Rhinella lilyrodriguezae, Cusi, Juan C., Moravec, Jiri, Lehr, Edgar & Gvozdik, Vaclav, 2017

Cusi, Juan C., Moravec, Jiri, Lehr, Edgar & Gvozdik, Vaclav, 2017, A new species of semiarboreal toad of the Rhinellafestae group (Anura, Bufonidae) from the Cordillera Azul National Park, Peru, ZooKeys 673, pp. 21-47 : 21-26

Rhinella lilyrodriguezae sp. n.

Rhinella cf. festae : Tasker and Twomey 2015: Pag. 1, figs 7-9.

Suggested English name.

Lily Rodriguez’s Beaked Toad

Suggested Spanish name.

Sapo picudo de Lily Rodríguez

Holotype.

MUSM 32204 (field number KT 75, Figs 3-6). An adult gravid female collected ca. 20 km from Park Rangers Center N° 53 “Shapaja” of the Cordillera Azul National Park (08°11'15.1"S, 76°12'36.8"W, 1260 m), Alto Biavo District, Bellavista Province, San Martín Region, Peru, collected on 27 September 2013 by K. Tasker and J. C. Cusi.

Paratypes.

Five individuals (Figs 7-8): an adult female MUSM 32206 (field number KT 76), two juveniles MUSM 32211 (field number KT 84), 32213 (field number KT 72) collected with the holotype; two adult females MUSM 32201 (field number KT 89), MUSM 32205 (field number KT 87), same locality as holotype, collected on 28 September 2013 by K. Tasker and J. C. Cusi.

Diagnosis.

A large species of the Rhinella festae group confirmed by 16S DNA barcoding. The new species can be diagnosed by the following combination of characters: (1) large size, SVL 47.1-58.3 mm in females (n = 4), males are unknown; (2) eight presacral vertebrae; (3) sacral vertebrae fused with coccyx; (4) snout long, acuminate, pointed to rounded terminally in dorsal view; snout protuberant, directed slightly anteroventral in profile as a "shark snout"; (5) cranial crests moderately developed; (6) canthal, supraorbital, postorbital and supratympanic crests continuous, distinctly elevated in female, slightly elevated in juveniles; pretympanic crest present, occipital crest absent; (7) tympanic membrane present, tympanic annulus weakly defined; (8) mandibular angle not protruding; (9) parotoid glands moderately large, roughly triangular to rounded in outline, slightly swollen laterally, incorporated into lateral row of tubercles; (10) dorsolateral rows of small, conical tubercles extending from parotoid gland to groin; (11) hands and feet with long digits, fingers basally webbed and toes moderately webbed; (12) skin on dorsum smooth with scattered conical tubercles in females; (13) subarticular tubercles diffuse, round to ovoid; (14) supernumerary tubercles present, round but poorly developed; (15) cloacal sheath absent; (16) in life, dorsum light brown to greenish brown with irregular brown, dark brown or black markings; with or without grey white middorsal stripe; venter cream yellow to brownish grey with minute light cream spots; iris silvery greenish with irregular black mottling.

Comparisons with other species.

The new species shares similarities with members of the Rhinella festae and R. acrolopha groups. Three species of the R. acrolopha group from Colombia and Panama ( R. truebae , R. lindae and R. tenrec ) are large-sized toads similarly to R. lilyrodriguezae sp. n. (maximum SVL 65.9 mm in R. truebae , 62.2 mm in R. lindae , 60.8 mm in R. tenrec and 58.3 mm in R. lilyrodriguezae ). The new species is most similar morphologically to R. truebae and R. tenrec , but is distinguished by lacking the occipital crest (which is low in R. truebae and R. tenrec ). Rhinella lilyrodriguezae sp. n. can also be distinguished from R. tenrec by having a sacrum fused with the coccyx, cranial crests moderately developed and tympanic membrane present (sacrum not fused with the coccyx, cranial crests poorly developed, tympanic membrane absent in R. tenrec ); from R. truebae and R. lindae by having a dorsolateral row of small conical tubercles extending from parotoid gland to groin, hands basally webbed, and feet moderately webbed (dorsolateral fold formed by the tubercles fusion, hands and feet extensively webbed in R. truebae and R. lindae ). Rhinella lindae possesses a snout slightly directed upwards (snout directed slightly anteroventral as "shark snout" in R. lilyrodriguezae ).

Other species of the Rhinella acrolopha group are R. acrolopha , R. nicefori , R. paraguas , and R. ruizi . These species are differentiated from R. lilyrodriguezae sp. n. (characters in parentheses) by the absence of the tympanic membrane (present), hands and feet extensively webbed in R. paraguas and R. ruizi , and reduced webbing in R. acrolopha and R. nicefori (hands basally webbed and feet moderately webbed), sacrum not fused with the coccyx except in R. paraguas (fused), seven presacral vertebrae except in R. paraguas (eight) and the presence of occipital crest except in R. paraguas (absent). Rhinella nicefori and R. ruizi have hands and feet with short digits (long digits). Rhinella paraguas and R. ruizi have cranial crests very low (moderately developed). Rhinella nicefori has a dorsolateral row of enlarged tubercles extending from the posterior margin of the parotoid gland to a point about three-fourths the distance between the axilla and groin (conical tubercles extending to groin). Rhinella acrolopha possesses a snout directed markedly anteroventrally (directed slightly anteroventral as "shark snout") and a dorsolateral row of depressed tubercles extending from the posterior margin of the parotoid gland posteriorly to a point about two-thirds distance between axilla and groin (conical tubercles extending from parotoid gland to groin).

The remaining species of the R. acrolopha and R. festae groups from Colombia and Ecuador ( R. festae , R. macrorhina and R. rostrata ) are distinguished from R. lilyrodriguezae sp. n. by lacking the tympanic membrane (present), by having hands and feet extensively webbed (hands basally webbed and feet moderately webbed), seven presacral vertebrae (except for R. macrorhina which has eight), snout directed markedly anteroventrally in R. festae and R. macrorhina and straight in R. rostrata (directed slightly anteroventral). Rhinella festae and R. macrorhina have occipital crest well developed (absent). Rhinella macrorhina is distinct in having a sacrum not fused with the coccyx (fused), and a dorsolateral row of small tubercles extending from posterior margin of parotoid gland posteriorly to a point about one-half distance between axilla and groin (tubercles extending from parotoid gland to groin). Rhinella festae possesses dorsolateral row of slightly enlarged, conical tubercles extending from posterior margin of skull to a point about three-fourths distance between axilla and groin (tubercles extending from parotoid gland to groin) and hands with short digits (long digits).

Peruvian species of the R. festae group ( R. chavin , R. manu , R. nesiotes and R. yanachaga ) are distinguished from R. lilyrodriguezae sp. n. by having smaller females (maximum SVL 21.4 mm in R. manu , 23.6 mm in R. nesiotes , 45.7 mm in R. yanachaga , except in R. chavin with 54.8 mm; vs. 58.3 mm in R. lilyrodriguezae ) and by having webbing of hands and feet fleshy (membranous in R. lilyrodriguezae ). Rhinella chavin possesses a snout rounded in lateral view (snout protuberant, directed slightly anteroventral as "shark snout"), large [about twice ED], ovoid parotoid glands (moderately large [about same size as ED] and triangular), dorsolateral row of large, nearly round elevated tubercles beginning above insertion of forelimb extending to inguinal region (small, conical tubercles extending from parotoid gland to groin), elevated, elongate glands on forearm, tibia and outer dorsal margin of the foot and hand (glands absent) and hands and feet with relatively short digits (long digits). Rhinella nesiotes has a snout rounded in lateral view (snout protuberant, directed slightly anteroventral as "shark snout"), lacks cranial crests (moderately developed), low ovoid parotoid glands (moderately large and triangular), lacks dorsolateral row of tubercles (present) and hands and feet with relatively short digits (long digits). Rhinella manu has a snout pointed in lateral view (snout protuberant, directed slightly anteroventral as "shark snout"), large [about twice ED], oblong parotoid glands to the point of being nearly spherical (moderately large [about same size as ED] and triangular) and inconspicuous cranial crests (moderately developed). Rhinella yanachaga is most similar to R. lilyrodriguezae sp. n. from cloud forests of central Peru, both have cranial crests, tympanic membrane distinct, moderately large [about same size as ED] parotoid glands, dorsolateral rows of small, conical tubercles extending from parotoid gland to groin, fingers and toes relatively long and long, slender extremities. Nevertheless, R. yanachaga differs from R. lilyrodriguezae sp. n. by having a snout slightly protruding in lateral view (snout protuberant, directed slightly anteroventral as "shark snout"), well developed webbing on hands and feet (hands basally webbed, and feet moderately webbed), dorsal skin smooth without keratin-tipped tubercles in females (dorsum smooth with scattered conical tubercles in females) and dorsal coloration in life is dark brown with small, irregular, green spots and markings (dorsum light brown to greenish brown with irregular brown, dark brown or black markings).

Description of the holotype.

Gravid female; body robust; SVL 58.3 mm; head triangular in dorsal view; head wider than long (HW 1.12 times HL), head width 30% of SVL; head length 27% of SVL; head narrower than body; snout acuminate, rounded terminally in dorsal view; not bulbous at tip; distance from the nostril to the tip of the snout (3.2 mm) is noticeably less than the distance from the nostril to the eye (5.8 mm), constituting 20% of head length; snout long, protuberant, directed slightly anteroventral as "shark snout" in profile (Fig. 3); snout with a ventral keel; canthus rostralis angular, rounded in lateral view; loreal region concave; nostrils small, rounded, not protruding, directed laterally, beyond anterior margin of lower jaw; internarial area concave; eye diameter equal to half the interorbital distance (ED/IOD = 0.5), ED noticeably shorter than E-D; canthal ridges angular evident; cephalic crests moderately developed; pre-, supra-, postorbital crests distinct, continuous; occipital crest absent; supratympanic crest evident, expanded laterally; pretympanic crests well defined; tympanic annulus weakly defined, superficial tympanic membrane present, not in contact with parotoid glands or postorbital crests; tympanum diameter smaller than eye diameter; parotoid glands moderately large (about same size as ED), roughly triangular in outline; upper eyelid covered with many, low, keratin-tipped tubercles; dorsal and lateral surfaces of head bearing many warts; forearms long, slender; forearm length 26% of SVL; dorsal surface of forelimbs spiculate, bearing densely scattered subconical tubercles; hand length 28% of SVL; hands with long fingers; relative lengths of fingers I <II <IV <III; finger tips rounded; fingers basally webbed, extended between fingers II–IV; Finger IV bears well-defined lateral fringes (Fig. 4A); palmar tubercle prominent, round, larger than oval thenar tubercle, about one half size of the palmar tubercle; subarticular tubercles diffuse, low, round to ovoid; supernumerary tubercles low, poorly developed, indistinct; hindlimbs long, slim; tibia length 40% of SVL; tibia longer than foot; dorsal surface of hindlimbs spiculate with subconical tubercles; foot length 39% of SVL; toes long; relative lengths of toes I <II <III <V <IV; toes tips rounded; toes moderately webbed, with the following formula: I 1-2- II 1 -– 2 III 1-3+ IV 3+-2 V; free portions of all toes bear well-defined lateral fringes (Fig. 4B); tarsal fold absent; inner metatarsal tubercle large, slightly elliptical, weakly protuberant; outer metatarsal tubercle round, smaller than inner metatarsal tubercle, half the size of inner metatarsal tubercle; subarticular tubercles diffuse, low, round to ovoid; supernumerary tubercles indistinct, skin on dorsal surface of the body with numerous small, round, elevated tubercles, bearing single keratinized tip; flanks with lower density of tubercles than dorsum; dorsolateral row of small, conical tubercles extending from parotoid gland to groin, not forming a distinct dorsolateral fold; skin of venter and throat, chest and venter granular; cloacal opening directed posteriorly at the mid-level of the thighs); tongue narrow, about 2.5 times as long as wide, not notched posteriorly, posterior one half free; choanae small, ovoid, widely separated and partially concealed by palatal shelf of maxilla.

Measurements (in mm) of the holotype.

SVL: 58.3; HW: 17.8; HL: 15.9; ED: 4.4; IOD: 9.3; EW: 3.9; EL: 6.4; IND: 4.3; E–N: 5.8; NSD: 3.2; SL: 9.1; FL: 15.4; HNDL: 16.3; FEML: 23.7; TL: 23.4; FOOTL: 22.6.

Coloration of the holotype in alcohol

(Fig. 5). General dorsal coloration light brown; dorsum and hind limbs with small, irregular, dark brown spots and markings; flanks cream, with a discontinuous, well defined, broad, black ventrolateral band beginning behind tympanum and extending to inguinal region; dark grey transverse bars on shanks; tympanum brown; upper lip cream without bars or spots; dorsolateral row of tubercles reddish brown, sharply contrasting with the black discontinuous ventrolateral band; throat, chest and venter gray with minute light cream spots; ventral surfaces of thighs gray with minute dark grey spots; ventral surfaces of hands and feet dark gray; subarticular and supernumerary tubercles cream on hands.

Coloration of holotype in life

(Fig. 6). Diurnal coloration of dorsum and flanks dark brown; flanks with irregular, dark green dorsolateral blotches extending from behind the parotoid glands to sacral region; dorsolateral row of tubercles light brown; throat dark gray; chest and belly grey with minute light cream spots extending to the thighs; lower side of the belly cream yellow; iris silvery greenish with irregular black mottling.

Variation

(Figs 7-9). Considerable variation between nocturnal and diurnal coloration was observed. The nocturnal coloration of an adult female (MUSM 32201, Fig. 7 A, B) was described as follows: dorsum light brown dorsally; whitish grey middorsal stripe extending from tip of the snout to cloaca; light cream parotoid glands; broad, whitish grey dorsolateral stripe on each side of flanks extending from behind the eyelids to groin; flanks with continuous, broad, black ventrolateral band extending from tympanic posterior region to groin; tip of the snout, eyes, eyelids, crown of head and tympanum light brown; upper lip, angle of the jaws and ventrolateral region of the flanks is whitish grey; dorsolateral row of tubercles reddish brown just above the black ventrolateral band; forelimbs and hands are brown, dorsum of the hands with whitish grey blotches in direction to the fingers; hindlimbs brown with transversal dark brown bars. The diurnal coloration of the same specimen (Fig. 7C, D): dorsum and flanks dark brown; gray middorsal stripe from tip of the snout to cloaca; parotoid glands cream brown; venter brownish grey marbled with cream; throat darker brown. An adult female (MUSM 32205; Fig. 8) lacked a middorsal stripe and had flanks with irregular dark grey spots at night. During the day, the same specimen was characterized by a dark green dorsolateral stripe on each side of body extending from parotoid gland to groin; venter brownish grey marbled with cream; irregular dark grey spots on thighs, lateral and lower side of the belly; white blotches on middle area of the belly. Two females (MUSM 32206, 32201) were of similar coloration as MUSM 32205.

The remaining paratypes show some variation in nocturnal color pattern. The overall dorsal coloration of the juveniles (MUSM 32211, 32213) is light grey with darker irregular markings forming a "dead-leaf pattern" from between eyes to cloacal region. Cranial crests are more prominent in adults than in juveniles. All specimens, except the holotype, have tip of snout acuminate in dorsal view. Radiographs show that all specimens have eight presacral vertebrae and a sacrococcygeal articulation (Fig. 9). For variation in measurements see Table 2.

Etymology.

The specific epithet lilyrodriguezae is a noun in the genitive case and a patronym for Dr. Lily Rodriguez, for her contributions to the knowledge of the Peruvian amphibians and her initiatives that have promoted the creation of numerous natural protected areas in Peru, such as the Cordillera Azul National Park.

Distribution, ecology and conservation status.

Rhinella lilyrodriguezae sp. n. is only known from the Shapaja sector within the CAZNP in northern Peru, at elevations between 1245 and 1280 m a.s.l. (Fig. 1). It was encountered in montane forest during the dry season. The type locality was accessed by a hike of 7 hours across a small trail from the Misterioso River (natural boundary between natural protected area and a forest concession dedicated to wood extraction). The species has nocturnal and semiarboreal mode of life (all individuals were found at night between 20:33 and 22:49 on leaves of bushes between 10 and 100 cm above the ground, along a small creek). A negative impact of logging, soil removal and noise pollution was observed around the localities of the new species. Subsistence game hunting and exaggerated fishing by local population are other factors threatening the local biodiversity.

One gravid female (MUSM 32204, SVL 58.3 mm, holotype) contained 185 ovarian eggs (left ovary: 95; right ovary: 90) with an average diameter of 2.8 ± 0.2 mm (3.2-2.7 mm, n = 20), which are pale cream yellow in preservative. The presence of numerous, large, pigmented eggs and association of the individuals with water bodies suggest endotrophic larvae (e.g., direct development or nonfeeding tadpoles that develop in water or moist soil; Peloso et al. 2014), as can be expected in R. manu , R. paraguas and R. tacana ( Grant and Bolívar-G. 2014, Catenazzi pers. comm.) The call of Rhinella lilyrodriguezae was not recorded. Other anuran species that occur in the area of the type locality include Hyloscirtus cf. phyllognathus , Pristimantis peruvianus , P. ventrimarmoratus , Rulyrana cf. flavopunctata , and Osteocephalus mimeticus (juveniles). We classify Rhinella lilyrodriguezae sp. n. as "Data Deficient" according to the IUCN red list criteria ( IUCN 2016) based on the limited information on its geographic range.