Proceratophrys kaingang, Santana & Mângia & da Silva Alves Saccol & Gomes dos Santos, 2021

Santana, Diego Jose, Mangia, Sarah, da Silva Alves Saccol, Suelen & Gomes dos Santos, Tiago, 2021, A new species of Proceratophrys Miranda-Ribeiro, 1920 (Anura, Odontophrynidae) of the P. bigibbosa species group from Southern Brazil, Vertebrate Zoology 71, pp. 387-401 : 387

Proceratophrys kaingang sp. nov.

Figs 1 , 2 , 3

Holotype.

ZUFSM 11127, adult female, collected at the Reserva Particular do Patrimônio Natural Rancho Sonho Meu (RPPN; Private Reserve of Natural Heritage), Tibagi municipality, in the Guartelá Canyon region from Campos Gerais of Paraná state, South Brazil (-24,559588, -50,275243), on 24 February 2016 by T. G. Santos, S. S. A. Saccol and A. A. B. Portela.

Paratypes.

ZUFSM 11123, ZUFSM 11126, ZUFSM 11131, ZUFMS-AMP14527-14530 (all adult males), ZUFSM 11132 and ZUFMS-AMP14526 (adult females), collected with the holotype. ZUFSM 11079, ZUFSM 11080, ZUFSM 11081, and ZUFSM 11082 (adult males), collected at a private farmland (-24.559588, -50.275243) on 22 February 2016, by the same collectors.

Diagnosis.

Proceratophrys kaingang sp. nov. is diagnosed by the following combination of characters: (1) small size for P. bigibbosa group (SVL 22.97-27.10 mm in adult males, 33.46-39.36 mm in adult females); (2) snout rounded in ventral and dorsal views, obtuse in profile; (3) upper eyelid border with small, rounded tubercles of similar size, and fused; (4) small postocular swellings; (5) yellowish blotches on the venter (in life); (6) toe webbing poorly developed; (7) distinct tympanic membrane, bordered by rounded tubercles.

Comparison with other Species.

Proceratophrys kaingang sp. nov. readily differs from P. appendiculata , P. belzebul , P. boiei , P. gladius , P. itamari , P. izecksohni , P. laticeps , P. mantiqueira , P. melanopogon , P. moehringi , P. paviotii , P. phyllostomus , P. pombali , P. renalis , P. rondonae , P. sanctaritae , P. subguttata , and P. tupinamba by the absence of a single unicuspidate palpebral appendage (a single and long unicuspidate palpebral appendage in all species, except in P. rondonae , which has a single and short multi-cuspidate palpebral appendage). In addition, P. kaingang sp. nov. can be distinguished from P. appendiculata , P. belzebul , P. gladius , P. itamari , P. izecksohni , P. laticeps , P. mantiqueira , P. melanopogon , P. moehringi , P. phyllostomus , P. pombali , P. sanctaritae , P. subguttata , and P. tupinamba by lacking a rostral appendage (present in those species). Proceratophrys kaingang sp. nov. differs from P. ararype , P. bagnoi , P. branti , P. carranca , P. concavitympanum , P. cristiceps , P. cururu , P. dibernardoi , P. goyana , P. huntingtoni , P. minuta , P. moratoi , P. redacta , P. rotundipalpebra , P. salvatori , P. schirchi , P. strussmannae and P. vielliardi by the presence of post-ocular swellings (absent in these species).

Among the species from P. bigibbosa group, P. kaingang sp. nov. differs by (1) its smaller size (mostly in males): 22.97-27.10 mm in males, and 33.46-39.36 mm in females ( P. brauni : 30.0-34.6 mm in males and 38.9-39.8 mm in females; P. bigibbosa : 35.5-43.8 mm in males and 51.2-53.4 mm in females; Kwet and Faivovich, 2001; and P. palustris : 27.3-33.8 mm in males, Giaretta and Sazima, 1991), except from P. avelinoi that presents similar sizes (23.9-29.2 males and 30.2-36.5 in females); (2) snout rounded in dorsal view ( P. brauni : pointed tip of the snout); (3) upper eyelid border with small, rounded tubercles of similar size, and fused ( P. avelinoi : small and triangular tubercles of varying sizes, and fused; P. bigibbosa : enlarged and pointed tubercles of varying sizes, not fused; P. brauni : long and triangular pointed tubercles of varying sizes, not fused); (4) small postocular swellings ( P. bigibbosa , P. brauni and P. palustris : presence of two well-developed, bulbous, bony post-ocular swellings); (5) toe webbing poorly developed ( P. bigibbosa : well-developed toe webbing); (6) yellowish blotches on the venter ( P. avelinoi and P. brauni : venter with orange reddish blotches; P. bigibbosa : venter red irregularly spotted with black; P. palustris : venter dark-grey with small beige blotches); and (7) distinct tympanic membrane, bordered by rounded tubercles ( P. avelinoi : tympanic membrane indistinct, covered with minute homogeneous tubercles).

Description of the Holotype.

Head wider than long (HL/HW = 0.70), head length 32% of SVL, snout rounded in dorsal and ventral views, obtuse in profile; nares elliptical and prominent, canthal crests well marked, prominent, and covered by small tubercles; no preocular crests; eyes directed anterolaterally, eye diameter 38% of head length; eyelid with distinct, rounded tubercles, with the contact point between the ocular-dorsal ridge of warts and the external eyelid margin tubercles in a tubercle posterior to the post-ocular swellings, six warts on the border of the left eyelid and five on the right; sparse tubercles on the eyelid; distinct tympanum; vomerine teeth in two short rows between and below the choanae; frontoparietal crests well developed; region between frontoparietal crests shallow; interocular ridge of warts not organized in a row, with sparse small rounded tubercles; ocular-dorsal ridge of warts incomplete, and discontinued to the coccyx region. Dorsal surface, including flanks, arms and legs, with various warts of different sizes and shapes, a single row of tubercles in different sizes bordered with some sparse tubercles on the forearm; ventral surfaces, except hands and feet and cloacal region, covered by numerous small, rounded, uniform warts. Finger lengths IV > II > I > III (Fig. 2b ); interdigital webbing absent; inner metacarpal tubercle rounded; single outer metacarpal small, both internal and external are rounded; scarce small, rounded supernumerary tubercles; subarticular tubercles large, rounded, but grooved anteriorly and posteriorly. Toe lengths I > II > V > III > IV; inner metatarsal tubercle long, elliptical, poorly spatulated; outer metatarsal tubercle small, rounded; scarce small, rounded supernumerary tubercles; subarticular tubercles large, nearly rounded, grooved anteriorly and posteriorly.

Color Pattern in Life (Fig. 3).

Dorsal coloration overall in variable shades of brown, with regular patterns of dark brown blotches in the dorsum. Presence of longitudinal irregular stripes of light brown in dorsolateral region. Gular region cream colored with mottling dark brown. Belly dark brown to black, irregularly spotted with yellow. Ventral surface of limbs dark brown to black, spotted with yellowish marks. Palm, fingers, soles of foot and toes are black, with two to three transverse dark-brown bars on fingers and toes.

Color Pattern in Preservative.

Overall coloration about the same as in life. However, the color became faded, and the light tones became darker. The longitudinal irregular stripes are brown in dorsal-lateral region. Gular region color beige with mottling dark brown. Belly dark brown irregularly spotted with beige.

Variation.

The main variation within this species relies on the sexual size dimorphism, with females (Fig. 4 ; 33.46-39.36 mm) bigger than males (Fig. 5 ; 22.97-27.10 mm); in addition, males have a darker gular region (Fig. 5 ). Overall, the tubercles on the dorsum can vary in size, and some can be rounded to triangular. The variation of the dorsal coloration is more prominent in the dark brown blotches that border the dorsal row of tubercles. The ventral pattern varying slightly on shape and size of yellowish blotches (Fig. 4 and 5 ).

Phylogenetic Inferences and Mitochondrial DNA Divergences.

Our 16S tree (Fig. 5 ) confidently recovered P. kaingang sp. nov. nested within the P. bigibbosa species group, and as the sister taxon of P. brauni (pp > 0.95). All nodes for species in the P. bigibbosa species group are well supported (pp > 0.95); however, some deeper nodes within Proceratophrys had low posterior probabilities, probably due to the single based gene tree. Average sequence divergence between the new species and its congeners within the P. bigibbosa species group ranges from 2.1% ( P. brauni ) to 5.6% ( P. bigibbosa ) (Appendix II).

Distribution and Natural History.

Proceratophrys kaingang sp. nov. is known only from its type locality, the Guartelá Canyon region, Tibagi municipality, in the Campos Gerais of Paraná state, Brazil (Fig. 7 ). Grassland physiognomies (e.g., rocky vegetational refuge, hygrophilous steppe, and grassy-woody steppe) are predominant in this region (Fig. 8 ), consisting of relictual vegetation that include Mixed Ombrophilous Forest and Cerrado mosaics ( Carmo et al. 2012; Souza et al. 2018). Calling males and a female were found in temporary puddles and slow running waters associated to flooded grasslands in the Private Reserve of Natural Heritage (RPPN Rancho Sonho Meu) and in a wetland in agricultural landscape. Calling activity was recorded in both diurnal and nocturnal periods (from early afternoon until at least ~11:00 h pm) during a historic event of heavy rains in the Paraná state. Males called from moist soil, exposed at the muddy edges of puddles as well as partially submerged in shallow flowing water, hidden among hygrophilous vegetation. Males of at least 12 other species were calling in the same breeding sites used by Proceratophrys kaingang sp. nov. (i.e., Aplastodiscus perviridis , Boana albpunctata , B. prasina , Dendropsophus minutus , Leptodactylus furnarius , L. fuscus , L. plaumanni , Melanoprhyniscus vilavelhensis , Physalaemus aff. gracilis , Scinax fuscovarius , S. rossaferesae , and S. squalirostris ).

Etymology.

The specific epithet Proceratophrys kaingang is a noun in apposition referring to the Kaingang (or Caingangue) ethnic group, which inhabits the plateau regions of the states of Paraná, São Paulo, Rio Grande do Sul and Santa Catarina, Brazil. We suggest the following Portuguese vernacular names “sapo-de-chifre-dos-caingangue” or “sapo-de-chifre-do-guartelá”.

Figures

Figure 1. Holotype of Proceratophrys kaingang sp. nov. (ZUFSM 11127). (A) Dorsal view of the body; and (B) ventral view of the body. Scale bar = 10 mm.

Figure 2. Holotype of Proceratophrys kaingang sp. nov. (ZUFSM 11127). (A) Head lateral view; (B) ventral view of right hand; and (C) ventral view of right foot. Scale bar = 5 mm

Figure 3. Live specimens of Proceratophrys kaingang sp. nov. from Parque Estadual do Guartela (type locality), Tibagi municipality, Para state, Brazil. (A) a live male (ZUFSM 11127); (B) the ventral coloration in life of an adult male (ZUFSM 11127); and (C) a couple in amplexus.

Figure 4. Dorsal and ventral color variation in preservative among female specimens from type series. (A) dorsal and (B) ventral views of ZUFSM 11027 (holotype); (C) dorsal and (D) ventral views of ZUFSM 11132; (E) dorsal and (F) ventral views of ZUFMS-AMP 14526. Scale bar = 10 mm

Figure 5. Dorsal and ventral color variation in preservative among male specimens from type series: (A) dorsal and (B) ventral views of ZUFSM 11079; (C) dorsal and (D) ventral views ZUFSM 11080; (E) dorsal and (F) ventral views of ZUFSM 11131; (G) dorsal and (H) ventral views of ZUFMS-AMP 14528; (I) dorsal and (J) ventral views of ZUFMS-AMP 14529; (K) dorsal and (L) ventral views of ZUFMS-AMP 14530.

Figure 7. Geographic distribution of species from Proceratophrys bigibbosa species group in South America.

Figure 8. Habitat in which Proceratophrys kaingang sp. nov. was found. (A) General view of the Private Reserve of Natural Heritage (RPPN Rancho Sonho Meu), Tibagi municipality, in the Guartela Canyon region from Campos Gerais of Parana state, Brazil. (B) The predominant grassland physiognomies in the region, and (C) the rocky vegetational refuge, hygrophilous steppe and grassy-woody steppe where the new species occurs.