Pristimantis mallii, Reyes-Puig, Carolina, Pablo Reyes-Puig, Juan, A. Velarde-Garcez, Daniel, Nicolas Davalos,, Mancero, Emilio, Jose Navarrete, Maria, H. Yanez-Munoz, Mario, F. Cisneros-Heredia, Diego & R. Ron, Santiago, 2019

Reyes-Puig, Carolina, Pablo Reyes-Puig, Juan, A. Velarde-Garcez, Daniel, Nicolas Davalos,, Mancero, Emilio, Jose Navarrete, Maria, H. Yanez-Munoz, Mario, F. Cisneros-Heredia, Diego & R. Ron, Santiago, 2019, A new species of terrestrial frog Pristimantis (Strabomantidae) from the upper basin of the Pastaza River, Ecuador, ZooKeys 832, pp. 113-133 : 117-125

Pristimantis mallii sp. n. Figures 2, 3, 4, 5, 6

Holotype.

QCAZ 52473 (field no. SC-PUCE 35222; Figs 3, 4), adult female, collected by Fernando Ayala, Diego Paucar, Yerka Sagredo, Juan Pablo Reyes-Puig, Fausto Recalde, Luis Recalde and Santiago Recalde on January 17, 2012 at Reserva Río Zuñag, Fundación EcoMinga, Baños, province of Tungurahua, Ecuador (1.36740S, 78.14573W; 2140 m elev.).

Paratypes

(7 females, 12 males). QCAZ 39777, adult female, collected by Diego Páez on January 1, 2009 at Reserva Río Zuñag, Fundación EcoMinga, Baños, province of Tungurahua, Ecuador (1.349399S, 78.15870W; 2127 m elev.). QCAZ 52476, 52477, adult females, collected by Fernando Ayala, Diego Paucar, Yerka Sagredo, Juan Pablo Reyes-Puig, Fausto Recalde, Luis Recalde and Santiago Recalde on January 17, 2012 at Reserva Río Zuñag, Fundación EcoMinga, Baños, province of Tungurahua, Ecuador (1.36761S, 78.14584W; 2153 m elev.). QCAZ 52494, adult female, collected by Fernando Ayala, Diego Paucar, Yerka Sagredo, Juan Pablo Reyes-Puig, Fausto Recalde, Luis Recalde and Santiago Recalde on January 17, 2012 at Reserva Río Zuñag, Fundación EcoMinga, Baños, province of Tungurahua, Ecuador (1.37220S, 78.15386W; 1823 m elev.). DHMECN 5236, 5264, adult females, collected by Mario Yánez-Muñoz, Miguel Urgilés and Andrés Laguna on May, 2008 at Reserva Río Zuñag, Baños, Province of Tungurahua, Ecuador (1.40045S, 78.186776W; 1300 m). ZSFQ 1305, adult female, collected by Carolina Reyes-Puig, Nicolás Dávalos, Daniel Velarde and Emilio Mancero on October 7, 2017 at at Reserva Río Zuñag, Fundación EcoMinga, Baños, province of Tungurahua, Ecuador (1.36761S, 78.14583W; 2190 m elev.). QCAZ 52512, subadult male, collected by Fernando Ayala, Diego Paucar, Yerka Sagredo, Juan Pablo Reyes-Puig, Fausto Recalde, Luis Recalde and Santiago Recalde on January 20, 2012 at Reserva Río Zuñag, Fundación EcoMinga, Baños, province of Tungurahua, Ecuador (1.37513S, 78.16363W; 1532 m elev.). QCAZ 52471, 52474, adult males with the same data of the holotype. QCAZ 52478, adult male, collected by Fernando Ayala, Diego Paucar, Yerka Sagredo, Juan Pablo Reyes-Puig, Fausto Recalde, Luis Recalde and Santiago Recalde on January 17, 2012 at Reserva Río Zuñag, Fundación EcoMinga, Baños, province of Tungurahua, Ecuador (1.36761S, 78.14583W; 2146 m elev.). QCAZ 52480, 52481, adult males, collected by Fernando Ayala, Diego Paucar, Yerka Sagredo, Juan Pablo Reyes-Puig, Fausto Recalde, Luis Recalde and Santiago Recalde on January 17, 2012 at Reserva Río Zuñag, Fundación EcoMinga, Baños, province of Tungurahua, Ecuador (1.36765S, 78.14594W; 2135 m elev.). DHMECN 5233-5235, 5238, adult males, collected by Mario Yánez-Muñoz, Miguel Urgiles and Andrés Laguna on May, 2008 at Reserva Río Zuñag, Baños, Province of Tungurahua, Ecuador (1.40045S, 78.186776W; 1269 m elev.). ZSFQ 1306, 1327, adult males, collected by Carolina Reyes-Puig, Nicolás Dávalos, Daniel Velarde and Emilio Mancero on October 7, 2017 at Reserva Río Zuñag, Fundación EcoMinga, Baños, province of Tungurahua, Ecuador (1.36761S, 78.14583W; 2190 m elev.).

Generic placement.

We assign the new species in Pristimantis based on our molecular data (Fig. 1).

Diagnosis.

A new species of Pristimantis having the following combination of characters: (1) skin on dorsum and flanks shagreen, with rounded tubercles scattered towards the axillary region, with ") (" shaped scapular folds (evident in life); dorsolateral folds absent; skin on venter areolate; discoidal fold slightly defined; (2) tympanic membrane and tympanic annulus present, round, equivalent to 45% of ED; supratym panic fold present; (3) snout broadly rounded in dorsal view, moderate in length and rounded in lateral view; (4) upper eyelid with one or two subconical tubercles on the center of eyelid and some rounded tubercles (less evident in preserved specimens); EW 100% of IOD; cranial crests absent (5) dentigerous processes of vomers oblique in outline, with five to seven teeth, moderately separated, posteromedial to choanae; (6) vocals slits and nuptial pads present; (7) Finger I shorter that Finger II; discs of digits expanded, truncate; two times the width of the digits on Fingers III and IV; (8) fingers with lateral fringes; (9) ulnar tubercles present, rounded; (10) heel bearing one or two subconical tubercles (less evident in preserved specimens) surrounded by few lower rounded tubercles; inner tarsal fold present, it extends up to 1/4 of the tarsus; (11) inner metatarsal tubercle oval, 5 –6× as large as outer metatarsal tubercle that is subconical; supernumerary plantar tubercles indistinct; (12) toes with slightly defined lateral fringes; webbing absent; Toe V longer that Toe III, disc on Toe V reach the distal subarticular tubercle on Toe IV; (13) in life, dorsum and flanks light brown to brown, with irregular dark brown marks bounded by dirty cream, light brown or greenish cream; hidden surfaces of thighs brown splashed with dirty cream; groin with irregular yellowish marks; venter light gray or cream, spotted to densely spotted with brown. Golden coppery iris with black reticulations and a reddish-brown horizontal stripe; (14) SVL in adult males 16.7 ± 4.5 (11.6-21.3 mm); females with 27.6 ± 3.9 (22.6-34.3).

Comparisons with other species.

Pristimantis mallii is most similar to its sister species P. miktos ( Ortega-Andrade and Venegas 2014) (characters in parentheses) from the eastern lowlands of Ecuador. However, P. mallii can be easily distinguished for having ") (" shaped scapular folds (W- or X-shaped scapular fold); snout broadly rounded in dorsal view (subacuminate); upper eyelid bearing one or two subconical tubercles and some rounded tubercles (one small non-conical tubercle); dentigerous processes of vomers with 5-7 teeth (2 or 3 teeth); vocal slits in males present (absent); lateral fringes present (absent); dorsum and flanks light brown to brown, with irregular dark brown marks bounded by dirty cream, light brown or greenish cream (dorsum reddish-brown with some greenish-orange stains in scapular region, with or without yellowish-pale spots); golden coppery iris with black reticulations and a reddish-brown horizontal stripe (deep orange finely reticulated with black).

Other species of Pristimantis from the eastern lowlands of Ecuador, that can be confused with the new species by having dermal ridges in the scapular region, are P. kichwarum ( Elmer and Cannatella 2008) and P. luscombei ( Duellman and Mendelson 1995). Nonetheless, these two species have W-shaped dermal ridges (") (" shaped fold in P. mallii ); snout subacuminate in dorsal view (broadly rounded in P. mallii ); ulnar tubercles absent or low (present, rounded in P. mallii ); and nuptial pads in males absent (present in P. mallii ). Other species of rain frogs from the eastern Andean slopes of Ecuador that are morphologically similar to Pristimantis mallii are P. marcoreyesi (Reyes-Puig et al., 2014), P. yanezi (Navarrete et al., 2016) and P. spinosus (Lynch, 1979). In males of Pristimantis marcoreyesi , P. yanezi and P. spinosus the vocal slits and nuptial pads are absent (present in P. mallii ). The snout in P. marcoreyesi and P. spinosus is subacuminate in dorsal view (broadly rounded in P. mallii ); furthermore, P. marcoreyesi has dorsolateral folds slightly defined (absent in P. mallii ), P. spinosus has the skin of dorsum finely tuberculate (shagreen in P. mallii ), and the groin is black enclosing white spots (groin with irregular yellowish marks in P. mallii ). Besides, P. yanezi can be distinguished by having one conical tubercle on the upper eyelid (one or two subconical in P. mallii ); discoidal fold absent (present, slightly defined in P. mallii ); fingers and toes without lateral fringes (present in P. mallii ); dorsum yellowish brown to dark brown with scattered pale brown or orange blotches and black flecks, bearing a faint mid-dorsal hourglass-shaped band (dorsum and flanks light brown to brown, with irregular dark brown marks bounded by dirty cream, light brown or greenish cream).

Description of the holotype.

Adult female. Measurements (in mm): SVL 28.7; tibia length 15.6; foot length 15.2; head length 11.7; head width 12.8; eye diameter 4.7; tympanum diameter 2.5; interorbital distance 3.6; upper eyelid width 3.7; internarial distance 3.5; eye–nostril distance 4.6. Head slightly wider than long (12.8 mm vs 11.7); head width 44.4% of SVL; head length 41% of SVL; snout broadly rounded in dorsal view, moderate in length and rounded in lateral view (Fig. 4); eye–nostril distance 16% of SVL; canthus rostralis slightly concave in lateral view, nostrils directed laterally; interorbital area flat, as wide as the upper eyelid; cranial crests absent; upper eyelid with one or two subconical tubercles (reduced by preservation effects), and some rounded tubercles; upper eyelid width 100% of IOD; tympanic membrane differen tiated, tympanic annulus present, with upper margins covered by a supratympanic fold; tympanum diameter 54% of eye diameter; three subconical postrictal tubercles. Choanae moderately in size, with a drop-shaped outline, not concealed by palatal shelf of maxilla; dentigerous processes of vomer oblique in outline, moderately separated, posteromedial to choanae, with six to seven teeth; tongue wider than long, notched posteriorly, approximately 40% of it fixed to the mouth floor.

Skin on dorsum and flanks shagreen, with rounded tubercles scattered towards the axillary region, with ") (" shaped scapular folds (evident in life); dorsolateral folds absent; skin on venter areolate; discoidal fold slightly defined; cloaca with rounded tubercles on the inferoposterior margin. Forearms slender, ulnar tubercles present, rounded; palmar tubercle heart-shaped, bilobed, approximately twice the size of oval thenar tubercle (the tubercles are slightly defined); subarticular tubercles rounded, defined, few supernumerary tubercles, indistinct; fingers with narrow lateral fringes; Finger I shorter than Finger II; disc on Finger I rounded and on Finger II expanded, twice the width of the digits on Fingers III and IV, truncate; pads on fingers well defined by circumferential grooves on all fingers (Fig. 4).

Hindlimbs slender, tibia length 54% of SVL; foot length 53% of SVL heel; upper surfaces of hindlimbs shagreen; posterior surfaces of thighs smooth, ventral surfaces areolate; heel bearing one or two subconical tubercles (less evident by preservation effect) surrounded by few lower rounded tubercles; inner tarsal fold present, it extends up to 1/4 of the tarsus; inner metatarsal tubercle oval, 5 –6× as large as outer metatarsal tubercle that is subconical; supernumerary plantar tubercles indistinct; toes with slightly defined lateral fringes; webbing absent; discs on Fingers I and II rounded, discs in Fingers III, IV and V expanded, twice the width of the digit; all toes with ventral pads well defined by circumferential grooves; Toe V longer that Toe III, disc on Toe V reach the distal subarticular tubercle on Toe IV (Fig. 4).

Color of holotype in life.

(based on digital photographs, Fig. 2) Dorsum light brown with irregular brown marks bounded by dirty cream; black ") (" shaped scapular fold; head with dirty cream marks, one located behind the nostrils; flanks with brown oblique stripes delineated of dirty cream, with marbled brown marks concentrated towards axillary region; with two diagonal dark brown subocular stripes. Ventral areas of body, limbs and palms cream spotted with minute brown dots; throat cream spotted with brown dots, aggregates towards the outer edge of the jaw; forearms and hindlimbs with transversal brown bars separated by light brown interspaces; posterior surfaces of thigh dark brown; groin with irregular pale-yellowish marks. Golden coppery iris with black reticulations and a reddish-brown horizontal stripe.

Color of holotype in ethanol 70%.

(Fig. 3) Dorsum light brown with irregular brown marks slightly bounded by cream; black ") (" shaped scapular fold; with two black longitudinal lines above and behind the tympanum; with two diagonal brown subocular stripes; dorsal surfaces limbs, fingers and toes with transversal brown bars separated by dirty cream; the anterior surfaces of flanks light brown, with marbled brown marks concentrated towards axillary region; posterior surfaces of flanks and groin grayish cream. Ventral areas of body, limbs and palms cream spotted with minute brown dots aggregates towards the outer edge of the jaw, carpus, ulnar surfaces, flanks, posterior surfaces of thigh and tarsus. Golden olive iris.

Variation.

Preserved individuals (Figs 5, 6). In the type series, adult males (10.2-21.3 mm) are smaller than females (22.6-34.3). See Table 1 for measurements of the type specimens. Males have vocals slits located in the posteromedial region of the floor of the mouth; and nuptial pads located in the lower external portion of the Finger I. The ") (" shaped scapular fold is present in all individuals, but is black in all females, while it is not in some males (ZSFQ 1306, QCAZ 52481) (Figs 5, 6). Background coloration varies from gray or light brown to brown. Marks on dorsum and flanks are similar in all the type series, except for the adult males (ZSFQ 1306, QCAZ 52481) that have a dorsum without irregular marks and exhibit an internarial cream brand (Fig. 6). Some males present a gray patch on the head between the narinal and postorbital region (QCAZ 52471, QCAZ 52474, QCAZ 52478, and QCAZ 52512); this pattern is not present in any female (Figs 5, 6). The males (QCAZ 52512 and DHMECN 5234) have a gray spot in the middorsal region (Fig. 6). One male presents a dorsal pattern with longitudinal stripes on the dorsum (QCAZ 52480) (Fig. 6). In general, males have more variable dorsal patterns than females. Ventral coloration varies from cream to light brown; from slightly spotted (ZSFQ 1305, QCAZ 52473) to roughly spotted with brown (DHMECN 5236, QCAZ 52474, QCAZ 52212) (Figs 5, 6).

Coloration in life.

(based on digital photographs of the type specimens, Fig. 2).

Dorsum and flanks light brown (QCAZ 52473) to brown (QCAZ 52494, QCAZ 52512), with irregular dark brown marks bounded by dirty cream (QCAZ 52473), light brown (DHMECN 5236) or greenish cream (ZSFQ 1305) (Fig. 2); hidden surfaces of thighs brown splashed with dirty cream; groin with irregular yellowish marks; venter light gray (QCAZ 52474) or cream (QCAZ 52473) spotted to densely spotted (QCAZ 52474, DHMECN 5236, QCAZ 5212) with brown. Golden coppery iris with black reticulations and a reddish horizontal stripe (Fig. 2).

Distribution and natural history.

Pristimantis mallii is only known from Fundación EcoMinga’s Río Zuñag Ecological Reserve, which is located in the southeastern buffer zone of the Llanganates National Park, in Baños, Río Negro, Tungurahua province, in the upper basin of the Pastaza River, on the east-central slope of the Andes in Ecuador. This locality comprises montane cloud forest ( MAE 2012). The elevation range is 1300-2190 m above sea level.

All specimens were found on herbaceous and shrub vegetation inside mature forest, where they perched on herbs, shrubs, palms, ferns, bromeliads and Araceae between 100 and 450 cm above the ground. A couple in amplexus was found in January 2012, and the female deposited an egg clutch in a field bag, in the time passed between being collected in the field and reaching the base camp. Additionally, two couples in amplexus and an adult female were found in October 2017.

Etymology.

The new species is named in honor of the late Dr V. N. Mallikarjuna “Malli” Rao, of Wilmington, Delaware, USA. A winner of the Lavosier Medal at DuPont, he helped develop an environmentally safe alternative to the fluorocarbons that were depleting the ozone layer. His donation to EcoMinga in 2007 started the Río Zuñag Reserve, the type locality of P. mallii .