Herpetology Notes, volume 11: 357-360 (2018) (published online on 02 May 2018)

Range expansion in the geographic distribution of Ninia teresitae (Serpentes: Dipsadidae): new localities from northwestern Ecuador

Teddy Angarita-Sierra1,2,*

The semifossorial of the Ninia are the entire range of the . During examination of widely distributed throughout Central and South Ninia specimens housed at the Museum of Zoology, America, ranging from southern Mexico to northern Pontificia Universidad Católica del Ecuador ��������� Peru at elevations between sea level and 1600 m (Dunn, discovered 14 specimens of N. teresitae from eight new 1935; Savage, 2002; Angarita-Sierra, 2014). Ninia localities (Table 1), all restricted to Pacific lowlands currently comprises twelve species, of which only four and the western slopes of the Ecuadorian Andes (Fig. occur in South America (N. atrata Hallowell, 1845; N. 1). In the following account, I followed the methods franciscoi Angarita-Sierra, 2014; N. hudsoni Parker, and terminology of Peters (1964) and Dowling (1951). 1940; N. teresitae Angarita-Sierra and Lynch, 2017). Hemipenial preparation followed Pesantes (1994) with Even though the identities of some populations of the named forms of N. atrata have been resolved (see Savage and Lahanas, 1991; McCranie and Wilson, 1995; Savage, 2002; Angarita-Sierra and Lynch, 2017), their sympatric distribution and morphological overlap between N. atrata and its congeners have masked the taxonomic identity of several undescribed species. Ninia teresitae was recently described based on ten specimens from several localities of the Chocó- Magdalena biogeographic province in Colombia. This taxon was previously confused with N. maculata and N. atrata by Castaño-M et al. (2004) due to its irregular spotted patterns on the ventral body surfaces. However, this species can be distinguished from all congeners by using morphometric ratios of the head´s scales, colour pattern of the ventral surfaces of head and body, hemipenial morphology, and nasal scale count (Angarita-Sierra and Lynch, 2017). As part of an ongoing study of the Ninia atrata complex, I have examined more than 400 specimens that cover

1Yoluka ONG, Fundación de Investigación en Biodiversidad y Conservación Carrera 24 No. 51-81, Piso 3, Bogotá, Colombia; Figure 1. Distribution map of Ninia teresitae. Purple hexagons 2Grupo de Investigación de Biología de Organismos Tropicales, correspond to the new localities reported in this study, black Universidad Nacional de Colombia, Bogotá, Colombia. triangles depict literature records, and the red triangle denotes * Autor e-mail: [email protected] the type locality. 358 Teddy Angarita-Sierra

Table 1. List of Ecuadorian localities of Ninia teresitae. Localities in which Ninia teresita is sympatric with Ninia atrata are marked with an asterisk (*). 1 Table 1. List of Ecuadorian localities of Ninia teresitae. Localities in which Ninia teresita is sympatric with Ninia atrata are marked with an asterisk (*). 2 Specimen number Province Locality Latitude Longitude Elevation (m) QCAZ 659 Cotopaxi San Francisco de las Pampas* 0.4237°S 78.9677°W 1245 QCAZ 1086 Cotopaxi San Francisco de las Pampas* 0,4333°S 78,9667°W 1605 QCAZ 1257 Cotopaxi El Corazón 1,1363°S 79,0745°W 1568 QCAZ 1258 Esmeraldas Quinindé, Estero las Peñas 0,3269°S 79,4648°W 214 QCAZ 1259 Pichincha Noroccidente, Hacienda Pachigal Unknown Unknown Unknown QCAZ 1260 Cotopaxi San Francisco de las Pampas 0,4333°S 78,9667°W 1605 QCAZ 1263 Cotopaxi San Francisco de las Pampas 0,4333°S 78,9667°W 1605 QCAZ 1585 Cotopaxi San Francisco de las Pampas* 0,4333°S 78,9667°W 1605 QCAZ5040 Esmeraldas km 17 en la vía San Lorenzo-Ibarra 1,1812°S 78,7528°W 55 QCAZ 7036 Manabi Vía El Carmen-Pita, km 7, Rancho RH* 0,2389°S 79,4451°W 250 QCAZ 7199 Manabi Quinta Los Helechos, km 36 vía Santo Domingo-Chone 0,2738°S 79,4587°W 248 QCAZ 7207 Manabi Vía El Carmen-Pita, km 7, Rancho RH* 0,2389°S 79,4451°W 250 QCAZ 7837 Cotopaxi San Francisco de las Pampas* 0,4333°S 78,9667°W 1605 QCAZ 11960 Imbabura Reserva Manduriacu 0,3103°S 78,8576°W 1239 3 4

adjustments as suggested by Myers and Cadle (2003), the morphological variation of the species and allow Zaher and Prudente (2003), and Smith and Ferrari- adjustment of the following characters: maximum male Castro (2008). The staining process followed Jadin and SVL = 382 mm (earlier 324 mm), minimum female Smith (2010) and Angarita-Sierra (2014). Tail length SVL = 115 mm (198 mm), minimum ventral scale (TL) and snout–vent length (SVL) measurements were count in males = 138 (143), and minimum ventral scale made to the nearest 0.5 mm using a type measure. count in females = 148 (150). All remaining pholidosis features as well as body size measurement fall within Results the ranges described in the original species description 1 of N. teresitae (Angarita-Sierra and Lynch, 2017). The specimens reported in this paper were identified Likewise, the hemipenial morphology observed on as Ninia teresitae using the following external two males (QCAZ 1585, 7207) agrees with the original morphological characters and hemipenial morphology description, exhibiting a bilobed and semicaliculate (Angarita-Sierra and Lynch, 2017): ventral surfaces organ, with a lateral projection ornamented by a large, of head and body spotted without regular pattern, basal, hooked spine that is larger than any other spine subcaudal surface homogenously black or dark brown; on the hemipenial body, and a centrifugal and bifurcate two nasal scales; bilobed hemipenis; sulcus spermaticus sulcus spermaticus. centrifugal with a bifurcation proximal to midpoint of the hemipenial body; lateral projection ornamented with Discussion a large basal hooked spine that is larger than any other spine on the hemipenial body (Fig. 2). Northwestern Ecuador lies at the intersection of two All specimens used to compile this report (six males, of the most diverse terrestrial ecoregions on the planet, eight females) had the same colour pattern variation on the Andes and the Chocó (Arteaga et al., 2016). This the ventral surfaces of head and body as observed in convergence has allowed the evolution of astonishing the type series of Ninia teresitae, having strong or faint species richness, with several diversification patterns (inmaculate in QCAZ 1257) irregular pigmented spots and lineages observed in relatively limited areas (e.g., or faint irregular pigmented spots on the ventral surface. Mindo or Bilsa Biological Station; Arteaga et al., 2016, Also, all specimens exhibited dorsal surfaces that were Arteaga et al., 2013). In addition, northwestern Ecuador uniformly dark brown–black; the nuchal collar present belongs to an historical interchange pathway for South or absent, when present it was W-shaped, or included and North American faunal elements that allowed great the parietal scales (Fig. 3). immigration and emigration of lineages (Cadle The Ecuadorian specimens expand our knowledge of and Greene, 1993), but in addition, greater rates of Range expansion in the geographic distribution of Ninia teresitae, Ecuador 359

speciation have been observed than in other regions (Pyron and Wiens, 2013). These peculiarities make the northwestern Ecuador a significant hotspot for reptilian tropical biodiversity and endemism, as well as part of a biogeographical unit clearly recognized as the Chocó-Magdalena biogeographic province (Cadle, 1985; Hernández-Camacho, 1992, Angarita-Sierra and Lynch, 2017). Given the biogeographic definition of this province (Hernández-Camacho et al., 1992), the presence of N. teresitae in Ecuadorian Pacific lowlands as well as on the western slopes of the Ecuadorian Andes was expected. Particularly, these new records found in northwestern Ecuador mark the southernmost geographic extent of the species, which coincides with the southern limit of the Chocó-Magdalena biogeographic province. The Ecuadorian records of N. teresitae come from both conserved and transformed habitats (ranging from Chocoan rainforest to crops and towns), showing the same high tolerance of the species towards transformed habitats observed in the population at the type locality and in several sisters taxa (e.g., N. atrata, N. diademata, Figure 2. Hemipenis of Ninia teritae from San Francisco de N. maculata). Northwestern Ecuadorian populations of las Pampas, Cotopaxi, Ecuador (QCAZ 1585), in sulcate (left) N. teresita occur in sympatry with N. atrata in three of and asulcate (right) views. Photos by Alexandre Pinheiro de the eight localities recorded, reaching elevations of 1605 Almeida. m (Table 1). Finally, a proper and careful revision of the N. atrata-like snakes from Ecuador is recommended, in order to understand and clarify the taxonomic composition and geographic distribution of the genus.

Acknowledgements. I want to thank Omar Torres for allowing access to the QCAZ collection. Also, I thank to Yerka Sagredo and Diego Paucar for their hospitality and collaboration during my stay in Ecuador, Jhon Infante Betancour for helping me with the construction of the map, Gustavo Pazmiño for providing me with photographs of some specimens of N. teresitae, and Alexandre Pinheiro de Almeida for help with the hemipenis pictures.

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Accepted by Hinrich Kaiser