Asian Herpetological Research 2012, 3(1): 38–45 DOI: 10.3724/SP.J.1245.2012.00038

A New Species of the Genus Elaphe (Squamata: Colubridae) from Zoige County, Sichuan, China

Song HUANG1, 2, 3*, Li DING4*, Frank T. BURBRINK5, 6, Jun YANG7, Jietang HUANG3, Chen LING8, Xin CHEN5, 6 and Yaping ZHANG2**

1 College of Life and Environment Sciences, Huangshan University, Huangshan 245041, Anhui, China 2 State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming 650223, Yunnan, China 3 Huangshan Institute of Ophiology, Huangshan 245000, Anhui, China 4 Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, Sichuan, China 5 Department of Biology, the College of Staten Island, the City University of New York, 2800 Victory Blvd, Staten Island, NY 10314, USA 6 Department of Biology, the Graduate Center, the City University of New York, 365 Fifth Avenue, New York, NY 10016- 4309, USA 7 School of Life Sciences, Sichuan University, Chengdu 610064, Sichuan, China 8 College of Forestry, Southwest Forestry University, Kunming 650224, Yunnan, China

Abstract Previous phylogenetic analyses of DNA sequence data reveal a likely new species of Elaphe Fitzinger from China, which forms the sister group of all extant Elaphe. One of the three original specimens is crushed and the other two are neonates, which precludes a morphological analysis. Three adult snakes (2 females, 1 male) collected from Jiangzha Hot Spring in Zoige County, Sichuan, China on 26 July 2010 now facilitate the species’ description. Mitochondrial DNA nucleotide sequences of cytochrome b (cyt b), NADH dehydrogenase subunit 4 (ND4), 12S ribosomal RNA (12S rRNA), and cytochrome oxidase subunit 1 (CO1) from the adults are identical to sequences from the crashed specimen. Interspecific uncorrected p-distances of partial CO1 gene of 26 sequences from all 11 species of Elaphe, including this new species, are large (8.5–15.2%). This new species is morphologically distinguished from all other species of Elaphe by several characters: “M”-shaped marking on the supraoculars and adjacent frontal; four parallel series of red-brown spots on the dorsum extending from the neck to the vent; four longitudinal stripes associated with the spots; each spot is usually composed of a whole (or incomplete) red-brown scale with puce borders; spots of each stripe are similar in shape, they occur at relatively regular intervals of one or two scales, and they are slightly red-brown in color; and it has three preoculars, whereas all other species have one or two. Now Elaphe contains 11 species and this discovery highlights the need for continued exploration of temperate regions. Keywords new species, mitochondrial DNA, DNA barcoding, Zoige ratsnake

1. Introduction molecular phylogenetics and increased taxon sampling. These changes are important not only for classification, Nowadays, taxonomic revisions of major groups of but also for comparative biology, which requires credible vertebrates are common due, in part, to the revolution in phylogenetic estimates and accurate assessments of extant taxa for hypothesis testing (Felsenstein, 1988; * Both authors contribute equally to this work. ** Corresponding author: Prof. Yaping ZHANG, from Kunming Institute Díaz-Uriarte and Garland, 1996, 1998). Advanced snakes of Zoology, Chinese Academy of Sciences, with his research focusing (Caenophidia), representing about 3 000 species, are the on molecular phylogeny and evolution of vertebrates. E-mail: [email protected] subject of several recent massive phylogenetic studies Received: 5 January 2012 Accepted: 27 Feburary 2012 that propose numerous changes in classification (Pyron No. 1 Song HUANG et al. New Species of the Genus Elaphe from Zoige, China 39 et al., 2011; Burbrink and Crother, 2011) as well as members from all genera of Old World ratsnakes and provide important tests of major hypotheses in evolution representatives from the New World strongly indicates (Burbrink and Pyron, 2010). Within the advanced snakes, that the three specimens represent the sister species to all the ratsnakes are one of the most-well-studied groups. other taxa in the genus Elaphe. This relationship suggests Recent major taxonomic changes occur at generic and an undescribed species of Elaphe but it is not described species levels (Utiger et al., 2002; Burbrink and Lawson, because the absence of adult specimens precluded a 2007; Pyron and Burbrink, 2009a). morphological description. Several studies demonstrate that the ratsnake genus On 26 July 2010, we captured three adult snakes (2 Elaphe (sensu lato) is likely paraphyletic, particularly females, 1 male) at Jiangzha Hot Spring, Zoige County, with respect to New World taxa that are more closely Sichuan, China. Based on molecular data from cyt b, related to kingsnakes, pinesnakes and their allies (e. g., ND4 and 12S rRNA, the new specimens were determined Lampropeltis, Pituophis, Rhinocheilus, Cemophora). to belong to Elaphe and to be conspecific with those The genus Elaphe, with the type species Coluber reported by Ling et al. (2010). sauromates Pallas 1811 from Europe, is now restricted to the following ten taxa found throughout Eurasia: 2. Materials and Methods E. anomala (Boulenger), E. bimaculata Schmidt, E. carinata (Günther), E. climacophora (Boie), E. davidi Three adult snakes (2 females, 1 male) were sampled at (Sauvage), E. dione (Pallas), E. quadrivirgata (Boie), the side of a stream (Figure 1) by Li DING within 40 m E. quatuorlineata (Lacépède), E. sauromates (Pallas) of the mouth of Jiangzha Hot Spring at 12:00 hr on 26 and E. schrenckii (Strauch) (Schulz, 1996; Helfenberger, July 2010. Several days later, two females laid 11 eggs, 2001; Utiger et al., 2002; Burbrink and Lawson, 2007). respectively (Figure 2) in field terrariums. The eggs were Phylogenetic support for the monophyly of Elaphe is smooth-shelled and adhesive. The holotype (Collection strong and placement of species within in this genus is number: HS201014) and paratypes (HS201015–16) were confirmed by both nuclear and mtDNA genes (Utiger et preserved and deposited in the Museum of Huangshan al., 2002; Burbrink and Lawson, 2007). Geographically, University (Voucher numbers: HUM20110001–3). the highest diversity is centered in East Asia, although the Tissues from the distal ends of ventral scales were range of the genus extends as in the far north to Russia sampled from all the new specimens. Partial mtDNA (E. dione, E. schrenckii), in the south to Vietnam (E. sequences were obtained by polymerase chain reaction carinata), east to the China Sea, Korean Peninsula and (PCR) and direct sequencing using the primers and Japan (E. anomala, E. bimaculata, E. carinata, E. davidi, methods described in Arévalo et al. (1994), Burbrink et al. E. climacophora, E. quadrivirgata), and west to Italy (E. (2000), and Huang et al. (2009) for cyt b (1 015 bp) and quatuorlineata). ND4 (696 bp), and Utiger et al. (2002) for 12S rRNA Molecular data provide a powerful means of (428 bp) and CO1 (513 bp). The new cyt b, ND4 and identifying cryptic diversity, such as that within Elaphe 12S sequences were identical to those reported by Ling (E. quatuorlineata) and the related genera Zamenis (Z. et al. (2010) (GenBank accession number: HQ330525, longissimus), Pantherophis (P. guttatus, P. obsoletus), and HQ330527 and HQ330523, respectively). The absence of Lampropeltis (L. getula), (Lenk and Joger, 1994; Lenk et al., variation indicated that the six snakes were conspecific. 2001; Burbrink et al., 2000; Burbrink, 2001, 2002; Pyron Because sequences from COI—DNA barcoding— and Burbrink, 2009b, c). The rapid radiation of these have enabled the discrimination of closely allied species advanced snakes may be responsible for the absence of (Hebert et al., 2003) and provided a standardized tool morphological differentiation. For example, both E. dione to inventory biodiversity (Naro-Maciel et al., 2010), and E. bimaculata are morphologically quite similar with we sequenced the gene for the three new specimens respect to scutellation and color pattern, and both occur in (GenBank accession numbers: JF510474–76). The China. uncorrected p-distances of partial CO1 gene of 26 Based on molecular phylogenetic analyses of sequences from all the 11 species of Elaphe (23 from nucleotide sequences from three mitochondrial DNA Utiger et al., 2002; 3 from this study) were calculated genes (cyt b, ND4 and 12S rRNA), Ling et al. (2010) using the program MAGA 4 (Tamura et al., 2007). COI suggest that three specimens collected from Zoige p-distances ranged between 8.2–15.0 among species and County, Sichuan, China likely represent a new species between 0–2.5 within species (Table 1). of Elaphe. The matrilineal genealogy, which contains All measurements and observations were conducted 40 Asian Herpetological Research Vol. 3 26 ] 0 * 7.6 25 0 8.4 0 24 8.8 9.2 0.8 23 0 8.8 0 11.9 11.7 22 12.5 12.3 # 0.0 0 11.9 11.7 21 12.5 12.3 8.2 8.2 0 12.1 11.5 20 11.5 10.9 8.2 8.2 0.0 0 12.1 11.5 19 11.5 10.9 8.6 8.6 0 . [26]; the species was previously was known as E. 10.3 10.3 12.9 11.9 18 11.9 11.1 vs 8.6 8.6 2.5 0 10.7 10.7 13.5 12.1 17 12.1 11.3 9.9 9.9 0 12.1 12.1 11.9 11.5 13.1 13.3 16 13.1 13.1 [21] suggests they are either conspecific or the former is a subspecies 11.5 11.5 10.9 10.9 11.7 12.3 10.3 0 12.3 12.3 15 12.3 11.3 2.3 0 11.9 11.9 10.5 10.5 11.3 11.9 11.1 12.3 12.7 14 13.1 11.7 . GenBank accession numbers follow species names. 1.4 1.8 0 11.3 11.3 10.3 10.3 10.3 10.9 10.3 11.7 12.5 13 12.5 11.1 1.6 1.9 0.2 0 11.5 11.5 10.5 10.5 10.5 11.1 10.5 11.9 12.7 12 12.7 11.3 (11.1%–14.4%). 2.1 1.9 1.2 1.4 0 11.5 11.5 10.7 10.7 11.3 11.9 10.9 11.9 12.9 11 12.9 10.7 Elaphe 2.1 1.9 1.2 1.4 0 0.0 11.5 11.5 10.7 10.7 11.3 11.9 10.9 11.9 12.9 10 12.9 10.7 8.6 9.4 8.6 8.8 9.6 9.6 0 13.1 13.1 11.5 11.5 12.5 12.3 10.7 13.5 13.1 9 12.9 14 9.4 9.4 9.0 9.2 9.6 9.6 1.4 0 12.9 12.9 10.9 10.9 12.3 12.5 11.9 13.5 12.9 8 13.1 13.8 9.4 9.7 9.0 9.2 9.6 9.6 1.8 0.4 0 12.5 12.5 11.3 11.3 12.7 12.9 12.1 13.1 12.5 7 13.1 13.8 12.5 12.5 11.9 11.9 11.9 12.1 11.9 11.1 11.1 10.5 10.7 11.3 11.3 12.7 13.1 13.5 0 12.7 14.6 6 14.6 12.1 0.6 0 12.5 12.5 11.9 11.9 11.9 12.1 11.3 11.3 11.3 10.7 10.9 11.5 11.5 13.1 13.5 13.8 12.9 15.0 5 15.0 12.5 E. sauromates [25] distance of 7.6% occurs between two individuals of E. sauromates A : * 0.6 0.0 0 : The distance of 0.0% between E. anomala [22] and schrenckii The distance : 12.5 12.5 11.9 11.9 11.9 12.1 11.3 11.3 11.3 # 10.7 10.9 11.5 11.5 13.1 13.5 13.8 12.9 15.0 4 15.0 12.5 , 2010). 12.9 12.9 11.1 11.1 14.2 13.3 12.9 12.7 13.1 12.5 12.7 12.5 12.5 13.1 13.1 13.1 13.1 13.1 13.6 13.6 0 13.1 13.5 3 13.5 et al. sp. nov. and all other ten species of E. zoigeensis sp. nov. 12.7 12.7 11.5 11.5 14.0 13.1 12.9 13.1 13.5 12.9 13.1 12.9 12.9 13.5 13.5 13.5 13.1 13.5 13.6 13.6 0.4 0 13.1 13.5 2 13.5 12.7 12.7 11.5 11.5 14.4 13.5 13.3 13.1 13.5 12.9 13.1 12.9 12.9 13.5 13.5 13.1 13.5 13.5 14.0 14.0 0.4 0 0.4 13.1 13.5 1 13.5 , 2002; Ling et al. AY122714 AY122712 AY122712 AY122687 AY122687 AY122688 AY122688 AY122686 AY122686 AY122684 AY122684 AY122753 AY122753 AY122683 AY122683 AY122720 AY122754 AY122754 AY122755 AY122755 AY122691 AY122691 AY122747 AY122747 AY122749 AY122749 AY122745 AY122745 AY122746 AY122746 AY122750 AY122750 AY122748 AY122748 AY122719 [22] E. anomala AY122719 [21] E. schrenckii [20] E. quadrivirgata AY122709 AY122710 AY122710 [19] E. quadrivirgata [18] E. carinata [17] E. carinata [16] E. davidi [15] E. dione [14] E. dione [13] E. dione [12] E. dione [11] E. dione [11] [10] E. dione [ 9] E. bimaculata [ 8] E. bimaculata [ 7] E. bimaculata AY122713 AY122713 [25] E. sauromates [ 6] E. climacophora [ 5] E. climacophora [ 4] E. climacophora [ 3] HUM20110002 JF510476 [ 3] HUM20110002 [ 1] HUM20110001 JF510474 [ 1] HUM20110001 JF510475 [ 2] HUM20110003 AY122711 [26] E. sauromates AY122711 [24] E. quatuorlineata [23] E. quatuorlineata [ . Their taxonomic status may need further study. Their taxonomic status may need further study. . quatuorlineata sauromates Interspecific distances (8.2%–15.0%) are not shadowed. Matrix of uncorrected P-distances of partial CO1 gene (513bp) for 26 sequences from 11 species of Elaphe 1 Matrix of uncorrected P-distances partial CO1 gene (513bp) for 26 sequences from 11 Table Conspecific distances (0.0%–2.5%) are shadowed in gray. of the latter (Pope, 1935; Utiger Numbers in grid show the distances between No. 1 Song HUANG et al. New Species of the Genus Elaphe from Zoige, China 41

Figure 1 Sampling site and type locality of Elaphe zoigeensis sp. nov. A: The type locality of Elaphe zoigeensis sp. nov. is indicated by an asterisk, located at Jiangzha Hot Spring, Zoige County, Sichuan, China (N34º12’18’’, E102º42’48’’, 3200 m asl); B: Habitat of type locality of the new species. Photo by S. Huang.

Figure 2 Clutch of eggs of HUM20110001 (holotype) and HUM20110002 (paratype). A: Eleven eggs laid by HUM20110002 on 31 July 2010, with egg length being 29.45 ± 1.35 mm, egg diameter 16.74 ± 0.99 mm, and mean egg weight 5.50 g; B: Eleven eggs laid by HUM20110001 on 29th July 2010, with egg length being 29.35 ± 1.80 mm, egg diameter 16.46 ± 0.78 mm, and mean egg weight 4.59 g. Photos by S. Huang. on live snakes and the description of eggs was taken at Holotype: HUM20110001, adult female captured at 90% humidity. Terminology, methods, and characters 12:00 on 26 July 2010. Specimen was basking on a large for the description of the new taxon mainly followed rock about 20 m from the mouth of Jiangzha Hot Spring Schulz (1996). Herein, the term subpreocular was used to (3200 m elevation, N34º12’18’’, E102º42’48’’) in Zoige describe the lowest small preocular. The measurements County, Sichuan, China. of length and weight were taken using a digital vernier Paratypes: Captured on gravel by a stream caliper and electronic balance to the nearest 0.1 mm and (HUM20110002, adult female, about 40 m from the 0.1 g, respectively, except for snout-vent and tail lengths, mouth of Jiangzha Hot Spring) and in a small soil hole by which were recorded with a measuring tape to the nearest the same stream (HUM20110003, adult male, about 30 m 1 mm. from the mouth of Jiangzha Hot Spring), same place and 3. Results time as the those of the holotype. Diagnosis: Elaphe zoigeensis sp. nov. is distinguished Elaphe zoigeensis sp. nov. (Figures 3–5) from all other species of Elaphe by several sets of Zoige Ratsnake, Ruoergai Jinshe (Bopomofo) characters involving color and scutellation. Color pattern 42 Asian Herpetological Research Vol. 3

Figure 3 Body of holotype (HUM20110001, adult female). A: Dorsolateral; B: Abdomen; C: Back. Photos by S. Huang.

Figure 4 Head of holotype (HUM20110001, adult female). A: Dorsal; B: Ventral; C: Right; D: Left. Po: Preocular; Sp: Supralabial; Sp*: Supralabial in contact with the eye; At: Anterior temporal; Pt: Posterior temporal. Photos by S. Huang. is distinct in having a red brown, “M” shaped marking on this species has 3 preoculars, rather than 1–2 found in all the back of the head at the margins of the supraoculars other species (Schulz, 1996). and adjacent posterior frontal and the presence of 4 Description of the holotype and variation: When longitudinal, parallel rows of spots on the dorsum differing from the holotype, features of the paratypes extending from the neck to vent. Each spot usually follow in parentheses (HUM20110002, HUM20110003 encompasses an entire scale and is red brown, while the respectively). Holotype an adult female with body weight edges of the spot and bordering scales are puce. The 97.96 g (103.68 g, 124.29 g), snout-vent length 722 mm spots from each of the parallel rows are similar in size (695 mm, 64 mm), tail length 158 mm (150 mm, 175 and shape, and rank at a relatively regular interval of one mm). Scutellation: two postoculars; 2 + 3 temporals or two scales of slightly red brown color. Additionally, (anterior temporal fused); 3 preoculars (HUM20110003 No. 1 Song HUANG et al. New Species of the Genus Elaphe from Zoige, China 43

Figure 5 Heads of the paratypes, showing the lateral scales on both sides. A, B: Right, left side of HUM20110002 (adult female); C, D: Right, left side of HUM20110003 (adult male). Abbreviations are same as those in Figure 4. Photos by S. Huang. left includes a subpreocular); 7 supralabials, the 3rd and 2010, almost every day, some short soft black hairs were 4th bordering the eye (HUM20110003 left 8 supralabials, found in their excrement, suggesting that they ate rodents the 4th and 5th bordering the eye); 9 infralabials; 1 loreal; or insectivores. 21–19–17 forebody-midbody-hindbody transverse dorsal Etymology: The specific name refers to the type locality, scale rows (21–21–17), weakly keeled on the back; 205 Zoige County, Sichuan, China. ventrals (212, 202); 69 pairs subcaudals (68, 79); anal plate divided. Body and headshape: body relatively 4. Discussion slender; venter round; head elongated and flattened, and slightly wider than the neck; pupils round. Coloration in Whereas traditional morphological approaches fail to life: body ground color off-white; clear red brown “M” define cryptic species, molecular data often succeed shaped marking with black edges formed at the margins (Burbrink and Castoe, 2009). Because of the relatively of the supraocular that borders the posterior margins of high rate of mutation, largely neutral evolution, and the frontal; a red brown postocular stripe with black edge large suite of characters, analyses of molecular data continuing to the last supralabial; four series of red brown augment those of morphological data and synergistically spots placed next to each other on back of body that they identify morphologically cryptic lineages. When range from posterior of the neck to vent and form four morphological differences are found, the data may be longitudinal rows; dorsal spots are usually composed of coded for phylogenetic analyses and combined molecular an entire (or incomplete) red brown scale and puce edges data to obtain a much more convincing conclusion at the borders of scales; spots of each stripe are similar in regarding species history, identification and diagnosis shape and range at relatively regular intervals of one or (Burbrink, 2001; Ling et al., 2010). two scales with slightly red brown color; ventrals, basal Ling et al. (2010) demonstrate that Elaphe zoigeensis part (close to trunk) black and distal ends non-uniform sp. nov. is a new species and likely the sister taxon to all gray, making its belly look like having many black rings. other extant Elaphe. While most species of Elaphe found The holotype laid 11 eggs on 29 July 2010 and the eggs in China are easily diagnosed, E. zoigeensis sp. nov., E. are white, smooth-shelled, and adhesive. dione, and E. bimaculata superficially resemble each other Ecology: The species is a diurnal terrestrial oviparous with respect to basic coloration (e. g., similar striping snake, laying eggs around the end of July, with a clutch pattern) and aspects of scutellation. Utiger et al. (2002) size of at least 11 eggs. Eggs are white, smooth-shelled, confirm that E. dione and E. bimaculata are likely valid and adhesive. From the day of capture until 3 August species based on an analysis of genetic distances. Elaphe 44 Asian Herpetological Research Vol. 3 zoigeensis sp. nov. is morphologically unique in its dorsal snakes disperse into the New World? Mol Phylogenet Evol, 43: striping, pattern of spotting, and number of preoculars. 173–189 Some specimens from Zoige County and surrounding Burbrink F. 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