Pristimantis leopardus, Rivera-Correa, Mauricio, Jiménez-Rivillas, Carlos & Daza, Juan M., 2017

Rivera-Correa, Mauricio, Jiménez-Rivillas, Carlos & Daza, Juan M., 2017, Phylogenetic analysis of the Neotropical Pristimantis leptolophus species group (Anura: Craugastoridae): molecular approach and description of a new polymorphic species, Zootaxa 4242 (2), pp. 313-343 : 316-324

Pristimantis leopardus sp. nov.

( Figs. 2–5 )

Holotype. MHUA-A 8865, adult male ( Figs. 2 , 4 A) from Colombia, Departamento de Antioquia, Municipio de Sonsón, Vereda Manzanares, Páramo de Sonsón, Cerro La Vieja (5.77279, -75.23354; 3143 m.a.s.l.), collected on July 1, 2014 by Estefany Cano, Natalí Duque, José Fang and Laura Pinto.

Paratypes. MHUA-A 7577–78, 7585, 7588, 7592, 7604, 7613, 7615–16, 7626, adult males; MHUA-A 7614, (adult female) from Colombia, Departamento de Antioquia, Municipio de Sonsón, Vereda San Francisco, Páramo de Sonsón , Alto de las Palomas (5.72584, -75.25165; 3221 m.a.s.l.), collected on July 2, 2012 by Carlos Jiménez- Rivillas. MHUA-A 8831 adult male. MHUA-A 8833 sub-adult female. MHUA-A 8836–40, 8861, 8863, 8872, adult males from Colombia, Departamento de Antioquia, Municipio de Sonsón, Vereda La Palmita, Páramo de Sonsón, Alto de las Palomas (5.72585, -75.25176; 3207 m.a.s.l.), collected on April 28th, 2014 by Juan D. Loaiza, M. Isabel Pérez, Santiago Cuartas and Carlos Marín. MHUA-A 8835, 8845-46, 8859, 8866, 8869-70, 8875, adult males . Colombia, Departamento de Antioquia, Municipio de Sonsón, Vereda La Palmita, Páramo de Sonsón , Alto de las Palomas (5.72591, -75.25146; 3199 m.a.s.l.), collected on April 29, 2014 by Juan D. Loaiza, M. Isabel Perez, Santiago Cuartas and Carlos Marín. MHUA-A 8847–48, 8851, 8854, 8856, 8858, 8860, adult males ( Figs. 4–5 ) from Colombia, Departamento de Antioquia, Municipio de Sonsón, Vereda Manzanares, Páramo de Sonsón, Cerro la Vieja (5.77344, -75.23289; 3220 m.a.s.l.), collected on June 29, 2014 by Laura Pinto, José Fang, Estefany Cano and Natalí Duque. MHUA-A 8841, 8843, 8853, 8864, 8867, 8871 adult males from Colombia, Departamento de Antioquia, Municipio de Sonsón, Vereda La Palmita, Páramo de Sonsón, Cerro las Palomas (5.72598, - 75.24912; 3361 m a.s.l.), collected on April 30, 2014 by Juan D. Loaiza, M. Isabel Perez, Santiago Cuartas and Carlos Marín. MHUA-A 8844 adult male ( Fig. 5 B), 9124–25, 9126 ( Fig. 5 C), 9127 ( Fig. 4 E) from Colombia, Departamento de Antioquia, Municipio de Sonsón, Páramo de Sonsón, Cerro La Vieja (5.77293, -75.2335; 3154 m.a.s.l.), collected on July 1, 2014 by Laura Pinto, José Fang, Estefany Cano and Natalí Duque. ICN 55753, adult female . ICN 55755–56 adult males from Colombia, Departamento de Antioquia, Municipio de Sonsón, Cerro las Palomas (5°43' 33.1" N; 75° 14' 56.3" W, 3360 m a.s.l.), collected on August 19, 2014 by Marco Rada, Gustavo Gonzalez and Giovanni Chaves . ICN 55757, adult male from Colombia, Departamento de Antioquia, Municipio de Sonsón, Cerro las Palomas (5°43' 37.0" N; 75° 15' 24.9" W, 2900 m a.s.l.) on August 19, 2014 by Marco Rada, Gustavo Gonzalez and Giovanni Chaves.

Generic allocation. We assigned the new species to the genus Pristimantis and P. leptolophus group on the basis of the phylogenetic hypothesis presented in this study ( Fig. 1 , Appendix 2).

Diagnosis. The new species is characterized by: (1) skin texture of the dorsum finely shagreen, venter areolate; dorsolateral folds and discoidal fold absents; (2) tympanic membrane and tympanic annulus evident, supratympanic fold weakly differentiated; horizontal diameter of tympanum 40–50% of eye diameter; (3) snout short, broadly rounded or truncated by protruding nostrils in dorsal view, truncate in profile (slightly sloping in some males); (4) upper eyelid with small tubercles, cranial crests absent; (5) dentigerous processes of vomers oblique, positioned posterior to level of choanae, broadly separated from each other; (6) in males, vocal slits and small subgular vocal sac present, nuptial pads absent; (7) Finger I shorter than II, with large rounded digital disc; (8) fingers without lateral fringes; (9) antebrachial, ulnar tubercles and fold absent, (10) tubercles on heel and outer edge of tarsus absent; inner tarsal fold absent; (11) inner metatarsal tubercle oval, two to three times as long as round outer metatarsal tubercle; supernumerary plantar tubercles very small and low; (12) toes without lateral fringes; webbing absent; fifth toe longer than third; (13) in life, dorsum golden to cooper with dark brown spots, marks and blotches (in some individuals forming a reticulated pattern); venter cream, golden, light brown, or yellow-green with or without blotches and marks, in some individuals forming a reticulated pattern; (14) adults small, males 16.0– 19.9 mm (media= 17.8 ± 1.0; n=30) and a single female 22.8 mm.

Comparisons. Pristimantis leopardus sp. nov. can be distinguished from other species in the P. leptolophus group ( Fig. 3 ) by having a golden to cooper dorsum with dark brown spots, and marks and blotches in some individuals that form a reticulated pattern; the skin texture of the dorsum is finely shagreen without tubercles or warts and it lacks dorsolateral folds; it lacks nuptial pads lateral fringes on toes and fingers. The other species in this group have dorsolateral folds and tuberculate and warty bodies, including tubercles on eyelids and heels, except for P. peraticus . Pristimantis leopardus sp. nov. and P. peraticus are not sister taxa in phylogenetic analyses of mtDNA (strong evidence for not being conspecific) and, furthermore, the later has a brown dorsum with no patterns or only ill-defined marks, the anterior and posterior surfaces of thighs are cream with brown spots, and finger and toes have fringes. For a summary of diagnostic characters for species in the P. leptolophus group see Table 3.

Description of the holotype. A small frog, 17.5 mm SVL ( Fig. 2 ); head barely wider than long; snout broadly rounded in dorsal view and truncate in lateral view, relatively short (snout–eye distance 16% SVL); canthus rostralis indistinct; loreal region concave; nostrils protuberant, directed anterolaterally, area between nostrils slightly concave; interorbital area flat, barely narrower than upper eyelid; eyes directed anterolaterally; eye-tonostril distance equal to eye diameter; cranial crests absent; upper eyelid with small and low tubercles; tympanic membrane and tympanic annulus distinct, round; supratympanic fold not differentiated ( Fig. 4 A); tympanum diameter 40% of eye diameter; postrictal tubercles absent. Lips not flared; choanae small, nearly rounded, not concealed by palatal shelf of maxillary; dentigerous processes of vomers oblique, positioned posterior to level of choanae, broadly separated from each other; tongue longer that wide, posterior one-half free from floor of mouth; vocal slits present, longitudinal, originating on the sides of the tongue and extending to the corner of the mouth, small subgular vocal sac. Texture of skin on dorsum and flanks finely shagreen, dorsolateral folds absent; venter areolate; thoracic fold and discoidal fold absent; cloacal sheath absent.

Forearm slender; radio-ulna length 30.0% SVL; ulnar tubercles and low ulnar fold absent; radio-ulna length longer than hand length (hand length 27% SVL); fingers without lateral fringes; relative lengths of fingers I, II, IV, III; palmar tubercle bifid, thenar tubercle oval; subarticular tubercles round, subconical; supernumerary palmar tubercles present at the base of all fingers, conspicuous; finger disc ovoid, disc cover of Finger I slightly expanded, those of Fingers II–IV extensively expanded; discs on outer fingers as wide as those of toes; all disc covers with elliptical ventral pads defined by circummarginal grooves. Nuptial pads absent. Hind limbs relatively slender; tibia length 57.7% SVL; foot length 82% of tibia length; tarsal fold and tarsal tubercles absent; heel (tibiotarsal articulation) without tubercles; toes without lateral fringes; subarticular tubercles round, subconical; inner metatarsal tubercle oval, about 2x as long as wide; subconical outer tubercle; supernumerary plantar tubercles round, low; disc covers slightly expanded; toes with defined pads; disc pads nearly rounded; relative lengths of toes I, II, III, V, IV ( Fig. 4 ); tip of Toe V reaching midway between penultimate and distal subarticular tubercle of Toe IV; tip of Toe III reaching proximal border of medial subarticular tubercle of Toe IV.

Coloration of holotype in life. Dorsum cooper with reticulated dark brown pattern mainly along the dorsal vertebral region to the sacrum ( Fig. 4 A); canthus rostral and lips iridescent with dark brown background; intraorbital and supra-tympanic thin, dark brown stripe. Arms, forearms, flanks, thighs and legs with multiple dark brown blotches and spots on a yellow-golden background; groin and hidden regions of the arms and legs goldcopper; throat region dark brown with tiny copper dots; belly copper to cream with small brown spots distributed at random; iris dark brown bronze with spots, irregular marks and a copper reticulate pattern.

Coloration of holotype in ethanol. The reticulated pattern of small stains, marks and spots of black color around arms on a dark brown background, flanks and legs are a mixture of brown and dark gray. Throat, venter, and underside of limbs dirty gray; palms and soles gray with some brown pigmentation on external fingers and toes ( Fig. 2 ).

Measurements of the holotype (in millimeters). SVL: 17.5; HL: 6.6; HW: 7.1; ED: 2.5; END: 2.8; NSD: 0.7; IND: 1.9; AMD: 3.3; TD: 1.0; FAL: 5.2; FAB: 1.6; HAL: 4.7; THL: 10.1; TL: 10.1; TAL: 6.2; FL: 8.2; TFD: 0.8 and FTD: 0.6.

Variation. Males are smaller than the single known female (see Table 4 ). Pristimantis leopardus sp. nov. is a highly polymorphic species: the dorsal and ventral patterns vary considerably, having or lacking conspicuous dark brown blotches, spots and marks, and not always forming reticulated patterns ( Fig. 4 ). These marks, on a goldcopper dorsum, are sometimes iridescent-green in life. The belly of most individuals is immaculate gray or brown, though some specimens have dark brown to almost black blotches, marks and spots ( Fig. 5 ).

Distribution and natural history. Pristimantis leopardus sp. nov. is known from two localities from the Páramo de Sonsón complex: Cerro Las Palomas and Cerro La Vieja, in the northern of the Cordillera Central of Departamento de Antioquia, Colombia ( Figs. 6–7 ). The new species inhabits cloud forests (2820–3035 m a.s.l.) and páramos (3165–3360 m a.s.l., Fig. 7 B). It has been observed mostly inside phytotelmata such as Bromelia sp.

and Paepalanthus sp. ( Fig. 8 ) at 0.5–1.0 m above ground. It is a nocturnal species with activity from 19:00 h to midnight; vocal activity was not heard during the sampling. When observed, air temperature ranged between 10.8°C to 13.3°C and relative humidity was 89.7% to 97.4%. P. scoloblepharus and P. parectatus , two species of the P. leptolophus group, are syntopic with P. leopardus sp. nov. but apparently do not use bromeliads as the new species frequently does. At present, the area does not have a national protection figure but this work is the result of a research project that is providing information for the ecological and biogeographic delimitation of the Paramo that might eventually be key for the preservation of this area.

Etymology. The specific epithet corresponds to the Latin noun leopardus and it refers to the spotted and striped pattern observed in the new species, which resembles color patterns the American felid genus Leopardus , known in Spanish as tigrillos or ocelotes.

Figures

FIGURE 1. Maximum clade credibility tree after a partitioned Bayesian analysis using 8945 sites depicting the phylogenetic relationships among Pristimantis including the Pristimantis leptolophus species group. Numbers on nodes represent posterior probabilities and ultrafast bootstrap (as obtained in the ML analysis) support respectively. Asterisks represent nodal support larger than 95 % in both ML and Bayesian analyses. Two dashes in ultrafast bootstrap indicate the node was not recovered in the ML analysis (see Appendix 2 for the ML tree).

FIGURE 2. Holotype of Pristimantis leopardus sp. nov. (MHUA-A 8865), SVL 17.5 mm, adult male.

FIGURE 3. Specimens from different species in the Pristimantis leptolophus species group. A, Pristimantis lasalleorum, MHUA-A 8743 (Alto el Junco, Vereda Corcovado, Abriaquí, Antioquia, Colombia), adult male; B, Pristimantis leptolophus, MAR 2771 (Vereda las Mercedes, Corregimiento de Herrera Río Blanco, Tolima, Colombia), adult female; C, Pristimantis maculosus, ICN 55578 (Manizales, Caldas, Colombia), adult male; D, Pristimantis parectatus, MHUA-A 9977 (Páramo de los Cristos, Vereda Chaverras, Sonsón, Antioquia, Colombia) adult female; E, Pristimantis peraticus, MAR 2677, (Vereda Las Mercedes, Corregimiento de Herrera Río Blanco, Tolima, Colombia), adult female; F, Pristimantis scoloblepharus, MHUA-A 8875 (Páramo las Palomas, Vereda Manzanares, Sonsón, Antioquia, Colombia), adult male; G, Pristimantis stictus, (Vereda El Vergel, Marulanda, caldas, Colombia) adult male not collected; H, P. uranobates, MHUA-A 9867 (Páramo Valle Alto, Corregimiento San Félix, Salamina, Caldas, Colombia), adult female. Photos by J. M. Daza (A, D), M. Rivera-Correa (F), Marco Rada (B, E), G. González (C, G). (See Appendix 3 for specimens examined).

FIGURE 4. Variation in dorsal color patterns of life specimens of Pristimantis leopardus sp. nov. A, adult male, MHUA-A 8865, holotype, SVL 17.5 mm; B, adult male, MHUA-A 8858, paratype; C, adult male, MHUA-A 9126, paratype; D, adult male, MHUA-A 8856, paratype; E, adult male, MHUA-A 9127, paratype; F, adult male, MHUA-A 8848, paratype. Photos by José Fang.

FIGURE 5. Variations in ventral color patterns in life specimens of Pristimantis leopardus sp. nov. A, adult male, MHUA-A 8856, paratype; B, adult male, MHUA-A 8844, paratype; C, adult male, MHUA-A 9126, paratype; D, adult male, MHUA-A 8858, paratype. Photos by José Fang.

FIGURE 6. Map showing geographic distribution of species in the Pristimantis leptolophus group in Colombia. Pristimantis lasalleorum (green dots), P. leopardus sp. nov. (violet dots), P. leptolophus (yellow triangles), P. maculosus (light brown squares), P. parectatus (red dots), P. peraticus, (pink pentagons), P. scoloblepharus (white pentagons), P. stictus (light blue triangles), P. uranobates (blue squares). For localities see Appendix 4.

FIGURE 7. A, panoramic view of the type locality of Pristimantis leopardus sp. nov., Cerro La Vieja (Sonsón, Antioquia, Colombia); B, typical habitat of Pristimantis leopardus sp. nov.

FIGURE 8. Specimens of Pristimantis leopardus sp. nov. in their microhabitat (Bromelia sp.). A, individual (not collected) using a phytotelmata; B, individual on the bract of Bromelia sp.

Tables

TABLE 4. Morphological variation in measurements (in mm) within the type series of Pristimantis leopardus sp. nov. See text for abbreviations of measurements. Min = minimum value; Max = maximum value; Mean = arithmetic mean value; S. D. = standard deviation.

Measurement	Males (n = 30)		Female (n = 1)
	Min.–Max.	Mean±SD
SVL	16.0–19.9	(17.8±1.0)	22.8
HL	5.3–7.4	(6.5±0.4)	7.8
HW	6.0–8.1	(7.2±0.5)	9.2
ED	1.7–2.9	(2.5±0.3)	2.5
END	2.4–3.5	(3.0±0.2)	2.8
NSD	0.7–1.3	(1.0±0.1)	0.9
IND	1.3–2.1	(1.8±0.2)	2.0
AMD	2.8–4.1	(3.4±0.3)	4.5
TD	0.8–1.4	(1.0±0.1)	1.6
FAL	3.7–5.3	(4.5±0.4)	4.9
FAB	0.8–1.8	(1.3±0.2)	1.7
HAL	3.9–5.9	(5.1±0.4)	6.4
THL	8.1–10.6	(9.3±0.6)	11.1
TL	8.4–10.9	(9.5±0.6)	10.8
TAL	4.9–6.7	(5.9±0.4)	6.6
FL	6.5–9.4	(8.1±0.6)	10.5
TFD	0.6–1.1	(0.8±0.1)	0.9
FTD	0.5–1.0	(0.7±0.1)	0.9