. The dorsal surfaces of the head, the body and the tail are Raw Umber (280), strongly speckled with Dark Salmon (59); the flanks, dorsum, legs, and tail have an irregular Smoke Gray (266) stripe; the iris is Amber (51) with Orange Rufous (56) reticulations. The throat and ventral surface are Smoke Gray (266) with Raw Umber (280) speckles and reticulations; hands and feet soles are Grayish Horn (268) ventrally (Fig. 3 ).

Color variation.

The specimens IAvH-Am-17414-16 have the dorsal surfaces of the head, flanks, dorsum, front legs and vertebral band Orange-Rufous (56), strongly speckled with Raw Umber (280); paravertebral area, tail, and hind legs back Light-Yellow Ocher (13) with Raw Umber (280) dashes, and bordered with a wide Raw Umber (280) band; white stipples on the lateral surface of the head and back of the legs. The specimen IAvH-Am-17425 has the dorsal surfaces of the head, body, legs, and tail Dark Salmon (59), strongly speckled with Raw Umber (280), with greater concentration at the nape of the neck. Ventral markings or blotches on the ventral surfaces of the body and tail vary in shape and size; often with irregular margins but are consistently white or cream-colored independent of sex and age (Figs 5 - 7 ).

Etymology.

Named after the native human inhabitants of the Altiplano Cundiboyacense and Sabana de Bogotá. The Muiscas regarded amphibians as sacred creatures associated with sex, fertility, and the arrival of the rainy season. The specific epithet is used as a noun in apposition.

Distribution.

At present, Bolitoglossa muisca is known only from small cloud forest remnants on the western slopes of the Cordillera Oriental of Colombia in Bojacá, Granada, San Antonio del Tequendama, Silvania, and Soacha municipalities, Cundinamarca department. All specimens collected between 2390-2700 m a.s.l. (Fig. 2 ).

Natural history.

Individuals from Bojacá municipality were regularly found at night on the base (on the mantillo) and leaves of Cyatheaceae ferns, which are dominant in the cloud forests of the Tequendama region of Cundinamarca department (Fig. 8 ). Most individuals from San Antonio del Tequendama municipality were found active at night perching on small branches of shrubs ( Araceae and Melastomataceae ), usually far away from rivers or streams. During the day, a few individuals were found inactive inside bromeliads below two meters height. When handled, these salamanders produced a sticky whiteish mucoserous substance; we consider this to be a defense mechanism against potential predators ( Arrivillaga and Brown 2018). Two frog species ( Pristimantis sp. and P. uisae ) and a lizard ( Anolis heterodermus ) were found in sympatry with Bolitoglossa muisca ; no other salamander species were found within our fieldwork area.

Other material examined.

Countries are indicated in bold capitals, departments in regular capitals, municipalities, and localities in plain text. * Denotes specimens examined via photographs.

Bolitoglossa adspersa (n = 29). Colombia: Boyacá: Duitama, Páramo de la Rusia: ICN 4301, 4310; Toquilla, Paramo de Toquilla : ICN 9487 . Cundinamarca: Bogotá, D.C., Páramo de Cruz Verde: IAvH-Am-8917, San Cristobal, Tánques de Vitelma, finca La Marranera: IAvH-Am-696, 2947, 2954, 2957-58, 2964, 2967, 2972, Usme, represa El Hato: ICN 37555 ; Cabrera: vereda de Hoyerías: IAvH-Am-13253 Fómeque, Laguna de Chingaza : ICN 4884, 4891; Guasca, Paramo de Guasca : ICN 4521, 4525, 4546; Fómeque, laguna Chingaza: ICN 4885; Fusagasugá, km 50 carretera Fusagasugá-La Florida: ICN 39096 ; Guatavita, vereda Montenquiva: IAvH-Am-13174-75, ICN 55420 ; Guayabetal: Páramo Atravezado: IAvH-Am-14808; Paramo de Palacio , P.N.N. Chingaza: IAvH-Am-3101, 3096, Quétame, km 22 carretera central Villavicencio-Alto del Tigre : ICN 7123; Ubaque, reserva ecológica Matarredonda: IAvH-Am-13254 .

Bolitoglossa capitana (n = 4). Colombia: Cundinamarca: Albán, Granjas del Padre Luna: ICN 9221 & MLS 182-184*.

Bolitoglossa guaneae (n = 27). Colombia: Santander: Charalá, Virolín, Cañaverales, sector El Reloj: ICN 5197, 8555, 8557, 12770-72, 34230, Cuchilla del Fara: ICN 47980 , Hacienda La Sierra : ICN 34229 -30, km 56: ICN 19558 , UIS-A 1369, UIS-A 2078, UIS-A 2082, UIS-A 2179, UIS-A 2317, UIS-A 2325-6, UIS-A 2891, UIS-A 2893, UIS-A 2895, UIS-A 2898-9, paratypes UIS-A 2203, UIS-A 2320, UIS-A 2324, UIS-A 2897 .

Bolitoglossa muisca (n = 54). Colombia: Cundinamarca: Bojacá, vereda Roble Hueco, predio La Esmeralda: IAvH-Am-17413-22, Predio Peñas Blancas: IAvH-Am-17423-28; Granada, hacienda El Soche: ICN 3544-48, MVZ 167997*; San Antonio del Tequendama, vereda Roquemonte, cerca de entrada Parque Chicaque: ICN 60319 -25; Silvania, vereda Noruega Alta, Hacienda La Tribuna : ICN 58245 -58268; Soacha, vereda Cascajal: IAvH-Am-17429 .

Bolitoglossa nicefori (n = 13). Colombia: Santander: Floridablanca, El Mortiño, quebrada Torrentosa : ICN 50000 , 58227, 58231; Zapatoca, finca Los Puentes, quebrada Uchuvala: ICN 58223 -24, 58226; Piedecuesta, vereda Los Monos: UIS-A 2987, UIS-A 2991; Los Santos, vereda El Carrizal, finca Utopía: UIS-A 5273-4; Guapotá, vereda Las Flores, finca La Chocolatera: UIS-A 5264; San Gil, vereda San José, finca La Esperanza: UIS-A 5270-1 .

Bolitoglossa pandi (n = 20). Colombia: Cundinamarca: Pandi, vereda Buenos Aires Alta : ICN 45500 , Supatá, vereda Las Lajas, reserva Cuzcungos: ICN 58492 -05, Villeta, La Esmeralda: IAvH-Am-10303-08 .

Figures

Figure 1. Map of Colombia showing the distribution records of Bolitoglossa muisca and other sympatric Bolitoglossa species in the Cordillera Oriental.

Figure 2. Maximum-likelihood tree of Bolitoglossa (Eladinea) adspersa group showing the phylogenetic position of Bolitoglossa muisca. Vertical bars indicate the two-species hypothesis within Bolitoglossa adspersa with the supported species partitions inferred using genetic data (p-distances, ABGD, and P ID). Support values for well-supported nodes correspond to ML ultrafast bootstrap (> 95) and Bayesian posterior probabilities (> 0.95), respectively. Photograph by JDF.

Figure 3. Holotype of Bolitoglossa muisca (IAvH-Am- 17413) in A dorsal B ventral and C lateral views. Photographs by JDF. Scale bar: 10 mm.

Figure 4. Ventral views of the hands and feet of A Bolitoglossa muisca (IAvH-Am- 17413) B Bolitoglossa adspersa (ICN 4885). Photographs by JDF. Scale bar: 10 mm.

Figure 5. Paratype of Bolitoglossa muisca (IAvH-Am- 17419) in life. Photographs by JDF.

Figure 6. Chromatic variation of Bolitoglossa muisca in life. Notice some individuals bear white blotches on the ventral surfaces of the body and the tail. Photographs by YLP.

Figure 7. A non-captured individual of Bolitoglossa muisca from the surroundings of Reserva Chicaque, San Antonio del Tequendama, Cundinamarca, Colombia, photographed by JDF in October 2015. Notice the striking contrasting dark brown background coloration with ochre markings primarily on the dorsal surfaces of the body.

Figure 8. The Andean cloud forests of Bojaca, Cundinamarca, and the habitat of Bolitoglossa muisca. Photographs by YLP.