Pristimantis jimenezi , Paez, Nadia B. & Ron, Santiago R., 2019
Paez, Nadia B. & Ron, Santiago R., 2019, Systematics of Huicundomantis, a new subgenus of Pristimantis (Anura, Strabomantidae) with extraordinary cryptic diversity and eleven new species, ZooKeys 868, pp. 1-112: 1
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Pristimantis jimenezi sp. nov.
English: Jiménez de la Espada’s Rain Frog. Spanish: Cutín de Jiménez de la Espada.
QCAZ 45170, an adult male from San Antonio, Cajas National Park border, Azuay Province, Ecuador (2.8835S, 79.4103W, 2099 m), collected by Elicio E. Tapia and Juan Carlos Morocho on August 21, 2009. Figures 15BView Figure 15, 18AView Figure 18.
(24: 11 males, 9 females, 4 juveniles). Ecuador: Azuay Province: QCAZ 45179, QCAZ 45186, adult females, QCAZ 45178, QCAZ 45187-189, adult males, from San Antonio, Cajas National Park border (2.8835S, 79.4103W, 2099 m), collected by Elicio E. Tapia and Juan Carlos Morocho on August 21, 2009; QCAZ 45199, adult female, QCAZ 45196, juvenile, from San Antonio, Cajas National Park border (2.8612S, 79.3787W, 2900 m), collected by Elicio E. Tapia and Juan Carlos Morocho on August 24, 2009; QCAZ 45202-203, QCAZ 45208-209, QCAZ 45212, adult females, QCAZ 45201, QCAZ 45204, QCAZ 45207, QCAZ 45211, adult males, QCAZ 45205-206, juveniles, from San Antonio, Malacatos river (2.9107S, 79.4146W, 1800 m), collected by Elicio E. Tapia and Juan Carlos Morocho on August 24, 2009; QCAZ 46955, adult male, from Chipla river (2.7457S, 79.4089W, 2500 m), collected by Elicio E. Tapia on January 16, 2010; QCAZ 46978, adult female, QCAZ 46977, QCAZ 46980, adult males, QCAZ 46979, juvenile, from Molleturo, Zadracay river (2.7353S, 79.4142W, 2674 m), collected by Elicio E. Tapia on January 20, 2010.
A member of the Pristimantis phoxocephalus group characterized by the following combination of characters: (1) skin on dorsum shagreen with or without scattered small subconical tubercles; faint middorsal fold; dorsolateral folds absent; thin lateral folds on anterior half of flanks; skin on venter areolate; discoidal fold absent; (2) tympanic membrane and tympanic annulus prominent, its upper and posterior margin concealed by supratympanic fold; (3) snout moderately long, acuminate with a fleshy keel in dorsal view, protruding in profile; (4) upper eyelid with small, distinct tubercles; cranial crests absent; (5) dentigerous processes of vomers low to prominent, oblique, moderately separated, posteromedial to choanae; (6) males bearing vocal slits and vocal sac, with white nuptial pads; (7) Finger I shorter than Finger II; discs of digits expanded to broadly expanded, elliptical to truncate; (8) fingers with broad lateral fringes; (9) ulnar tubercles low or absent; (10) heel bearing one medium subconical tubercle surrounded or not by lower tubercles; outer edge of tarsus bearing low subconical tubercles; inner edge with or without low tubercles, short inner tarsal fold present; (11) inner metatarsal tubercle ovoid, elevated, three times the size of elliptical, elevated outer metatarsal tubercle; supernumerary tubercles indistinct; (12) toes with broad lateral fringes; basal webbing present; Toe V longer than Toe III (disc on Toe III reaches or exceeds the proximal edge of penultimate subarticular tubercle on Toe IV; disc on Toe V reaches or exceeds the proximal edge of distal subarticular tubercle on Toe IV); toe discs slightly smaller than those on fingers, elliptical to truncate ( Fig. 7DView Figure 7); (13) in life, dorsal background light to dark brown with a scapular W-shaped marking and an interorbital stripe; groins and posterior surfaces of thighs salmon, pinkish brown, purplish brown or dark brown with small light brown to yellow spots; venter cream; iris reddish copper with black reticulations with a red medial streak to completely red ( Fig. 18View Figure 18); (14) average SVL in adult females: 35.0 ± 2.1 mm (30.9-37.4 mm; n = 9); in adult males: 25.5 ± 1.6 mm (21.9-27.1 mm; n = 12).
Comparison with other species.
Pristimantis jimenezi is similar to P. atillo , P. phoxocephalus , P. teslai sp. nov., P. torresi sp. nov., P. totoroi sp. nov., and P. verrucolatus sp. nov. It differs from P. atillo in the coloration of the groins and posterior surfaces of thighs (orange in P. atillo ; pinkish to dark brown with small light brown to yellow spots in P. jimenezi ), and flanks (black dots and flecks surrounding light reticulations in P. atillo ; lacking black dots in P. jimenezi ). Pristimantis jimenezi can be distinguished from P. phoxocephalus by its groin coloration (yellow with black reticulations in P. phoxocephalus ) and advertisement call. The call of P. phoxocephalus has shorter inter-note intervals and a lower frequency of the second harmonic and final frequency of the note (Table 6). Pristimantis teslai sp. nov. has tuberculate dorsal skin (shagreen in P. jimenezi ) and lacks lateral folds, which are present in P. jimenezi . The most similar species to P. jimenezi is P. torresi sp. nov., which has golden to beige iris with a red medial streak (copper to red in P. jimenezi ). P. totoroi sp. nov. has more prominent folds and tubercles than P. jimenezi , and its head length is larger (males Z = 3.05006, p = 0.0023, HL/SVL = 33.4-37.5% in P. jimenezi , 34.2-38.7% in P. totoroi sp. nov.; females Z = 2.85798, p = 0.0043, HL/SVL = 32.8-37.3% in P. jimenezi , 35.2-37.6% in P. totoroi sp. nov.), its tympanum smaller (males Z = -4.24763, p < 0.0001, TD/SVL = 4.8-6% in P. jimenezi , 4.4-5.1% in P. totoroi sp. nov.; females Z = -2,96383, p = 0.003, TD/SVL = 5-5.6% in P. jimenezi , 4.8-5.3% in P. totoroi sp. nov.), and its eyes smaller (males Z = -4.06051, p < 0.001, ED/SVL = 11.1-13.3% in P. jimenezi , 9.2-11.6% in P. totoroi sp. nov.; females Z = -2.43458, p = 0.0149, ED/SVL = 9.5-11.8% in P. jimenezi , 9.5-10.6% in P. totoroi sp. nov.) than those of P. jimenezi . Furthermore, advertisement calls of P. totoroi sp. nov. have shorter notes and lower dominant, second harmonic, and final frequencies of the notes than those of P. jimenezi (Table 6). Pristimantis verrucolatus sp. nov., which has an adjacent distribution to P. jimenezi at high elevations, can be distinguished by the presence of large tubercles and warts on flanks (absent in P. jimenezi ) and by its smaller size (males Z = 3.58643, p = 0.0003, SVL = 21.8-27.0 mm in P. jimenezi , 25.1-34.5 mm in P. verrucolatus sp. nov.; females Z = 2.00347, p = 0.0451, SVL = 31.1-37.4 mm in P. jimenezi , 40.4-46.8 mm in P. verrucolatus sp. nov.). Additionally, they can be distinguished by their advertisement calls: the call of P. verrucolatus sp. nov. consists of a single note, longer than the one or two notes of the call of P. jimenezi ; the dominant frequency is higher in advertisement calls of P. jimenezi , and the change in frequency along the note is lower in P. jimenezi (Table 6).
Description of the holotype.
An adult male ( QCAZ 45170, SC29041). Measurements (in mm): SVL 24.1; TL 12.7; FL 11.0; HL 8.2; HW 8.5; ED 2.8; TD 1.4; IOD 2.7; EW 2.5; IND 2.2; EN 2.6; TED 1.0. Body slender; head as wide as body, slightly wider than long; snout moderately long, acuminate with a vertical fleshy keel in dorsal view, protruding in profile; nostrils slightly protuberant, ovoid, directed laterally with slight dorsal inclination; canthus rostralis concave in dorsal view, angular in cross section; loreal region slightly concave, bearing small tubercles; upper eyelid bearing small distinct tubercles; cranial crests absent; tympanic membrane and annulus prominent with upper and posterior margins concealed by supratympanic fold; two medium-sized, rounded postrictal tubercles surrounded by smaller tubercles outlining supratympanic stripe. Choanae large, circular, not concealed by palatal shelf of maxilla; dentigerous processes of vomers prominent, oblique, moderately separated, posteromedial to choanae; each vomer bearing several indistinct teeth; tongue slightly wider than long, posterior border notched, posterior half free; vocal slits slightly curved, located at posterior half of mouth floor in between tongue and margin of jaw; vocal sac present.
Skin on dorsum and flanks shagreen; faint middorsal fold; row of three small middorsal tubercles on head; dorsolateral folds absent; thin lateral folds on anterior half of the flanks; skin on throat, chest, belly and ventral surfaces of thighs areolate; discoidal fold absent. One low round ulnar tubercle; white nuptial pads present; palmar tubercles prominent, outer palmar tubercle bifid, slightly bigger than ovoid thenar tubercle; subarticular tubercles distinct, rounded; low supernumerary tubercles at base of fingers; fingers with broad lateral fringes extended to outer edge of the palm; Finger I shorter than Finger II; discs on fingers expanded and elliptical; ventral pads on fingers surrounded by circumferential grooves ( Fig. 7DView Figure 7).
Hindlimbs slender; dorsal surfaces of hindlimbs with scattered low tubercles; posterior surfaces of thighs smooth, ventral surfaces of thighs areolate; heel bearing one medium prominent subconical tubercle surrounded by few lower rounded tubercles; outer edge of tarsus bearing low subconical tubercles; short inner tarsal fold present; inner metatarsal tubercle ovoid, elevated, three times the size of elliptical, rounded outer metatarsal tubercle; plantar surface with indistinct supernumerary tubercles; subarticular tubercles distinct, rounded; toes with broad lateral fringes; basal webbing present; discs smaller than those on fingers, wider on Toe IV and V; discs on toes expanded, elliptical; all toes having ventral pads and circumferential grooves; relative lengths of toes: I < II < III < V < IV; Toe V longer than Toe III (disc on Toe III exceeds proximal edge of the distal subarticular tubercle on Toe IV, disc on Toe V exceeds proximal edge of penultimate subarticular tubercle on Toe IV; Fig. 7View Figure 7). Coloration of the holotype in life and preservative is shown in Figures 15BView Figure 15, 18AView Figure 18.
Coloration of holotype in preservative. Dorsum grayish light brown with cream markings including an irregular W-shaped scapular mark and irregular reticulations; middorsal black stripe; head with interorbital stripe; faint canthal and labial bars; dark brown supratympanic stripe; flanks with the same background color as dorsum; flanks, groins, anterior, and posterior surfaces of thighs with cream medium-sized spots and cream oblique reticulations; background color of groins, anterior, and posterior surfaces of thighs lighter than dorsum; venter cream with scattered minute brown flecks on belly and brown midventral stripe ( Fig. 15BView Figure 15).
Coloration of holotype in life. Based on studio photographs. Dorsum light brown with cream markings including an irregular W-shaped scapular mark and irregular reticulations; head with interorbital stripe; faint gray canthal stripes and thin labial bars; black supratympanic stripe; flanks, groins, anterior, and posterior surfaces of thighs with yellow medium-sized spots and cream oblique reticulations; background color on groins and posterior surfaces of thighs purplish brown; ventral surfaces of limbs and chest pinkish cream; vocal sac yellowish cream; venter pinkish cream suffused with white; iris copper with thin black reticulations; light-blue sclera ( Fig. 18AView Figure 18).
Data of preserved individuals is based on 25 specimens of the type series; information of coloration in life is based on photographs from nine individuals. Variation of preserved and live individuals is shown in Figures 18View Figure 18, 19View Figure 19. Coloration in life is provided in parenthesis. Dorsum is creamy to light and dark brown, with or without lighter irregular marks or chevrons; some individuals show a middorsal band; a pale W-shaped mark on scapula is often present; head bears an interorbital stripe or band paler or darker than dorsum; most individuals bear labial bars, and a few, canthal bars (black to brown). Flanks often bear pale (light-brown to yellow) spots or reticulations all over them or confined to the groins and axils. Groins and posterior surfaces of thighs cream with small pale spots (salmon, pinkish brown, purplish, purplish brown or dark brown with small light-brown to yellow spots). Dorsal surfaces of thighs are darker than dorsum bearing pale (yellow or light brown) reticulations. Ventral surfaces of the body are cream (yellowish to pale cream) with or without dark flecks or reticulations on the belly and throat. Iris varies from copper with a red medial streak of variable size to completely red; it bears thin black reticulations; white to light-blue sclera.
Based on recordings of QCAZ 45178 (January 20, 2010; 19h10; 12 °C) and QCAZ 46977 (August 23, 2009; 16h30). The advertisement call of P. jimenezi consists of one or two peep-like notes ( Fig. 6AView Figure 6). The average duration of a note is 0.18 s (range 0.016-0.21 s), and the interval between them is 0.30 s (range 0.23-0.36 s). The fundamental and dominant frequencies of the notes are on average 2894 Hz (range 2776-3013 Hz). The peak time of the note occurs exactly in the middle of its duration. There is an increase in frequency along the note (2790-2977 Hz on average). Descriptive statistics for bioacoustic parameters are shown in Table 6.
Distribution, natural history, and conservation status.
This species is known from Western Andean slopes of Azuay Province between elevations of 1800 and 2900 m ( Fig. 1View Figure 1); this corresponds to the Western Montane Forest region. Individuals of this species inhabit primary and secondary forest. Specimens were found at afternoon and night on low vegetation up to 1.5 m above the ground and inside bromeliads up to 8 m above the ground. Some individuals were collected from banana plants. Gravid females and calling males were collected in January and August; males were found calling at 16 and 19 h.
Following B1ab(iii) IUCN criteria, we consider P. jimenezi to be Critically Endangered. Available records come from two localities (sensu IUCN 2017) where suitable surrounding habitats are disturbed by human settlements, agriculture, and cattle raising. Its Extent of Occurrence is estimated to be less than 100 km2 (39 km2).
The specific epithet is a noun in the genitive case and is a patronym for Marcos Jiménez de la Espada, a Spanish naturalist who visited South America between 1862 and 1865. During his trip, he made significant collections of amphibians including new species that he described between 1870 and 1875. Among others, he named the genus Pristimantis and 19 species of Ecuadorian amphibians. His species descriptions are remarkably detailed in comparison with those of his contemporaries. He also made accurate and detailed descriptions of the natural history and the external and internal morphology of Neotropical anurans.
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