Pristimantis bounides
sp. nov.
Figs 4
View FIGURE 4
& 5
View FIGURE 5
Pristimantis
sp. nov.
D in Lehr & von May (2017)
Common name. English: Hill Dweller Rubber Frog. Spanish: Rana cutín de montaña.
Holotype.
MUSM 31197
(IWU 182; Figs 4
View FIGURE 4
, 5
View FIGURE 5
), adult male from the buffer zone of the Pui Pui Protected Forest,
Quebrada Tasta
, "
Runda
", 11° 26' 51.9'' S, 74° 53' 11.9''W, 3463 m a.s.l. ( Fig. 2
View FIGURE 2
, 16
View FIGURE 16
B), Provincia Satipo, Región Junín, collected on 19 May 2012 by E. Lehr and R. von May.
GoogleMaps
Paratypes. A total of 6 ( Figs 6
View FIGURE 6
, 7
View FIGURE 7
), from Provincia Satipo, Región Junín: 2 males (
MUSM 31970
,
NMP 6
View Materials
V 75097
View Materials
[
GenBank
accession numbers
KY962797
View Materials
,
KY962790
View Materials
,
KY962774
View Materials
]) collected with the holotype; 4 females (
MUSM 31198
[
GenBank
accession numbers
KY962794
View Materials
,
KY962789
View Materials
,
KY962771
View Materials
], 31971,
NMP 6
View Materials
V 75066
View Materials
[
GenBank
accession numbers
KY962796
View Materials
,
KY962773
View Materials
]), 75540 [
GenBank
accession numbers
KY962795
View Materials
,
KY962772
View Materials
], from Sector Carrizal, Satipo-Toldopampa Road at km 134 on left side of street coming from Satipo ( Fig. 16
View FIGURE 16
A), 11° 29' 03.5'' S, 74° 53' 27.3''W, 3350 m a.s.l., collected on 23 June 2013 by E. Lehr, J. C. Cusi, and J. Moravec.
GoogleMaps
Generic placement. We assign this species to
Pristimantis
based on molecular evidence ( Fig. 3
View FIGURE 3
).
Diagnosis. A new species of
Pristimantis
assigned to the
Pristimantis danae
species Group having the following combination of characters: (1) Skin on dorsum shagreen with scattered tubercles, skin on flanks tuberculate, skin on venter areolate with scattered low tubercles; discoidal fold and thoracic fold present; irregularly shaped, discontinuous dorsolateral folds present; (2) tympanic membrane and tympanic annulus present; (3) snout moderate, rounded in dorsal and in lateral views; (4) upper eyelid with few enlarged conical tubercles; EW shorter than IOD; cranial crests absent; (5) dentigerous processes of vomers present; (6) males with vocal slits and visible subgular vocal sac, nuptial pads absent; (7) Finger I shorter than Finger II; discs of digits slightly expanded, round, having circumferential grooves; (8) fingers with narrow lateral fringes; (9) minute low ulnar and tarsal tubercles present; (10) heel with a small conical tubercle; inner tarsal fold absent; (11) inner metatarsal tubercle ovoid, 1.5 times as large as outer; outer metatarsal tubercle small, rounded; few low supernumerary plantar tubercles; (12) toes with narrow lateral fringes; basal toe webbing present; Toe V longer than Toe III; discs of digits slightly expanded, round, having circumferential grooves, toe discs slightly smaller than those on fingers; (13) in life, dorsal and lateral ground coloration pale grayish brown, orange brown, yellowish brown or reddish brown with dark grayish-brown marmorations, flanks paler than dorsum; canthal and supratympanic stripes grayish brown, not well defined, diffuse; groin pale grayish brown, orange brown, yellowish brown or reddish brown with or without gray flecks; venter pale gray, pale greenish gray or creamish white with or without dark gray mottling; iris pale grayish green with fine black vermiculation, a short vertical black streak from pupil across lower half of iris and a reddish tint horizontally across lower half of iris; (14) SVL in males 18.2–21.0 mm (n = 3), in females 21.6–24.4 mm (n = 4).
Comparisons.
Pristimantis bounides
sp. nov. is readily distinguished from its congeners in Ecuador (181 species), Peru (130 species), and Bolivia (17 species); AmphibiaWeb (2017), by having discontinuous dorsolateral folds, tuberculate flanks, an areolate venter, slightly expanded digits, and a polymorphic dorsal coloration consisting of pale grayish brown, orange brown, yellowish brown or reddish brown with dark grayish brown marmorations.
Pristimantis bounides
sp. nov. is similar to members of the former
Pristimantis orestes
species Group (considered to be not monophyletic by Duellman & Lehr 2009, not recognized by Padial et al. 2014). This group contained 17 species at high elevations in southern Ecuador and in Peru ( Guayasamin &Artega 2013; Duellman & Lehr 2009), which have narrow digits usually with circumferential grooves (except
P. simonsii
[ Boulenger, 1900]), and all (except
P. simonsii
) have a tympanum. The following five species are morphologically similar to
P. bounides
sp. nov.: Two species have discontinuous dorsolateral folds (
P. chimu Lehr, 2007
;
P. cordovae
[ Lehr & Duellman, 2007b]), whereas three have continuous dorsolateral folds (
P. seorsus Lehr, 2007
;
P. simonsii
,
P. ventriguttatus Lehr & Köhler, 2007
). However,
P. bounides
sp. nov. lacks cranial crests (present in
P. chimu
), has males with vocal slits (absent in
P. chimu
,
P. seorsus
and in
P. simonsii
), discs on digits slightly expanded, round (notched in
P. cordovae
, broadly expanded, truncate in
P. ventriguttatus
), and tarsal fold absent (present in
P. seorsus
and in
P. ventriguttatus
). Furthermore,
P. bounides
sp. nov. is phylogenetically distant from members of the former
P. orestes
Group (
P. melanogaster
[ Duellman & Pramuk, 1999],
P. orestes
[ Lynch, 1979],
P. simonbolivari
[ Wiens & Coloma, 1992],
P. simonsii
, Fig. 3
View FIGURE 3
).
Pristimantis bounides
sp. nov. shares with other members of the
Pristimantis danae
species Group ( Fig. 3
View FIGURE 3
), dorsolateral folds and males (unknown for
P. albertus
) which have vocal slits present (absent in
P. attenboroughi
) and nuptial pads absent. Furthermore, all but
P. attenboroughi
and
P. puipui
sp. nov. have a tympanum and discs with circumferential grooves.
Pristimantis bounides
sp. nov. is larger than
P. albertus
(female SVL 21.6–24.4 [n = 4] vs. 19.7–20.7 [n = 2] in
P. albertus
; Duellman & Hedges 2007) and has the belly colored pale gray, pale greenish gray or creamish white with or without dark gray mottling (cream with brown mottling in
P. albertus
).
Pristimantis bounides
sp. nov. has dorsal skin shagreen with scattered tubercle (smooth in
P. aniptopalmatus
) and ventral skin areolate with scattered low tubercles (smooth in
P. aniptopalmatus
and
P. ornatus
).
Pristimantis bounides
sp. nov. easily can be distinguished from
P. stictogaster
which has a white belly with prominent dark brown to black, irregularly shaped spots.
Pristimantis bounides
sp. nov. is most similar to three other species of the
Pristimantis danae
species Group from upper montane forests and puna of the PPPF and its surroundings ( Figs 2
View FIGURE 2
, 3
View FIGURE 3
):
P. attenboroughi
,
P. humboldti
sp. nov., and
P. puipui
sp. nov. All have discontinuous dorsolateral folds, tuberculate flanks, and an areolate venter.
Pristimantis bounides
sp. nov. and
P. humboldti
sp. nov. are sister taxa ( Fig. 3
View FIGURE 3
), and are of similar size (male SVL 18.2–21.0 mm, n = 3 in
P. bounides
sp. nov. vs. male SVL 17.2–20.6 mm, n = 3 in
P. humboldti
sp. nov.). Furthermore,
P. bounides
sp. nov. and
P. humboldti
sp. nov. have a tympamum (absent in
P. attenboroughi
and
P. puipui
sp. nov.), and have expanded discs with circumferential grooves (absent in
P. attenboroughi
and
P. puipui
sp. nov.). Males of
P. bounides
sp. nov.,
P. humboldti
sp. nov., and
P. puipui
sp. nov. have vocal slits (absent in
P. attenboroughi
). However,
P. bounides
sp. nov. has diffuse canthal and supratympanic stripes (well defined canthal and supratympanic stripes in
P. humboldti
sp. nov.), interorbital bar absent (present in
P. humboldti
sp. nov.), scapular without W- or X-shaped marks (W- or X-shaped marks present in
P. humboldti
sp. nov.), and the venter pale gray, pale greenish gray or creamish white with or without gray mottling (ventral coloration of throat, chest, belly, and anterior ventral surfaces of thighs dark gray and pale gray mottled with white and pale gray spots in
P. humboldti
sp. nov.).
Description of holotype. Head about as long as wide; head length 38% of SVL, head width 37% of SVL; cranial crests absent; snout moderate, rounded in dorsal and lateral views ( Figs 4
View FIGURE 4
A, B); eye-nostril distance 89% of eye diameter; nostrils slightly protuberant, directed dorsolateral; canthus rostralis rounded in lateral view, straight in dorsal view; loreal region slightly concave; lips rounded; outer margin of upper eyelid with few enlarged conical tubercles; upper eyelid width 54% of IOD; supratympanic fold short and narrow, extending from posterior margin of upper eyelid curved around posterior margin of tympanum; tympanic membrane present, tympanic annulus weakly defined, its upper and posterior margin concealed by supratympanic fold; two conical, slightly fused postrictal tubercles on each side of head. Choanae small, ovoid, not concealed by palatal shelf of maxilla; dentigerous processes of vomers positioned posterior to level of choanae, vomerine teeth minute, embedded in buccal mucosa, narrowly separated; tongue long, narrow, about three times as long as wide, not notched posteriorly, posterior half one third free; vocal slits slightly curved, located at posterior half of mouth floor between tongue and margin of jaw; subgular vocal sac distinct.
Skin on dorsum shagreen with scattered tubercles, skin on flanks tuberculate, irregularly shaped, discontinuous dorsolateral folds extending continuously from posterior level of tympanic area to level of mid body, extending discontinuously (separated into tubercles) to level of hind limb insertion; skin on throat smooth, chest and belly areolate with scattered low tubercles; discoidal fold and thoracic fold weakly defined; cloacal sheath short.
Outer ulnar surface of right arm with row of five minute low tubercles, outer ulnar surface of left arm bearing four tubercles; palmar tubercle bifid; thenar tubercle ovoid; subarticular tubercles weakly defined, most prominent on base of fingers, round in ventral view, subconical in lateral view; few, low supernumerary tubercles present, about one third size of subarticular tubercles; fingers with narrow lateral fringes, Finger I shorter than Finger II; nuptial pads absent; discs on digits of fingers slightly expanded, round, having circumferential grooves ( Fig. 5
View FIGURE 5
A).
Hind limbs short, slender, tibia length 45% of SVL; foot length 48% of SVL; dorsal surfaces of hind limbs tuberculate; inner surface of thighs smooth, posterior surfaces of thighs tuberculate, ventral surfaces of thighs areolate; heels each with a small conical tubercle; outer surface of tarsus with few scattered minute low tubercles; inner tarsal fold absent; inner metatarsal tubercle ovoid, one and a half times the size of round outer metatarsal tubercle; subarticular tubercles weakly defined, most prominent on base of toes, round in ventral view, subconical in lateral view; few low plantar supernumerary tubercles, about one third the size of subarticular tubercles; toes with narrow lateral fringes; basal webbing present; discs slightly expanded, round, less expanded than those on fingers, having circumferential grooves; relative length of toes: 1<2<3<5<4; discs on Toe III reaching middle of penultimate subarticular tubercle on Toe IV, Toe V surpassing penultimate subarticular tubercle on Toe IV, but not reaching distal subarticular tubercle on Toe IV; Fig. 5
View FIGURE 5
B).
Measurements of the holotype (in mm). SVL 18.2; tibia length 8.2; foot length 8.8; head length 6.9; head width 6.8; eye diameter 1.8; tympanum diameter 0.9; inter orbital distance 2.4; upper eyelid width 1.3; internarial distance 2.1; eye–nostril distance 1.6.
Characters
Pristimantis bounides
sp. nov.
Pristimantis humboldti
sp. nov.
Males (n = 3) Females (n = 4) Males (n = 3) Females (n = 6)
SVL 18.1–21.0 (20.0 ± 1.3) 21.6–24.4 (22.9 ± 1.3) 17.2–20.6 (18.9 ± 1.4) 19.7–25.7 (22.5 ± 1.9) TL 8.2–9.2 (8.8 ± 0.4) 9.9–10.8 (10.3 ± 0.4) 7.8–8.7 (8.3 ± 0.4) 9.2–10.6 (10.1 ± 0.4) FL 8.8–9.9 (9.2 ± 0.5) 9.9–12.0 (10.8 ± 0.9) 8.3–9.0 (8.9 ± 0.5) 9.5–11.2 (10.4 ± 0.5) HL 6.9–7.9 (7.5 ± 0.4) 7.6–8.9 (8.4 ± 0.5) 6.8–8.5 (7.4 ± 0.8) 7.5–8.7 (8.2 ± 0.4) HW 6.8–8.0 (7.6 ± 0.5) 7.7–8.5 (8.0 ± 0.3) 6.1–7.5 (6.9 ± 0.6) 6.9–8.7 (7.8 ± 0.5) ED 1.8–2.2 (2.0 ± 0.2) 2.1–2.5 (2.3 ± 0.2) 2.1–2.2 (2.1 ± 0.0) 2.2–2.6 (2.4 ± 0.2) TY 0.9–1.0 (0.9 ± 0.0) 1.0–1.1 (1.1 ± 0.1) 0.6–0.9 (0.8 ± 0.1) 0.8–1.0 (0.9 ± 0.1) IOD 2.4–2.7 (2.5 ± 0.1) 2.7–3.0 (2.8 ± 0.1) 2.3–2.7 (2.4 ± 0.2) 2.5–3.3 (2.9 ± 0.2) EW 1.3–2.0 (1.7 ± 0.3) 1.4–1.7 (1.6 ± 0.1) 1.3–1.7 (1.5 ± 0.2) 1.4–2.0 (1.7 ± 0.2) IND 2.0–2.3 (2.1 ± 0.1) 2.2–2.5 (2.3 ± 0.1) 1.9–2.3 (2.1 ± 0.2) 2.0–2.6 (2.4 ± 0.2) E–N 1.4–1.8 (1.6 ± 0.2) 1.6–1.9 (1.8 ± 0.1) 1.4–1.7 (1.6 ± 0.1) 1.5–2.2 (1.8 ± 0.2) TL/SVL 0.43–0.45 0.44–0.46 0.42–0.45 0.40–0.49
FL/SVL 0.42–0.48 0.46–0.50 0.46–0.48 0.41–0.49
HL/SVL 0.38 0.35–0.37 0.37–0.41 0.34–0.38
HW/SVL 0.37–0.38 0.33–0.36 0.35–0.38 0.33–0.36
HW/HL 1.00 0.9–1.0 0.9–1.0 0.92–1.0
E–N/ED 0.64–0.95 0.72–0.79 0.64–0.81 0.58–0.88
TY/ED 0.45–0.50 0.40–0.52 0.27–0.43 0.35–0.42
EW/IOD 0.54–0.76 0.50–0.63 0.48–0.74 0.50–0.71 Coloration of the holotype in life ( Fig. 4
View FIGURE 4
). Dorsal and lateral ground coloration pale yellowish brown with dark grayish-brown marmorations dorsally, flanks paler than dorsum. Canthal and supratympanic stripes grayish brown, not well defined, diffuse. Tympanum creamish white. Interorbital bar narrow grayish brown. Groin pale yellowish brown, anterior surfaces of thighs with a reddish tint. Throat and vocal sac pale yellowish brown, anterior half of chest pale grayish brown, posterior half of chest and belly pale creamish white, thighs pale grayish brown. Iris pale grayish green with fine black vermiculation, a short vertical black streak from pupil across lower half of iris and a reddish tint horizontally across lower half of iris.
Coloration of the holotype in preservative. Dorsal and lateral ground coloration pale tan with grayish-brown marmorations dorsally, flanks paler than dorsum. Canthal and supratympanic stripes grayish brown, not well defined, diffuse. Narrow grayish brown interorbital bar. Groin pale tan, anterior surfaces of thighs pale tan, posterior surfaces of thighs pale grayish brown. Throat, chest, and belly pale tan with small gray dots, and narrow gray midventral line across chest and belly. Lower arms pale tan with narrow gray midventral line, tarsi and ventral surfaces of thighs pale grayish brown and tan mottled. Iris pale gray with fine black vermiculation.
Variation. All paratypes ( Figs 6
View FIGURE 6
, 7
View FIGURE 7
) are similar to the holotype regarding morphology and proportions ( Tables 3, 4). Besides differences in SVL, notable morphological variation includes prominence of dorsolateral folds. Two females (MUSM 31971, NMP 6
V
View Materials
75540; Figs 7
View FIGURE 7
E, H) have prominent dorsolateral folds of pale grayish coloration on dark grayish brown dorsum. Most striking is the coloration polymorphism which resembles the color range of plants the frogs inhabit ( Fig. 16
View FIGURE 16
C). The dorsal and lateral ground coloration is pale grayish brown (MUSM 31971, NMP 6
V
View Materials
75540; Figs 7
View FIGURE 7
D, G), orange brown (NMP 6
V
View Materials
75097; Fig. 6
View FIGURE 6
B), pale yellowish brown (MUSM 31198; Fig. 7
View FIGURE 7
B) or reddish brown (MUSM 31970; Fig. 6
View FIGURE 6
E) with dark grayish-brown marmorations. The dorsum and flanks have reddish brown flecks in one male (MUSM 31970; Fig. 6
View FIGURE 6
E), pale yellowish brown in one female (MUSM 31198; Fig. 7
View FIGURE 7
B), and orange in one female (NMP 6
V
View Materials
75540; Fig. 7
View FIGURE 7
E). Flanks are paler than dorsum. Males have the throat and vocal sac pale grayish orange (MUSM 31970; Fig. 6
View FIGURE 6
F) or salmon ((NMP 6
V
View Materials
75097; Fig. 6
View FIGURE 6
C). The venter is pale gray (NMP 6
V
View Materials
75540, Fig. 7
View FIGURE 7
F; MUSM 31970, Fig. 6
View FIGURE 6
F; MUSM 31971, Fig. 7
View FIGURE 7
I; NMP 6
V
View Materials
75066), pale greenish gray (NMP 6
V
View Materials
75097; Fig. 6
View FIGURE 6
C) or creamish white (MUSM 31198; Fig. 7
View FIGURE 7
C) with or without dark gray mottling.
Etymology. The species epithet
bounides
is derived from the Greek noun “bounos” and means "dweller in the hills". The name refers to the habitat of montane forests where the new species was found.
Distribution, natural history, and threat status.
Pristimantis bounides
sp. nov. is known from two locations in the humid montane forest ecoregion ( Brack 1986, Figs 2
View FIGURE 2
, 16
View FIGURE 16
A, B): the type locality in the Tasta valley, "Runda" at 3463 m a.s.l (7.5 km [airline] from the border of PPPF), and the Satipo-Toldopampa Road at km 134 on left side of street coming from Satipo at 3350 m a.s.l. (4.2 km [airline] from the type locality). The vegetation at the type locality consists of Peruvian feather grass (
Stipa
ichu), large layers of mosses, small bushes, trees, and forest patches ( Fig. 16
View FIGURE 16
B). The vegetation is disturbed by cattle breeding and deforestation. The male holotype was found calling from the center of Peruvian feather grass in the early afternoon on 19 May 2012 during light rain. No sympatric anuran species were found at the type locality.
At km 134 of the Satipo-Toldopampa Road, calling males were heard from the moss layer in the early afternoon on 23 June 2013. On the same day a female
P. bounides
sp. nov. (NMP 6
V
View Materials
75540; Fig. 16
View FIGURE 16
C) guarding 20 eggs was found there inside moss. The eggs were pale cream colored and had an average diameter of 3.4 ± 0.1 mm (3.2–3.7 mm, n = 20). Sympatric anurans include
Phrynopus
sp. A (IWU 285) and
Gastrotheca griswoldi Shreve, 1941
(MUSM 31972).
We propose that
Pristimantis bounides
sp. nov. should be classified as "Data Deficient" according to the IUCN Red List criteria and categories (IUCN Standards and Petitions Subcommittee 2016) based on the low number of specimens known. Threats include agriculture and cattle breeding in both valleys, more acute in the Toldopampa valley than in the Tasta valley.