Pristimantis vrazi Moravec, Lehr, Wang & Uvizl, 2024

Lehr, Edgar, Moravec, Jiří, Wang, Yingtong & Uvizl, Marek, 2024, A new species of Pristimantis (Amphibia, Anura, Strabomantidae) from a montane forest of the Pui Pui Protected Forest in central Peru, ZooKeys 1219, pp. 143-163 : 143-163

Pristimantis vrazi Moravec, Lehr, Wang & Uvizl sp. nov.

Figs 4 , 5 , 6 , 7 , 8 , Tables 1 , 2

Type material.

Holotype. MUSM 41581 (field number IWU 200, Figs 4 , 5 , 7 ), GenBank accession numbers PQ 330255 (16 S), PQ 345842 (12 S); adult ♀; from buffer zone of the Pui Pui Protected Forest (11°05'44.2"S, 75°13'39.8"W), 1550 m a. s. l., property of Abelardo Cabrejos Vega, Distrito Pichanaqui, Provincia Chanchamayo, Región Junín, Peru; Edgar Lehr, Juan Carlos Cusi, Rudolf von May & Jiří Moravec leg., 10 June 2013 .

Paratype. MUSM 41582 (IWU 205, Fig. 6 ), GenBank accession numbers PQ 330256 (16 S), PQ 345843 (12 S); 1 subadult ♂; from the type locality; Edgar Lehr, Juan Carlos Cusi, Rudolf von May & Jiří Moravec leg., 10 June 2013 .

Referred specimens.

MUSM 31929 (IWU 197), MUSM 41583 (IWU 201), MUSM 31932 (IWU 204), MUSM 31933 (IWU 206) (Fig. 8 ), 4 juveniles; all from the type locality; Edgar Lehr, Juan Carlos Cusi, Rudolf von May, and Jiří Moravec leg.; 10 June 2013 • MUSM 31937 (IWU 214), MUSM 31983 (IWU 215), 2 juveniles; from the buffer zone of the Pui Pui Protected Forest (11°06'17.0"S, 75°12'26.2"W); 1730 m a. s. l.; Edgar Lehr, Juan Carlos Cusi, Rudolf von May, and Jiří Moravec leg.; 11 June 2013 .

Generic placement.

We assign this species to Pristimantis based on our molecular data (Fig. 3 ) and the general morphological similarity to other members of the genus.

Diagnosis.

A new species of Pristimantis not assigned to any species group having the following combination of characters: (1) skin on dorsum shagreen with scattered conical tubercles; skin on venter areolate; weak discoidal fold present; short dorsolateral ridges present; (2) tympanic membrane and tympanic annulus present; (3) snout moderately long, acutely rounded in dorsal view, rounded in profile; (4) upper eyelid bearing small conical tubercles; EW narrower than IOD; cranial crest absent; (5) dentigerous processes of vomers present, oblique; (6) vocal slits and nuptial pads absent; (7) finger I shorter than finger II; discs of digits broadly expanded, round, bearing circumferential grooves; (8) fingers with narrow lateral fringes; (9) ulnar tubercles small, round; (10) heel without tubercles; inner tarsal fold absent; inner edge of tarsus with low, elongated tubercles; (11) inner metatarsal tubercle ovoid, 4–5 times larger than outer elongated metatarsal tubercle; numerous supernumerary plantar tubercles; (12) toes with weak lateral fringes; basal toe webbing present; toe V longer than toe III; toe discs smaller than those on fingers, circumferential grooves present; (13) in life, the dorsum ranges from cream to dark grayish brown with a black W-shaped scapular fold, with or without cream blotches; anterior and posterior surfaces of thighs grayish brown with pale gray spots; groin grayish brown with pale-gray spots with or without a hint of olive-green; venter cream to gray with or without numerous small, brown, dark dots; iris pale bronze with fine black reticulation, a reddish brown median horizontal streak, and a narrow black median vertical streak on lower half of eye; head light to dark brown dorsally, with or without brown or black interorbital bar and canthal stripes, with or without white spot at dorsal tip of snout; (14) SVL in single subadult male 14.4 mm, in single adult female 26.4 mm.

Comparison.

Pristimantis vrazi is distinguished from its congeners in Peru (152 species; AmphibiaWeb 2024) by the following combination of characters: dorsum with conical tubercles and short dorsolateral ridges, tympanic membrane and tympanic annulus present, males without vocal slits and nuptial pads, dentigerous processes of vomer present, groin grayish brown with or without a hint of pale olive-green, and iris pale bronze with fine black reticulation, a reddish-brown median horizontal streak, and a narrow black median vertical streak on lower half of eye. The following 11 species of Pristimantis have been recorded from montane forests in the PPPF and its surroundings including the Distrito de Pampa Hermosa ( Moravec et al. 2020; Venegas et al. 2023; Lehr and Moravec 2024; this paper): Pristimantis albertus Duellman & Hedges, 2007 , P. aniptopalmatus ( Duellman & Hedges, 2005) , P. ashaninka , P. bipunctatus ( Duellman & Hedges, 2005) , P. clarae Venegas, García-Ayachi, Marchelie, Ormeño & Catenazzi, 2023 , P. croceoinguinis , P. cruciocularis , P. sagittulus ( Lehr, Aguilar & Duellman, 2004) , P. sinschi , P. cf. stictogaster ( Duellman & Hedges, 2005) , and P. sp.

Comparing to Pristimantis vrazi (characters in parentheses), P. albertus has distinct dorsolateral folds (short ridges), males with vocal slits (absent), and a yellow groin (grayish brown with pale-gray spots with or without a hint of olive-green) ( Duellman and Hedges 2007; Moravec et al. 2020). Pristimantis aniptopalmatus has females with SVL up to 22.0 mm (26.4 mm), males have vocal slits (absent), and the iris is grayish white (pale bronze with reddish-brown streak) ( Duellman and Hedges 2005). Pristimantis ashaninka is only known from the northeastern corner of the PPPF (northwestern corner), and both species have the dorsum with conical tubercles and the venter pale gray and mottled grayish brown. However, P. ashaninka lacks a tympanum (present), has the dorsum with a reddish-brown blotch in shape of an hourglass (absent), and the iris with a black narrow upper and lower streak (only lower streak present) ( Lehr and Moravec 2017). Pristimantis vrazi and P. bipunctatus have the iris bronze, with a median horizontal reddish-brown streak. However, P. bipunctatus has females with SVL up to 41.5 mm (26.4 mm), males with vocal slits and nuptial pads (absent), and the scapular with a pair of black warts (absent) ( Duellman and Lehr 2009). Pristimantis clarae was recently described from the Distrito de Pampa Hermosa, but it is a member of the P. danae group and differs in having distinct dorsolateral folds (short ridges), heals with low conical tubercles (absent), males with vocal slits (absent), coppery iris (pale bronze with reddish-brown streak), and dorsum with triangular blotches (dorsum dark grayish brown) ( Venegas et al. 2023). Pristimantis croceoinguinis is known from montane forests in the north-central buffer zone and the eastern corner of the PPPF, and it also has the iris with a reddish-brown median streak and a vertical dark-brown streak at lower half of the eye. However, P. croceoinguinis lacks a tympanum (present), and has the groin yellow (grayish brown) ( Lynch 1968). Pristimantis vrazi and P. cruciocularis occur syntopically in the surroundings of the PPPF. However, P. cruciocularis lacks a tympanum (present), fingers and toes lack lateral fringes (present), has a red or orange groin (cream to dark brown with or without a hint of olive-green), and has the iris with a dark-brown cross (vertical streak at lower half or iris) ( Lehr et al. 2006).

Pristimantis sagittulus from the northwestern part of the PPPF has dentigerous processes of vomers absent (present), males with vocal slits and nuptial pads (both absent), and posterior surfaces of thighs with longitudinal red stripes (no red skin coloration) ( Duellman and Lehr 2009). Pristimantis stictogaster has vocal slits present (absent), and dark-brown groin with white spots (grayish brown with pale-gray spots with or without a hint of olive-green), and white belly with dark-brown spots (venter cream to gray with or without numerous small dark-brown dots) ( Duellman and Lehr 2009). Pristimantis platydactylus is known for its remarkable variation in color pattern ( De la Riva 1998; Lehr et al. 2006; Duellman and Lehr 2009). However, P. platydactylus has the tympanic annulus barely evident (distinct), and males have vocal slits and nuptial pads (both absent).

Our phylogeny (Fig. 3 ) revealed close phylogenetic similarities of P. vrazi with P. lindae , P. quaquaversus , P. rhabdocnemus , P. sinschi , and P. sp. Pristimantis vrazi and P. lindae have the iris with a red median horizontal streak and fingers and toes with lateral fringes. However, P. lindae has the dorsum shagreen (tuberculate), males with nuptial pads and vocal slits (absent), and a prominent tympanum (not prominent) ( Duellman 1978). Pristimantis quaquaversus has males with vocal slits (absent), heel with a conical tubercle (absent), tympanic membrane absent (present), and the venter white with or without brown spots (gray and brown mottled) ( Lynch 1974). Pristimantis vrazi and P. rhabdocnemus are related as sister taxa in our phylogeny (Fig. 3 ), are of similar size, and have a grayish-brown dorsal ground coloration. However, P. rhabdocnemus lacks a tympanum (present), lacks dentigerous processes of vomers (present), and has a grayish-tan iris (iris pale bronze with fine black reticulation, a reddish-brown median horizontal streak, and a narrow black median vertical streak on lower half of eye) ( Duellman and Hedges 2005). Pristimantis vrazi and P. sinschi have the dorsum with conical tubercles, the venter areolate, and males that lack vocal slits and nuptial pads, and the iris pale bronze with fine black reticulation and broad median red or reddish-brown band through pupil and a narrow black vertical streak from pupil across lower half of iris ( Moravec et al. 2020; this paper). However, P. sinschi lacks a tympanum (present), lacks a pale brown tympanic stripe (present), has the groin black with cream blotches (cream to dark brown with or without a hint of olive-green), and venter mottled black and cream (pale gray and brown mottled).

Holotype description.

Adult female (Figs 4 , 5 ); head narrower than body, slightly longer than wide; head width 36.4 % of SVL; head length 33.3 % of SVL; cranial crest absent; snout acutely rounded in dorsal view, round and moderate in length in lateral view, E – N 78.3 % of eye diameter; nostrils protuberant, directed dorsolaterally; canthus rostralis straight in dorsal view, rounded in profile; loreal region slightly concave; lips rounded; upper eyelid bearing several small, conical tubercles; EW 85.2 % of IOD; supratympanic fold distinct, short, extending diagonally from posterior margin of upper eyelid towards insertion of arm, covering upper and posterior margin of the tympanum; tympanic membrane and tympanic annulus present, more distinct on the right side of head, tympanum 29.3 % of ED; several conical postrictal tubercles present bilaterally, some fusing to short ridges. Choanae small, ovoid; dentigerous processes of vomers oblique and round; tongue longer than wide, not notched posteriorly, posterior one-third free.

Skin on dorsum (Fig. 4 B ) shagreen, with many scattered tubercle, with short, weakly defined dorsolateral fold on anterior half of body (Fig. 4 A ); skin on flanks shagreen with scattered tubercles (Fig. 4 C ); skin on throat and chest smooth, belly areolate (Fig. 4 D ); discoidal and thoracic folds weakly defined; cloacal sheath short.

Outer ulnar surface with numerous minute tubercles; outer palmar tubercle round and slightly bifid, inner palmar tubercle ovoid, approximately one-half size of outer tubercle; distinct supernumerary tubercles, ovoid, approximately one-half size of subarticular tubercles; subarticular tubercles well defined, round in ventral view, conical in lateral view; fingers with narrow lateral fringes; finger I shorter than finger II; discs on fingers broadly expanded, rounded, bearing circumferential grooves (Fig. 7 A ).

Hind limbs long, slender, tibia length 53.1 % of SVL; foot length 43.6 % of SVL; upper surface of hind limbs shagreen with many scattered tubercles; anterior surface of thighs smooth, posterior and ventral surfaces of the thighs areolate; heels without conical tubercles; outer surface of tarsus with minute tubercles; inner tarsal fold not present; inner metatarsal tubercle ovoid, 4–5 times size of ovoid outer metatarsal tubercle; subarticular tubercles well defined, round in ventral view, conical in lateral view; few plantar supernumerary tubercles distinct, about one-fourth size of subarticular tubercles; toes with narrow lateral fringes; basal webbing present, most prominent between toes IV and V; discs expanded, slightly truncated, slight smaller as discs on fingers, bearing circumferential grooves; relative length of toes: I < II < III < V < IV (Fig. 7 B ).

In life (Fig. 4 ), dorsum dark grayish brown with a black W-shaped scapular fold; head dorsally with a pale brown interorbital bar and a pale cream blotch on snout, head laterally grayish brown with a dark-brown canthal stripe, a dark-brown supratympanic stripe, two dark-brown labial bars below eye and distinct pale brown tympanic stripe from posterior outer corner of upper eyelid to corner of mouth; arms dorsally grayish brown with a dark-brown diagonal bar on lower arm; hind limbs dorsally grayish brown with three dark-brown diagonal bars; flanks pale brown with three diagonal dark grayish-brown bars alternating with three pale-brown diagonal to vertical bars; groin, anterior and posterior surfaces of thighs grayish brown with pale-gray spots; throat, chest, belly, and extremities pale-gray and dark-brown mottled; finger and toe discs dorsally pale reddish brown; iris pale bronze with fine black reticulation, a reddish-brown median horizontal streak, and a narrow black median vertical streak at lower half of eye.

In alcohol (Fig. 5 ) after 10 years, the dorsal ground coloration is pale brown with dark-brown flecks, dark brown W-shaped ridge in scapular area, cream interorbital bar and cream blotch on snout; flanks cream with three pale-brown stripes; groin, anterior and posterior surfaces of thighs cream; thighs and tibias ventrally cream, throat, chest, and belly cream and pale grayish-brown mottled; iris pale gray.

Holotype measurements (in mm). SVL 26.4; TL 14.0; FL 11.5; HL 8.8; HW 9.6; ED 3.7; TY 1.1; IOD 2.7; EW 2.3; IND 2.1; E – N 2.9.

Variation.

The single subadult male paratype MUSM 41582 (Fig. 6 ) has following measurements (in mm): SVL 14.4; TL 8.2; FL 6.3; HL 5.0; HW 5.4; ED 2.3; TY 0.4; IOD 2.2; EW 1.2; IND 1.2; E – N 1.6. It has strongly tuberculated dorsum (tubercles of conical shape), dark grayish-brown dorsum with reddish-brown flecks. The flanks bear three dark grayish-brown bars (Fig. 6 C ). The six juvenile referred specimens (Fig. 8 , SVL = 11.9–14.3 mm) have the dorsum tuberculate, the venter pale to dark gray, with brown mottling, the iris pale bronze with fine black reticulation, a reddish-brown median horizontal streak, and a narrow black median vertical streak on lower half of the eye, and a distinct pale-cream tympanic stripe (Fig. 8 C ). One specimen ( MUSM 31929, Fig. 8 A ) has a middorsal pale cream stripe that is narrow on the head and wider on its back.

Etymology.

We dedicate this new species to the Czech explorer and patriot Enrique Stanko Vráz (1860–1932), who explored Africa, South America, and eastern Asia ( Todorová 2006). In South America, he spent several years working in Venezuela and traveling by boat from Venezuela to Peru via the Rio Orinoco, Rio Negro, and Rio Amazonas before crossing the Andes and working around Cajamarca. His specimen collections (animals, artifacts) were sent to the National Museum in Prague. He published his travels and observations in South America in a book ( Vráz 1900) that provides valuable original insights into South American nature and indigenous peoples at the end of the 19 th century and still inspires people. The specific epithet is used as a noun in apposition.

Distribution and natural history.

Pristimantis vrazi is only known from two localities of the northwestern corner of the buffer zone of the Pui Pui Protected Forest (Fig. 2 ) between 1550 and 1730 m a. s. l. The type locality lies in the valley of Rio Huatziroqui (Fig. 9 ). The slopes of the valley are covered by primary mountain rainforest characterized by a 15–20 m high canopy. The type specimens were collected on the right bank of the river. They inhabited a disturbed forest that was interspersed with small coffee plantations, solitary houses of local coffee farmers, and various forest paths. All specimens were found at night on low vegetation (up to 80 cm above ground). The holotype was found at 8: 55 p. m. on a leaf 80 cm above ground. Other syntopic frogs include Pristimantis bipunctatus , P. cruciocularis , and the hylid Boana aguilari . Considering the sparse data available, we here classify P. vrazi as Data Deficient according to the IUCN Red List criteria.

Figures

Figure 2. Map of the Pui Pui Protected Forest with the collecting sites of Pristimantis vrazi sp. nov.

Figure 3. Maximum-likelihood tree of reconstructed phylogenetic relationships of the Pristimantis frogs based on the concatenated dataset 12 S and 16 S sequences. Branch support values are shown by black dots on the nodes. The numbers at the tips represent either voucher numbers or GenBank accession numbers. Branch support values are shown by black dots on the nodes. The clade outlined with a dotted line includes the fourth sample molecularly processed in this study, along with other species such as P. croceoinguinis, P. altamazonicus, P. ashaninka, P. cruciocularis, and P. platydactylus.

Figure 4. Life adult female holotype (MUSM 41581, SVL = 26.4 mm) of Pristimantis vrazi sp. nov. A dorsolateral view B dorsal view C lateral view D ventral view. Photos by E. Lehr.

Figure 5. Preserved holotype (MUSM 41581, SVL = 26.4 mm) of Pristimantis vrazi sp. nov. A dorsal view B ventral view. Photos by Y. Wang.

Figure 6. Life subadult male paratype (MUSM 48582, SVL = 14.4 mm) of Pristimantis vrazi sp. nov. A dorsolateral view B dorsal view C lateral view D ventral views. Arrows indicate the three dark grayish-brown bars on the flank. Photos by E. Lehr.

Figure 7. Holotype (MUSM 41581) of Pristimantis vrazi sp. nov. A ventral view of hand B ventral view of foot, preserved toes are not on the same plane, resulting in an artifact with toe V appearing thinner than it is. Drawings by Y. Wang.

Figure 8. Juvenile referred specimens of Pristimantis vrazi sp. nov. A MUSM 31929, SVL = 12.3 mm B MUSM 31933, SVL = 12.0 mm C MUSM 41583, SVL = 14.1 mm D MUSM 31937, SVL = 11.9 mm E MUSM 31932, SVL = 14.3 mm F MUSM 31938, SVL = 12.1 mm. Photo A by J. Moravec, all others by E. Lehr.

Figure 9. Valley of the Rio Huatziroqui in the buffer zone of the Pui Pui Protected Forest, type locality of Pristimantis vrazi sp. nov. Type specimens were collected along the river at elevations between 1550 m and 1730 m a. s. l. Photo by J. Moravec.