A sheep in wolf's clothing: Elaphe xiphodonta sp. nov. (Squamata, Colubridae) and its possible mimicry to Protobothrops jerdonii
expand article infoShuo Qi§, Jing-Song Shi§|, Yan-Bo Ma§, Yi-Fei Gao§, Shu-Hai Bu, L. Lee Grismer#, Pi-Peng Li§, Ying-Yong Wang
‡ Sun Yat-sen University, Guangzhou, China
§ Shenyang Normal University, Shenyang, China
| Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China
¶ Northwest Agriculture and Forestry University, Yangling, China
# La Sierra Univer­sity, Riverside, United States of America
Open Access


Based on combined morphological and osteological characters and molecular phylogenetics, we describe a new species of the genus Elaphe that was discovered from the south slope of the Qinling Mountains, Shaanxi, China, namely Elaphe xiphodonta sp. nov. It is distinguished from the other congeners by a combination of the following characters: dorsal scales in 21-21-17 rows, the medial 11 rows keeled; 202–204 ventral scales, 67–68 subcaudals; two preoculars (including one subpreocular); two postoculars; two anterior temporals, three posterior temporals; reduced numbers of maxillary teeth (9+2) and dentary teeth (12); sharp cutting edges on the posterior or posterolateral surface of the rear maxillary teeth and dentary teeth; dorsal head yellow, three distinct markings on the head and neck; a distinct black labial spot present in supralabials; dorsum yellow, 46–49 complete (or incomplete) large black-edged reddish brown blotches on the body and 12–19 on the tail, two rows of smaller blotches on each ventrolateral side; ventral scales yellow with mottled irregular black blotches, a few irregular small red spots dispersed on the middle of the ventral. Based on molecular phylogenetic analyses, the new species forms the sister taxon to E. zoigeensis. The discovery of this new species increases the number of the recognized species in the genus Elaphe to 17.


Colubrid, morphology, osteology, Qinling Mountains, taxonomy


The colubrid genus Elaphe sensu lato Fitzinger (in Wagler), 1833, once contained approximately forty species ranging throughout temperate, subtropical, and tropical zones in both the eastern and western hemispheres (Schulz 1996). Most of the members of this genus have a slender body, partially or completely keeled dorsal scales, and round pupils. The head is distinguishable from the neck, the trunk vertebra lack a hypapophysis, and the posterior maxillary teeth are not significantly differentiated from the others. With the rise of molecular taxonomy in the last decades of the 20th century, a series of major taxonomic changes occurred at generic and species levels, resulting in the establishment or resurrection of several genera and species (Helfenberger 2001; Lenk Joger and Wink 2001; Utiger et al. 2002; Huang et al. 2012; Jablonski et al. 2019). Recent molecular phylogenetic studies suggest that the genus Orthriophis should be subsumed within the genus Elaphe, because the generic status of Orthriophis renders Elaphe paraphyletic, where E. zoigeensis Huang, Ding, Burbrink, Yang, Huang, Ling, Chen & Zhang, 2012 is sister to all the other Elaphe plus Orthriophis (Chen et al. 2017; Li et al. 2020; but see Figueroa et al. 2016). Currently, the genus Elaphe sensu stricto is comprised of 16 species of which most, are widely distributed in eastern Asia and the south slopes of the Himalaya, although the range of the genus extends east to eastern Russia, south to the Indonesia-Malayan region, and west to as far as Italy (Helfenberger 2001; Zhao 2006; Huang et al. 2012; Jablonski et al. 2019; Uetz et al. 2021). Previous biogeographic, phylogenetic, and phylogenomic studies support the hypothesis that Elaphe originated in the Eastern Palearctic (Lenk, Joger and Wink 2001; Utiger et al. 2002; Burbrink and Lawson 2007; Burbrink and Pyron 2010; Chen et al. 2017). In regards to China, 11 species of Elaphe are recognized: E. anomala (Boulenger, 1916), E. bimaculata Schmidt, 1925, E. carinata (Günther, 1864), E. cantoris (Boulenger, 1894), E. davidi (Sauvage, 1884), E. dione (Pallas, 1773), E. hodgsonii (Günther, 1860), E. moellendorffi (Boettger, 1886), E. taeniura (Cope, 1861), E. schrenckii (Strauch, 1873), and E. zoigeensis, two of which (E. bimaculata and E. zoigeensis) are endemic to China (Wang et al. 2020).

The main part of Qinling Mountains, lies on the south of Shaanxi Province, having an average elevation of approximately 2000 m and have long been regarded as the geographical, biological and climatological boundary between North (Palaearctic Realm; warm temperate monsoon climate) and South China (Oriental Realm; subtropical monsoon climate, Zhang 1999). Due to the unique environment and climate, the Qinling Mountains are the habitat of many rare animals (e.g., Ailuropoda melanoleuca, Budorcas bedfordi, and Rhinopithecus roxellana qinlingensis). Additionally, the herpetological diversity of that area is high. To date, more than 10 species of amphibians and 20 species of reptiles have been reported in this area, some of which are endemic to the Qinling Mountains and adjacent areas (e.g., Scutiger ningshanensis, Hyla tsinlingensis, Batrachuperus taibaiensis, Stichophanes ningshaanensis, Scincella tsinlingensis, Protobothrops jerdonii and Gloydius qinlingensis (Bu and Zheng 2015).

Bates (1862) discovered a spectacular type of adaptation known as “mimicry”, building on Charles Darwin's views on evolution. This phenomenon, now called “Batesian mimicry”, which is defined as an edible species (mimic) evolving to resemble a conspicuous inedible species (model), thereby gaining protection from predation, its efficiency relying on confusing the mimic with the model (Carpenter and Ford 1933; Ruxton et al. 2004). Batesian mimicry is observed among a wide variety of animals, ranging from invertebrates to vertebrates, including several non-venomous snakes mimicking the color pattern, head shape or behavior of sympatric venomous snakes to avoid predation (Brodie 1993).

During our recent herpetological surveys in the south slope of the Qinling Mountains, Shaanxi, China, two colubrid specimens were collected, which look quite different to any of the known species but similar to Protobothrops jerdonii (Figs 13). Detailed morphological examinations and further molecular analyses revealed that these specimens represent a separately evolving lineage within the genus Elaphe and can be distinguished from all congeners by morphological characters. We herein describe this overlooked Elaphe population as a new species. Furthermore, we suspect this new species is able to avoid predation by mimicking the syntopic pit-viper (P. jerdonii).

Figure 1. 

General view of the holotype (SYS r002534) of Elaphe xiphodonta sp. nov. in life. Photo by Shuo Qi.

Materials and methods


Morphological examinations were performed on two specimens collected from Chengguan Town, Ningshaan County, Shaanxi Province, China (Fig. 4). Both specimens were fixed in 10% buffered formalin after taking the tissue samples (liver or muscle), and then transferred to 70% ethanol for permanent preservation. The specimens are deposited in the Museum of Biology, Sun Yat-sen University (SYS r002534, Figs 1, 2, 3A) and Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences (IVPP OV 2721, Fig. 3C).

Figure 2. 

Detailed pholidosis of the head of the holotype (SYS r002534) of Elaphe xiphodonta sp. nov. Photos by Shuo Qi, illustrated by Xue-Man Zheng.

Figure 3. 

Comparison between Elaphe xiphodonta sp. nov. and sympatric Protobothrops jerdonii in different age stages A adult Elaphe xiphodonta sp. nov. (SYS r002534, holotype) B adult Protobothrops jerdonii C juvenile E. xiphodonta sp. nov. (IVPP OV 2721, paratype), road-killed specimen D juvenile P. jerdonii specimen in preservative. The black arrow points to the labial spot. Photos A, B, D by Shuo Qi, photo C by Liang Sun.

Morphological descriptions follow Dowling (1951) and Zhao (2006). The following measurements were taken with digital calipers (Neiko 01407A Stainless Steel 6-Inch Digital Caliper, USA) to the nearest 0.1 mm: TL total length (from tip of snout to tip of tail); SVL snout-vent length (from tip of snout to posterior margin of cloacal plate); TaL tail length (from posterior margin of cloacal plate to tip of tail); HL head length (from tip of snout to posterior margin of the mandible); HW maximum head width; ED eye horizontal diameter; RW = maximum rostral width; RH maximum rostral height.

Scalation features and their abbreviations are as follows: DSR dorsal scale rows, counted at one head length behind head, at midbody, and at one head length before vent; SPL supralabials; IFL infralabials; CS chin shields; PrO preoculars; PtO postoculars; LoR loreal; aTMP anterior temporals; pTMP posterior temporals; PrV preventral scales; V ventral scales; PrC precloacal plate; SC and subcaudals. Gender was determined by dissection or by the presence/absence of everted hemipenes. The numbers of maxillary teeth (MT) were counted based on the three-dimensional reconstructed model.

Other morphological characters (e.g., coloration, scalation, and size) for Elaphe taxonomy were obtained from Boulenger (1894), Stejneger (1907), Wen and Ji (1997), Zhao (2006), Huang et al. (2012), Jablonski et al. (2019), Shi et al. (2019) and Che et al. (2020).

The following abbreviations for museum collections are used throughout the paper:

AMNH American Museum of Natural History;

BFU Beijing Forest University;

BM British Museum;

CAS Chinese Academic of Sciences;

IVPP Institute of Vertebrate Paleontology and Paleoanthropology;

MHNG Muséum d’Histoire Naturelle, Genève;

SYS Sun Yat-sen University.

X-ray scanning and three-dimensional reconstructions

The X-ray scanning was carried out with Nano-computerized tomography. Specimens were scanned using a GE v|tome|x m dual tube 300/180kV system in the Key Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate Paleontology and Paleoanthropology (IVPP), Chinese Academy of Sciences. The specimen was scanned with an energy beam of 80 kV and a flux of 80*μA using a 360° rotation and then reconstructed into the 4096*4096 matrix of 1536 slices. The final CT reconstructed skull images were exported with a minimum resolution of 6.099 μm. The skull images were exported from the virtual 3D model which was reconstruct by Volume Graphics Studio 3.0.

The dataset of the 3D model included in this study is available online in the repository (ADMorph, Hou et al. 2020) at; (IVPP OV 2721, paratype).

DNA Extraction, Polymerase Chain Reaction (PCR) and sequencing

For the molecular analyses, two tissue samples of the new species were included, which were taken prior to fixation, preserved in 99% alcohol, and stored at -40 °C.

Genomic DNA was extracted from muscle or liver tissue samples, using a DNA extraction kit from Tiangen Bio-tech (Beijing) Co., Ltd. Partial segments of the mitochondrial genes, 16S ribosomal RNA gene (16S), cytochrome C oxidase 1 gene (CO1) and Cytochrome b gene (cytb) were amplified. Nested PCR experiments were performed as described in Li et al. (2017). Primers used for PCR and sequencing followed Li et al. (2020), see Table 1 for details. The first PCR procedure was performed with an initial denaturation at 94 °C for 4 min, 35 cycles of 94 °C for 45 s, 45 °C for 40 s and 72 °C for 2 min, followed by a final extension at 72 °C for 10 min. The second PCR procedure was performed with an initial denaturation at 94 °C for 4 min, 30 cycles of 94 °C for 45 s, 50 °C for 40 s and 72 °C for 1.5 min, followed by a final extension at 72 °C for 10 min. PCR products were purified with spin columns and then sequenced using BigDye Terminator Cycle Sequencing Kit as per the guidelines on an ABI Prism 3730 automated DNA sequencer by Guangzhou Tianyi Huiyuan Bio-tech Co., Ltd.

Table 1.

Nested PCR primers for this study (Li et al. 2020).

Gene Primer name Assay Sequence

Phylogenetic analyses

Fifty-nine sequences from 16 known Elaphe species plus six outgroup sequences from Euprepiophis mandarinus (Cantor, 1842) used to root the tree, were obtained from GenBank, and composed the dataset (Table 2).

Table 2.

Localities, specimen vouchers and GenBank accession numbers of the specimens included in this study.

No. Species name Locality Specimen voucher Genbank accession number References
16S rRNA CO1 Cytb
1 Elaphe xiphodonta sp. nov. Ningshaan, Shaanxi, China SYS r002534 MZ242100 MZ19164 MZ19166 This study
2 Ningshaan, Shaanxi, China IVPP OV 2721 MZ242101 MZ19165 MZ19167 This study
3 Elaphe anomala Huangshan, Anhui, China HS11075 MK193929 MK064632 MK201281 Li et al. 2020
4 Elaphe bimaculata Huangshan, Anhui, China HS15168 MK193931 MK064634 MK201283 Li et al. 2020
5 Elaphe cantoris Pailong, Tibet, China JK201705 MK194263 MK064913 MK201564 Li et al. 2020
6 Elaphe carinata Guangze, Fujian, China HS13055 MK193932 MK064635 MK201284 Li et al. 2020
7 Longyou, Zhejiang, China HS13062 MK193934 MK064637 MK201286 Li et al. 2020
8 Elaphe climacophora Abashiri, Hokkaido, Japan KUZ R64481 N/A LC328423 LC327534 Moriyama et al. 2018
9 Deshikutsu, Hokkaido, Japan KUZ R68813 N/A LC328426 LC327537 Moriyama et al. 2018
10 Elaphe davidi Taishan Mt., Shandong, China N/A KM401547 KM401547 KM401547 Xu et al. 2015
11 Elaphe dione Taibai, Shaanxi, China HS11036 MK193928 MK064631 MK201280 Li et al. 2020
12 Elaphe hodgsonii Jilong, Tibet, China HS13004 MK193983 MK064680 MK201335 Li et al. 2020
13 Elaphe moellendorffi Yunlin, Guangxi, China S-113 N/A KF698944 KF913314 Cao et al. 2014
14 Elaphe quadrivirgata N/A N/A AB738958 AB738958 AB738958 Direct Submission
15 Ashiu, Kyoto, Kansai, Japan As1352 N/A N/A HQ122007 Kuriyama et al. 2010
16 Elaphe quatuorlineata Crkvino, Northern Macedonia 1509 MK334307 MK334307 MK334307 Jablonski et al. 2019
17 Galatas, Argolida, Greece ZMUP 60 N/A N/A MH444348 Thanou et al. 2020
18 Elaphe sauromates Taganrogskyi Gulf, Russia ZISP 26197 N/A MK640250 N/A Jablonski et al. 2019
19 Solenoe Ozero, “Crimea” 1179 MK070315 MK070315 MK070315 Jablonski et al. 2019
20 Elaphe schrenckii Changbai, Jilin, China HS16031 MK193935 MK064638 MK201287 Li et al. 2020
21 Elaphe taeniura Zhouzhi, Shaanxi, China HS2010025 MK193982 MK064679 MK201334 Li et al. 2020
22 Heishiding, Guangdong, China SYS r001057 MK194113 MK064790 MK201445 Li et al. 2020
23 Elaphe urartica Kısıklı, Süphan Mts., Bitlis, Turkey ZDEU 26/2012 N/A MK640299 N/A Jablonski et al. 2019
24 Guzdak, Qobustan, Azerbaijan IZANAS T17 N/A MK640269 N/A Jablonski et al. 2019
25 Elaphe zoigeensis Zoige, Sichuan, China HS11251 MK193927 MK064630 MK201279 Li et al. 2020
26 Zoige, Sichuan, China HS2010015 MK193930 MK064633 MK201282 Li et al. 2020
27 Euprepiophis mandarinus HuangShan, Anhui, China HS12062 MK193939 MK064643 MK201291 Li et al. 2020
28 HuangShan, Anhui, China HS14017 MK193940 MK064644 MK201292 Li et al. 2020

DNA nucleotide sequences were aligned in the ClustalW algorithm with default parameters (Thompson et al. 1997) in MEGA 6 (Tamura et al. 2013). The aligned 16S, CO1 and cytb datasets were partitioned by codons with no gap positions allowed and applying default parameters in Gblocks version 0.91b (Castresana 2000). Three gene segments, with 1329 base pairs (bp) of 16S, 657 bp of CO1, and 585 bp of cytb, were concatenated seriatim into a 2571 bp sequence. With respect to the different evolutionary characters of each molecular marker, the dataset was split into seven partitions by gene and codon positions taking advantage of PartitionFinder 2.1.1 (Lanfear et al. 2012). The evolution models of each partition selected by PartitionFinder 2.1.1 were as follows: partition 1: 16S, GTR+I+G, 1287 bp; partition 2: COI\1, SYM+G, 219 bp; partition 3: COI\2, HKY+I, 219 bp; partition 4: COI\3, TVM+G, 219 bp; partition 5:;cytb\1, GTR+G, 195 bp; partition 6: cytb\2, HKY+I+G, 195 bp; partition 7:cytb\3, GTR+G 195 bp. General time-reversible (GTR) model. Sequence data were analyzed using Bayesian inference (BI) in MrBayes 3.2.4 (Ronquist et al. 2012), and maximum likelihood (ML) in RaxmlGUI 1.3 (Silvestro and Michalak 2012). Two independent runs were conducted in the BI analysis with 10,000,000 generations each and sampled every 1000 generations with the first 25% of samples discarded as burn-in, resulting in a potential scale reduction factor (PSRF) of < 0.005. In the ML analysis, a bootstrap consensus tree inferred from 1000 replicates was generated. Nodes with Bayesian posterior probabilities (BPP) ≥0.95 and ML support values of ≥70 were considered strongly supported (Huelsenbeck et al. 2001; Wilcox et al. 2002). Pairwise distances (p-distance) were calculated in MEGA6 using the uncorrected model. Gaps/Missing Data Treatment use the complete-deletion option, Substitutions to Include d: Transitions + Transversions option.

Taxonomic accounts

Elaphe xiphodontasp. nov.

Material examined

Holotype. SYS r002534, adult female (Figs 1, 2, 3A), collected by Yan-Bo Ma, Yi-Fei Gao on 4 September 2020 from Chengguan Town (33.58°N, 108.46°E (DD); ca 1731 m a.s.l.), Ningshaan County, Shaanxi Province, China. Paratypes. IVPP OV 2721, juvenile female (Fig. 3C), collected by Jing-Song Shi on 7 September 2020 from Chengguan Town (33.56°N, 108.50°E (DD); ca 1751 m a.s.l.), Ningshaan County, Shaanxi Province, China (Fig. 4).

Figure 4. 

Type terra of Elaphe xiphodonta sp. nov (marked with red triangles), with the collection localities of some other Chinese species of Elaphe.

Etymology. The specific epithet “xiphodonta” of the new species comes from the Ancient Greek “ξίφοσ (ksίfos, refer to ‘knife’ or ‘blade’)” and “δοντι (dónti, refer to ‘tooth’)”, meaning “blade-shaped teeth”, indicating that the new species has unique blade-shaped MT and DT (Figs 5, 6), which differs from the inconspicuous dental specializations (all teeth are cone-shaped) in its congeners. We suggest the Chinese formal name as “秦皇锦蛇” (Qín Huáng Jǐn Shé), which derived from Qin Shi Huang (personal name: Ying Zheng or Zhao Zheng; 259 BC–210 BC), the founder of the Qin dynasty and the first emperor of unified China, whose territory including the distribution range of Elaphe xiphodonta sp. nov. The English name is suggested as “Qin Emperor Rat Snake” or “Blade-teethed Rat Snake”.

Figure 5. 

Three-dimensional model of the skull of Elaphe xiphodonta sp. nov. (IVPP OV 2721, paratype). Left, dorsal view; middle, ventral view; right, lateral view. (Right palatomaxillary arch, mandible and suspensorium are not shown). Implemented by Peng-Fei Yin, Ye-Mao Hou and Jing-Song Shi.

Figure 6. 

Palatomaxillary apparatus and mandibles of Elaphe xiphodonta sp. nov. (IVPP OV 2721, paratype). Implemented by Peng-Fei Yin, Ye-Mao Hou and Jing-Song Shi A ventral (A1) and dorsal (A2) view of left palatomaxillary apparatus B posterolateral view of left maxilla (B1) and right dentary (B2), with cutting edges (ce) and caudolateral ridges indicated C labial (C1, right), ventrolateral (C2, left), ventral (C3, right, mirrored) and dorsal (C4, left) view of maxilla D Labial (D1) and lingual (D2) view of right dentary E labial (E1), lingual (E2), ventral (E3) and dorsal (E4) view of right mandible. Abbreviations: an. angular, at. atlas, ax. axis, bo. basioccipital, bs. basisphenoid, bt. blade teeth (“xiphodont”), ce. Posterior cutting edge of the blade teeth, chp. choanal process of palatine, clr. caudolateral ridge of the teeth, cp. compound bone, d. dentary, dpd. dorsal process of dentary, ecp. ectopterygoid, epm. ectopterygoid process of maxilla, etp. empty tooth position, exo. exoccipital, f. frontal, f5b. foramen for maxillary branch of trigeminal, f5c. foramen for mandibular branch of trigeminal, lf. lacrimal foramen, lfe. lateral furcula of ectopterygoid, m. maxilla, na. nasal, mfe. mesial furcula of ectopterygoid, mp. maxillary process of palatine, nf, nutrient foramen, p. parietal, pcr. prearticular crest of compound bone, pfr. prefrontal, pmx. premaxilla, po. postorbital, pp. palatine process of maxilla, pro. prootic, psp. parasphenoid rostrum, pt. pterygoid, pVf. posterior Vidian foramen, rpt. replacement teeth, sac. surangular crest of compound bone, smx, septomaxilla, so. supraoccipital, sp. splenial, st. supratemporal, v. vomer, vpd. ventral process of dentary, fs. fracture surface.


Elaphe xiphodonta sp. nov. can be differentiated from its congeners by the combination of the following morphological characters: (1) medium body size , SVL 785 mm in single adult female; (2) dorsal scales in 21-21-17 rows, the medial 11 rows keeled; (3) supralabials seven or eight, third/fourth (right) or fourth/fifth (left) in contact with eye, infralabials 9 or 10; (4) ventral scales 202–204; (5) subcaudals 67–68; (6) loreal single, not in contact with eye, not in contact with internasals; (7) two preoculars (including one subpreocular), two postoculars; (8) two anterior temporals, three posterior temporals; (9) precloacal plate divided; (10) reduced teeth number in maxilla and dentary bones (MT 9+2, DT 12; (11) sharp edges on the posterior or posterolateral surface of the rear MT and DT; (12) top of head yellow, three distinct markings on head and neck; (13) a distinct black labial spot present on supralabials; (14) ground color of dorsum yellow, 46–49 entire (or incomplete) reddish brown blotches with black edges on body and 12–19 similarly colored spots on tail; (15) ventral surface of body yellow with mottled irregular black blotches, a few irregular small red spots dispersed on middle of ventral scales.

Description of holotype

Adult female (Figs 1, 2, 3A). Body and tail slender, TL 967.5 mm (SVL 785.2 mm, TaL 182.3 mm, TaL/TL: 0.19); dorsal scales in 21–21–17 rows, the medial 11 rows keeled, the vertebral scales not enlarged; head elongate, moderately distinct from neck, longer than width and narrow anteriorly, HL 26.5 mm, HW 18.1 mm (HW/HL: 0.68); eye medium, ED 3.2 mm, pupil elliptic; rostral triangular, broader than high, RW 7.1 mm, RH 4.5 mm (RW/RH: 1.58; RW/HW: 0.39), visible from dorsum; nostril laterally pointed, located in the middle of nasal; nasal divided into two scales by nostril; two internasals, anteriorly rounded, bordered by two large prefrontals posteriorly; frontal single and enlarged, narrowed posteriorly; parietals paired, longer than width, in contact with each other medially, with upper anterior and posterior temporals laterally; one loreal on each side, in contact with nasal anteriorly, preocular posteriorly, prefrontals dorsally and the second supralabial ventrally; one enlarged preocular in contact with eye and supraocular posteriorly, prefrontal and loreal anteriorly, a smaller subpreocular ventrally, and not in contact with frontal; subpreocular in contact with eye and the third supralabial posteriorly, the second supralabial anteriorly, and preocular dorsally; two postoculars, upper one in contact with eye anteriorly, supraocular and parietal dorsally, and upper anterior temporal posteriorly, bottom one in contact with eye anteriorly, with upper and lower anterior temporals posteriorly, fifth and sixth supralabials below on left, and with fourth and fifth supralabials below on right; eight supralabials on left, seven supralabials on right (the third and the forth merged relative to left), first and second in contact with nasal, fourth and fifth reaching orbit on left, third and fourth reaching orbit on right; ten infralabials on left, nine infralabials on right, first pair in broad contact with each other, first to fifth in contact with anterior pair of chin shields, fifth in contact with posterior chin shields, fifth infralabial bipartitioned, lower part obviously larger than upper one; two pairs of chin shields, elongate, anterior pair larger, first pair meeting in midline, posterior pair is separated by three small scales; two similarly sized anterior temporals on left, lower one divided into two small scales on the right; three similarly sized posterior temporals on each side; 204 ventrals, excluding two preventrals; 67 pairs of subcaudals, excluding tail tip; precloacal plate divided.

Table 3.

Uncorrected P-distance of CO1 gene among 17 Elaphe species used in this study.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17
1 Elaphe xiphodonta sp. nov. 0.0
2 E. anomala 12.2 /
3 E. bimaculata 16.4 12.9 /
4 E. cantoris 14.2 17.9 18.4 /
5 E. carinata 13.9 11.3 13.4 17.9 0.3
6 E. climacophora 15.7 16.0 15.7 19.1 14.4 0.0
7 E. davidi 14.4 12.2 11.5 19.5 13.5 13.5 /
8 E. dione 15.3 16.0 10.9 18.2 14.3 13.9 14.2 /
9 E. hodgsonii 13.8 17.1 16.7 13.7 16.2 17.3 14.8 16.4 /
10 E. moellendorffi 14.6 16.4 17.1 13.7 16.6 16.1 17.4 16.8 12.2 /
11 E. quadrivirgata 10.7 8.7 12.5 17.9 8.5 14.9 11.4 12.6 13.4 16.6 /
12 E. quatuorlineata 17.1 14.1 12.5 18.4 13.8 17.9 12.5 13.6 16.1 15.8 12.5 /
13 E. sauromates 15.5 13.9 14.1 19.1 15.5 14.7 14.5 13.4 17.3 16.6 15.6 8.5 0.3
14 E. schrenckii 12.2 0.0 12.9 17.9 11.3 16.0 12.2 16.0 17.1 16.4 8.7 14.1 13.9 /
15 E. taeniura 15.2 16.3 16.7 16.0 16.0 15.5 15.6 15.3 13.2 16.2 13.8 14.7 16.1 16.3 6.8
16 E. urartica 14.1 14.0 13.9 16.8 12.7 13.7 13.3 11.8 15.2 15.6 12.5 9.4 8.9 14.0 16.5 0.0
17 E. zoigeensis 8.4 13.3 14.9 16.9 15.0 16.4 13.3 15.5 13.5 14.6 12.3 16.0 14.2 13.3 16.3 14.0 0.0
Table 4.

Uncorrected P-distance of cytb gene among 16 Elaphe species used in this study.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16
1 Elaphe xiphodonta sp. nov. 0.0
2 E. anomala 17.1 /
3 E. bimaculata 13.7 17.0 /
4 E. cantoris 17.2 16.6 18.2 /
5 E. carinata 18.7 14.1 14.6 19.1 0.0
6 E. climacophora 17.9 11.0 16.2 19.4 12.9 0.4
7 E. davidi 18.8 15.2 12.1 17.2 14.8 15.9 /
8 E. dione 19.6 19.0 15.0 15.2 18.6 17.6 16.1 /
9 E. hodgsonii 16.7 15.5 17.1 14.5 17.7 15.0 14.1 19.6 /
10 E. moellendorffi 21.7 18.5 21.8 16.3 16.4 17.6 18.6 20.7 14.8 /
11 E. quadrivirgata 19.7 13.1 15.4 16.9 12.6 13.3 13.8 15.5 15.8 18.9 0.9
12 E. quatuorlineata 16.7 13.0 16.5 14.5 13.7 11.0 13.7 15.5 17.4 17.1 11.7 0.0
13 E. sauromates 20.0 15.3 18.6 14.3 16.0 15.5 15.4 20.8 17.2 17.5 15.6 6.8 /
14 E. schrenckii 17.7 0.4 17.6 17.2 14.7 11.6 15.2 18.3 16.1 19.1 12.5 13.5 15.9 /
15 E. taeniura 16.3 17.8 15.4 12.8 16.1 17.1 15.0 16.6 12.9 15.9 14.7 14.0 15.0 18.4 5.4
16 E. zoigeensis 13.3 16.3 16.4 17.1 20.4 16.5 18.3 15.4 12.5 20.0 16.6 14.3 16.0 16.9 15.7 0.0

Coloration in life

Dorsal surface of head yellow, three distinct markings on head and neck; the anterior transverse black stripe, somewhat reddish medially, extends from the posterior margin of rostral and through the each eye to last two supralabials and adjacent small scales; interorbital arcuate cross-band, covering anterior part of frontals, most part of prefrontals, top of supraoculars, bottom half of upper postoculars, intact bottom postoculars, bottom half of upper temporals, most of bottom of temporals, dorsal edge of sixth left supralabials (fifth on the right), dorsal half of seventh supralabials (sixth) and bottom half of posterior-most supralabials, not reaching internasals, connected to largest posterior marking from mediolateral part; largest marking is a distinct black “M”-shaped marking that is reddish medially, covering the posteromedial part of the frontals, posterior part of supraoculars, most of parietals, dorsal margin of upper temporals, posteriorly extended, forming two thick black-edged reddish brown stripes on nape. Lateral surface of head yellow, a few small black spots dispersed on supralabials (2, 4 and 5 on left and 2, 3 and 4 on right) and subpreocular, a distinct scutellate black labial spot on the junction of the 2, 3 and right subpreocular (absent on left). Ventral surface of head consistently light-yellow, a few small black spots dispersed on the mental, infralabials, and anterior chin shields. An irregular spot occurs on the posterior edge of the junction of two anterior chin shields. Mouth lining is pale-heather and tongue is black.

Ground color of dorsal surface yellow, 49 complete or incomplete, black-edged reddish brown blotches on body and 12 similarly colored spots on tail; dorsal blotches on body approximately three to five scales in length, and eight to eleven scales rows in width; each blotch is usually composed of reddish brown scale with dark-brown edges. Two rows of smaller, black-edged reddish brown blotches on both side of the larger mid-body blotches, alternating with the mid-body blotches, each blotch covers 2–4 dorsal scales and separated from ventral scales by two rows of the dorsolateral scales. Ground color of ventral surface is yellow, mottled with irregular black blotches, a few irregular small red spots scattered midventrally.

Intraspecific variation

Measurements, body proportions and scale and pattern counts of the two specimens are listed in Table 5. The third and fourth supralabials are merged on right in holotype, which leads to the different supralabial counts. Regardless of the slight variation in scalation among the type series of E. xiphodonta sp. nov., the color pattern varies considerably between the juvenile and the adult: In adult one (SYS r002534, holotype), the color of the rostral, top and side of head, and chin, as well as dorsal and ventral sections of body are consistently light-yellow, whereas in the juvenile (IVPP OV 2721), the color of head and dorsal body is greyish to olive-green; the ventral scales and subcaudal scales are oyster white, with irregular greyish black spots. The juvenile has a similar dorsal pattern as the adult.

Table 5.

Measurements and scale counts and body proportions of Elaphe xiphodonta sp. nov.

Voucher SYS r002534 IVPP OV 2721
Sex female female
SVL 785.2 307.5
TaL 182.3 62.5
TL 967.5 370.1
TaL/TL 0.19 0.17
HL 26.5 17.64
HW 18.1 11.14
HW/HL 0.68 0.63
ED 3.2 2.8
RW 7.1 3.4
RH 4.5 2.0
RW/RH 1.58 1.70
RW/HW 0.39 0.31
DSR 21-21-17 21-21-17
SPL 8/7 8/8
IFL 10/9 11/11
CS 4 (2 pairs) 4 (2 pairs)
V 204 202
SC 67 67/68
MT 11 (9+2) 11 (9+2)
Dorsal blotches 61 (49+12) 65(46+19)


The osteological description is based on a road-killed juvenile individual (Paratype, IVPP OV 2721, Figs 5, 6). The skull is nearly completely preserved except for the slightly crushed parietal, and basioccipital bones.

Snout (Fig. 5). The premaxilla is short and blunt. The ascending process is laterally expanded and slanted posteriorly as in many borrowers (e.g., Euprepiophis and Archelaphe), rendering it hourglass-shaped in anterior view. The transverse process of premaxilla is flat, triangular-shaped in dorsal view. The posterior end of vomerine process of premaxilla contacts the tip of both vomer and septomaxilla. The posteromedial surface of vomerine process is expanded and oval shaped. The anterior section of the dorsal plate of nasal is tapered while the posterior section is expanded as oval shaped. The dorsal surface is slightly bulged.

Braincase (Fig. 5). The parietal is blunt and rounded, lacks a lateral crest on each side. The prefrontal is slender, somewhat rectangular. The anterolateral process is blunt. The anterior margin of frontal presents as trapezium-shaped. The lateral margin slightly concave. The postorbital is crescent-shaped, the anterodorsal process does not contact the frontal. The ventral process of postorbital is tapered, not furcated. The supraoccipital is rectangular and compressed, bearing a trifurcated dorsal ridge. The basisphenoid is wide and flat, with no conspicuous ridges on the ventral surface. The rostrum of parasphenoid is not divided. The angular surface between the basisphenoid and basioccipital is smooth. The columella is stubby, the shaft of columella quite short, approximately 1.2 times as length as the diameter of foot plate. The foramina for maxillary branch of trigeminal (f5b) and mandibular branch of trigeminal (f5c) nerves are oval shaped, not reaching the margin of prootic. The f5b is approximately 1/2 the width of f5c.

Palatomaxillary arch

(Fig. 6) The maxilla is slender. The maxillary nutrient foramen is transversely elongate, oval shaped and opens on the anterolateral side at the level of the third maxillary tooth and perforates the maxilla laterally. The anterior portion of maxilla curves slightly inward. The palatine process of maxilla is elongate while the ectopterygoid process is short and bunt.

The maxilla has 11 teeth on each side, with one or two rows of replacement teeth on the lingual side. In contrast to other species of Elaphe, the maxillary dentition of the new species is conspicuously differentiated. The anterior five MT have inconspicuous posterolateral ridges, while the posterior six teeth have a sharply edged ridge on their posterior margin (which could be also described as: the posterolateral ridge gradually moved posteriorly by the MT row, forming a sharp cutting edge on the posterior MT), forming blade-like teeth. The MT increase in size posteriorly, the posterior-most two being the largest, not separated from the anterior teeth by a diastema. The cutting edges of the posterior four MT slightly posteriorly convex, rendering them kukri shaped.

The palatine bears nine teeth, with one row of replacement teeth on the labial side. The choanal process of palatine (chp) forms a right triangular in dorsal view. The anterolateral margin of the maxillary process forms an approximate 30° angle with the medial line. The posterior margin of maxillary process is perpendicular to the medial line and collinear with the anterior margin of the choanal process. The pterygoid is slender and lanceolate in shape, 1.8 times the length of palatine, bearing 12 solid teeth (with one row of replacement teeth on the labial side). The ectopterygoid process and the medial transverse process of pterygoid are very small and difficult to see. The ectopterygoid is horizontally expanded, and outwardly curved in dorsal view. The labial furcula of ectopterygoid is oval, and distinct from the lingual process. The medial furcula (medial process) is elongate and spiculate, slightly curved ventrally.

Suspensorium and mandible

(Figs 5, 6) The mandible is slender and moderately curved. The supratemporal is flat and slender, fusiform, and the anteroventral margin is slightly angulated upward. The quadrate is triangulated in lateral view, and approximately the same length as supratemporal. The supratemporal angular surface of the quadrate is expanded, elongated and oval shaped. The prearticular crest (pcr) of the compound bone is prominent while the surangular crest (sac) is absent. The angular is slender and triangular. The splenial is triangular, coracoid shaped and shorter than angular. The dental bone bears 13 teeth, decreasing in size at the fifth tooth, with one or two rows of replacement teeth on the lingual side. The posterior tip of dorsal process of dentary extends farther posteriorly than the ventral process. The largest dentary nutrient foramen is elongated-oval shaped and lies below the seventh tooth.


(IVPP OV 2721, paratype) Maxilla: 11/11 (9+2); pterygoid: 12/12; palatine: 9/9; dentary: 13/13. Dentitional comparisons within the genus Elaphe and some related colubrid groups are listed in Table 6.

Table 6.

Dentition comparison of the Elaphe species and related colubrid species.

Taxon (n) Maxillary Blade teeth Palatine Pterygoid Dentary Reference
E. xiphodonta sp. nov. (1) 11/11 Y 9/9 12/12 13/13 this study
E. bimaculata (1) 19/20 N 9/10 15/16 18/19 this study
E. carinata (5) 17 N 9–11 13–16 19–21 this study
E. climacophora (2) 17 N 11 17–20 23–25 Helfenberger and Schätti 1998
E. dione (2) 16–20 N 7–9 12–13 18–23 this study
E. davidi (3) 16–17 N 9–12 12–16 16–19 Helfenberger and Schätti 1998
E. moellendorffi (2) 22–23 N 11 19–22 27–29 this study
E. schrenckii (2) 16–17 N 10–11 12–13 19–21 this study
E. taeniura (3) 17–23 N 10–11 16–19 19–23 this study
E. zoigeensis (2) 14–17 Y 10–10 10–13 16–17 this study
Eu. mandarinus (2) 16–19 Y 10–11 19–24 19–22 this study
Eu. perlaceus (2) 20–20 Y 12–12 18–21 21–22 this study
C. flavolineatus (1) 23 N 11/13 25/26 27/28 this study
C. philippinus (1) 25 N 12/13 26/27 29/28 this study
C. radiatus (5) 20–23 Y 10–12 18–24 23–27 this study
Ol. ornatus (1) 8/8 Y 4/4 5/5 13/13 this study
Oo. rufodorsatus (2) 18–19 Y 10–11 17–18 21–23 this study
G. oxycephalum (1) 23/23 N 10/10 14/13 26/26 this study
P. carinata (2) 24–25 Y 17–15 19–20 22 this study
P. dhumnades (1) 26 Y 21/20 23 26 this study
D. scabra (2) 7–6 N 8–9 0 3–3 Helfenberger and Schätti 1998
T. zhaoermii (2) 19–21 Y 12–16 18–23 21/24 this study


Detailed comparisons among Elaphe species are given in Table 7 and Fig. 7.

Figure 7. 

Comparisons of general morphological characteristics with its congeners in China A Elaphe xiphodonta sp. nov. (SYS r002534, holotype), Ningshaan, Shaanxi, by Shuo Qi B E. anomala, Benxi, Liaoning, by Shuo Qi C E. bimaculata, Hong’an, Hubei, by Chong-Jian Zhou D E. cantoris, Bomê, Tibet, by Jing-Song Shi E E. carinata, Mentougou, Beijing, by Jing-Song Shi F E. davidi, Benxi, Liaoning, by Jing-Song Shi G E. dione, Yongdeng, Gansu, by Shuo Qi H E. hodgsonii, Gyirong, Tibet, by Shuo Qi I E. moellendorffi, Chongzuo, Guangxi, by Jia-Jun Zhou J E. schrenckii, Baishan, Jilin, by Shuo Qi K E. taeniura from Hangzhou, Zhejiang, by Wei-Liang Xie L E. zoigeensis, Jiuzhaigou, Sichuan, by Jin-Wang.

Elaphe xiphodonta sp. nov. is distinct from all of its congeners by having fewer MT, enlarged posterior MT, cutting edges on both MT and DT, and three rows of large, black-bordered reddish brown dorsal blotches.

Additionally, Elaphe xiphodonta sp. nov. can be distinguished from E. cantoris, E. climacophora (Boie, 1826), E. hodgsoni, E. moellendorffi, and E. taeniura by having fewer ventral scales (202–204 vs. 226–239 in E. cantoris, 222–236 in E. climacophora, 228–247 in E. hodgsoni, 270–278 in E. moellendorffi, and 223–261 in E. taeniura), fewer subcaudals (67–68 vs. 78–87 in E. cantoris, 97–116 in E. climacophora, 72–92 in E. hodgsoni, 92–102 in E. moellendorffi, and 73–121 in E. taeniura), smaller body size (SVL 785 mm in single adult female vs. maximum SVL 1158 mm in E. cantoris, > 2000 mm in E. climacophora, 1190 mm in E. hodgsoni, 1602 mm in E. moellendorffi, and > 2000 mm in E. taeniura), and vastly different color pattern (Table 7).

Table 7.

Diagnostic characters separating all 17 species of the Elaphe, with distinguishing characters marked in bold. *: Counts of PrO contain subpreocular.

Species maximum SVL (in mm) DSR SPL IFL PrO* PtO TMP V SC
Elaphe xiphodonta sp. nov. 785 21-21-17 8 (7) 9 (10) 2 2 2+3 202–204 67–68
Elaphe anomala 1925 23 (21–25)-23 (19–23)-19 (17–19) 8 9–11 2 (1) 2 (1) 2 (3)+3 (2) 203–225 45–77
Elaphe bimaculata 760 23 (23–25)-23 (21–25)-19 (21) 8 (7) 9–12 2 (1) 2 2+3 170–209 61–81
Elaphe cantoris 1158 19 (20, 21)-19 (21–23)-17 8 9–10 2 2 2+3 (2) 226–239 78–87
Elaphe carinata > 2000 23 (21–25)-23 (21–25)-19 (17) 8 (9) 9–12 2 (1) 2 2+3 186–227 69–102
Elaphe climacophora > 1500 NA-23 (25)-NA 8 (9) 11 2 2 2+3 (2) 222–236 97–116
Elaphe davidi 1227 25 (22–27)-23 (22–25)-19 (17–21) 8 (7) 11–13 2 (1, 3) 2 (1–4) 2 (1, 3)+4 (2–3) 155–183 53–72
Elaphe dione 893 25 (21–27)-25 (21–27)-19 (17–21) 8 (9) 9–11 2 (1) 2 2+3 168–206 51–84
Elaphe hodgsoni 1190 23 (21–25)- 23 (21–25)-17 8 (9) 9–12 2 (1) 2 2 (1, 3)+3 (2, 4) 228–247 72–92
Elaphe moellendorffi 1602 25 (23–27)-27 (5)-19 (21) 9 (10) 10–13 2 2 2 (3)+3 (4) 270–278 92–102
Elaphe quadrivirgata >1000 NA-19-NA 8 11 2 2 2 (1)+3 (2) 195–215 70–96
Elaphe quatuorlineata > 2000 25-25 (23–27)-19 8 (9) 11 2 (3) 2 (3) 2 (3)+3 (4) 187–234 56–90
Elaphe sauromates 1250 25 (21–27)-25 (23, 24)-19 (18–21) 8 (7–10) 11 (9–12) 2 (1, 3) 2 (1) 2(1, 3)+ 4 (2–5) 199–222 61–79
Elaphe schrenckii 1335 23 (21)-23 (21)-19 8 (7) 8–11 2 (1) 2 2+3 (2) 208–224 57–75
Elaphe taeniura > 2000 25 (23)-23 (21, 25)-19 (17) 9 (6–10) 9–13 2 (1) 2 (3) 2 (1, 3)+3 (2–5) 223–261 73–121
Elaphe urartica 970 25 (23, 24)- 25 (23, 24)-19 (18) 8 (9) 11 (10–13) 2 (1, 3) 2 (1, 3) 2 (3)+ 4 (2, 3) 154–211 60–74
Elaphe zoigeensis 722 21-19(21)-17 7–8 9 3 2 2+3(2) 202–212 68–79

Elaphe xiphodonta sp. nov. can be differentiated from E. quatuorlineata Lacepede, 1789, E. sauromates (Pallas, 1811) and E. urartica (Jablonski, Kukushkin, Avci, Bunyatova, Ilgaz, Tuniyev & Jandzik, 2019) by having fewer dorsal scale rows (21-21-17 vs. 25-25 (23–27)-19 in E. quatuorlineata, 25 (21–27)-25 (23, 24)-19 (18–21) in E. sauromates and 25 (23, 24)- 25 (23, 24)-19 (18) in E. urartica) and a vastly different color pattern. Beyond that, E. xiphodonta sp. nov. can be further differentiated from E. quatuorlineata and E. sauromates by its smaller body size (SVL 785 mm in single adult female vs. maximum SVL > 2000 mm in E. quatuorlineata and 1250 mm in E. sauromates).

Elaphe xiphodonta sp. nov. can be differentiated from Elaphe carinata, E. davidi and E. quadrivirgata (Boie, 1826) by its smaller body size (SVL 785 mm in single adult female vs. maximum SVL > 2000 mm in E. carinata, 1227 mm in E. davidi, and > 1000 mm in E. quadrivirgata), having fewer subcaudals (67–68 vs. 69–102 in E. carinata, 70–96 in E. quadrivirgata), and a vastly different color pattern.

Despite the morphological similarities to E. bimaculata, E. dione, and E. zoigeensis, Elaphe xiphodonta sp. nov. differs from them by having different dorsal scale row counts (21-21-17 vs. 23 (23–25)-23 (21–25)-19 (21) in E. bimaculata), fewer preoculars (2 vs. 3 in E. zoigeensis), fewer MT (11/11 vs. 19/20 in E. bimaculata, 16–20 in E. dione, and 14–17 in E. zoigeensis (Table 6), blade-like posterior MT and the unique color pattern within the genus Elaphe.

Molecular phylogeny

The ML and BI analyses produced identical topologies, which were integrated in Fig. 8. The pairwise distances based on CO1 and cytb genes among all species are given in the Tables 3, 4, respectively.

Figure 8. 

Bayesian inferenced topology of 13 Elaphe species, based on the four partial mitochondrial DNA sequences (12S rRNA, 16S rRNA, CO1 and Cytb genes). BPP and BS values, respectively, occur at the nodes.

The phylogenetic analyses recovered a strongly supported monophyletic lineage containing all Elaphe (BS 100; BPP 1.00) which can be divided into three strongly supported clades (BS 95–97; BPP 1.00). Clade 1 includes all species previously in the genus “Orthriophis” with strong nodal support (BS 95; BPP 1.00). Notable intraspecific genetic differentiation within E. taeniura (mean p-distances 6.8% in CO1, 5.4% in cytb), pertains to different geographical clades.

The two samples from Chengguan Town, Shaanxi are clustered together with strong support (BS 100; BPP 1.00) with nearly no molecular divergence (mean p-distances 0% in CO1, 0% in cytb) between them. The clade of the above-mentioned specimens constitutes a sister clade with E. zoigeensis (Clade2, mean p-distances 8.4% in CO1, 13.3% in cytb).

Within Clade 3, the relationship between Elaphe anomala and E. schrenckii are worth noting. These two species form a strongly supported monophyletic group (BS 100; BPP 1.00) bearing low interspecific molecular divergence (mean p-distances 0.0% in CO1, 0.4% in cytb), suggesting they may be different color morphs of the same species, as mention before (An et al. 2010). However, given their distinctive color pattern differences and obvious geographical separation, we follow the current taxonomy.

Based on their phylogenetic relationships, genetic differentiation, and morphological distinctiveness (see Taxonomic accounts below), we hypothesize that the population from Chengguan Town, Shaanxi represents a separately evolving lineage and should be described as a new species.

Distribution, ecology and habit

Elaphe xiphodonta sp. nov. is currently known only from the Chengguan Town, Ningshaan County, Shaanxi Province, China. The new species inhabits sunny or semi-sunny gravels and bushes on slopes of less than 20°, along Chang’an River with an average width of 3 m. Elevation of the habitat ranges from 1700 to 1900 m. The vegetation types are Abies fargesii forest with artificial Picea asperata, Salix fargesii, Rubus koereana, Betula albosinensis and Fargesia qinlingensis. The canopy density is 0.75 (Bu and Zheng 2015). The new species is sympatric with Nanorana unculuanus, Scutiger ningshanensis, Euprepiophis perlaceus, Rhabdophis nuchalis, Stichophanes ningshaanensis, Gloydius qinlingensis and Protobothrops jerdonii (Fig. 9). When being captured, the new species flattens its head triangular and releases scent from the cloacal scent glands with smells a bit similar to P. jerdonii.

Figure 9. 

Habitats of Elaphe xiphodonta sp. nov. (Ningshaan County, Shaanxi), Provide by Shu-Hai Bu.

In feeding habits, the fecal samples from the holotype were checked and contained only feathers, indicating that this species is at least a bird-eater. The holotype was observed to prey on captive nesting quail and quail egg.


This study described a new Elaphe species which has a unique coloration and specialized teeth and had been overlooked for a long time during previous surveys. The new species was mistaken for P. jerdonii, which it mimics, at first glance during the field work. Nevertheless, the mimicry in E. xiphodonta is not unique within Elaphe. The coloration and head shape of E. davidi mimics that of the sympatric Gloydius spp. (Zhao 2006). Gloydius qinlingensis and Protobothrops jerdonii are sympatric with E. xiphodonta. Elaphe xiphodonta is similar to P. jerdonii in body shape, coloration, pattern and cloacal gland odor. Therefore, we hypothesize that E. xiphodonta is mimicking the model P. jerdonii as a means to avoid predation. Additional observations concerning the potential mimic-model relationship between E. xiphodonta and P. jerdonii are necessary.

Elaphe dione was previously widely recorded from northern China (Zhao 2006). Its distribution is currently restricted to Qinling Mountains-Huaihe River Line, and the records from other localities may belong to different taxa (e.g., E. zoigeensis, Huang et al. 2012). Previously collected specimens designated as E. dione (e.g., Fang and Song 1981) from Ningshaan County and surrounding areas are worth carefully re-examining.

Based on the dentition comparison in Table 6, it is clear that E. xiphodonta has far fewer MT and DT than most other species of Elaphe, which have more than 14 MT and DT. The tooth morphology of E. xiphodonta is similar to that of the genus Oligodon in a number of ways. Their reduced number and the specialized shape might relate to their unusual diet, since the morphology of the teeth in snakes have yet to be correlated with diet and prey-handling behaviors (Cundall and Irish 2008; Shi 2020). Some lizard-eating snakes tend to have numerous and closely spaced teeth (e.g., Scaphiodontophis, Sibynophis and Liophidium Zaher et al., 2012) while many oophagous snakes have reduced numbers of MT and DT (e.g., Dasypeltis). Blade-like rear teeth (sickle or kukri in shape, with sharp cutting edges on the posterior margins of two or more posterior MT) occur widely in many clades of the superfamily Colubroidea (e.g., Colubridae, Dipsadidae, Natricidae and Pseudoxenodontidae), most of which are anuran predators (e.g., Heterodon, Hydrodynastes and Rhabdophis) while some of the others tend to feed on eggs (e.g., Oligodon, Prosymna and Cemophora) or lizards (e.g., Orientocoluber and Hemorrhois). The diet and prey behaviors of the above-mentioned snakes have been widely reported in previous studies (Henderson 1984; Cundall and Greene 2000; Cundall and Irish 2008; Zaher et al. 2012), whereas far fewer comprehensive studies have focused on the function of blade-like teeth. A recent study indicated that a large proportion of anuran predators have blade-like rear teeth. Some snakes have been observed to deflate their anuran prey by puncturing and slicing their skin (e.g., Pseudoagkistrodon rudis; Shi 2020) or to consume the egg liquid (or embryos) of reptile eggs by cutting through their shells (e.g., Oligodon, Prosymna and Cemophora). Hence, we speculate that E. xiphodonta preys on frogs or eggs in the wild. In view of the limited numbers of the specimens in the present study, further field observation is needed to verify the above-mentioned hypothesis.


We are grateful to Li Ding for sharing important specimens for osteological comparison, to Fan Yang, Shi-Chao He, Zi-Chuan Wei, Jun Yan, Liang Sun, Jun-Dong Deng, Zong-Chang Yang, Bo Tan, Zhao-Chi Zeng and Zhi-Tong Lyu for their help in the field and the preparation of the manuscript; to Jin-Wang, Jia-Jun Zhou, Wei-Liang Xie, Chong-Jian Zhou and Liang Sun for providing important photographs; to Ye-Mao Hou, Peng-Fei Yin and Hao Ding for their assistance with CT scanning and three-dimensional reconstruction; to Xue-Man Zheng for the help with illustrations, and to Sheng Zheng for providing priceless literature references. We thank Dr. Luis Ceríaco, Dr. Robert Jadin and an anonymous reviewer for their helpful comments and suggestions on the manuscript.

This study is supported by “the Second Tibetan Plateau Scientific Expedition and Research Program” (Grant No. 2019QZKK0705), “the Specimen Platform of Ministry of Science and Technology, China, teaching specimen’s sub-platform” (No. 2005DKA21403-JK) and “the National Park Project of Central Financial Subsidy” (No. ZX2020-09-43).


  • An JH, Park DS, Lee JH, Kim KS, Lee H, Min MS (2010) No Genetic Differentiation of Elaphe schrenckii Subspecies in Korea Based on 9 Microsatellite Loci. Animal Systematics Evolution and Diversity 26(1): 15–19.
  • Boulenger GA (1894) Catalogue of the Snakes in the British Museum (Natural History) (Vol. II) Containing the Conclusion of the Colubridae Aglyphae. Trustees of the British Museum (Natural History). Taylor and Francis, London.
  • Bu S, Zheng X (2015) An illustrated guide to wildlife in Huoditang forest region of Qinling Mountain. Science Press, Beijing, 71–79. [in Chinese]
  • Burbrink FT, Pyron RA (2010) How does ecological opportunity influence rates of speciation, extinction, and morphological diversification in New World rat snakes (tribe Lampropeltini)? Evolution 64: 934–943.
  • Carpenter GDH, Ford EB (1933) Mimicry. Methuen, London.
  • Che J, Jiang K, Yan F, Zhang YP (2020) Amphibians and Reptiles in Tibet: Diversity and Evolution. Science Press, Beijing, 648–659. [In Chinese]
  • Chen X, Lemmon AR, Lemmon EM, Pyron AR, Burbrink FT (2017) Using phylogenomics to understand the link between biogeographic origins and regional diversification in rat snakes. Molecular Phylogenetics and Evolution 111: 206–218.
  • Cundall D, Irish F (2008) The Snake Skull. Biology of the Reptilia. Contributions to Herpetology, New York.
  • Fang RS, Song MT (1981) A preliminary survey of snakes from southern slope of Eastern Qinling Mountains. Journal of Shaanxi Normal University (Natural Science Edition): 263–272.
  • Figueroa A, Mckelvy AD, Grismer LL, Bell CD, Lailvaux SP (2016) A species-level phylogeny of extant snakes with description of a new Colubrid subfamily and genus. PLoS ONE 11(9): e0161070.
  • Helfenberger N (2001) Phylogenetic relationships of Old-World rat snakes based on visceral organ topography, osteology, and allozyme variation. Russian Journal of Herpetology 8: 1–62.
  • Helfenberger N, Schätti B (1998) Morphological adaptations for egg-eating in the snake Elaphe davidi Sauvage 1884. Russian Journal of Herpetology 5(1): 36–42.
  • Hou YM, Cui XD, Canul-Ku M, Jin SC, Hasimoto-Beltran R, Guo QH, Zhu M (2020) ADMorph: a 3D Digital Microfossil Morphology Dataset for Deep Learning. IEEE Access 8: 148744–148756.
  • Huang S, Ding L, Burbrink FT, Yang J, Huang J, Ling C, Chen X, Zhang Y (2012) A new species of the genus Elaphe (Squamata: Colubridae) from Zoige County, Sichuan, China. Asian Herpetological Research 3(1): 38–45.
  • Huelsenbeck JP, Ronquist F, Nielsen R, Bollback JP (2001) Bayesian Inference of Phylogeny and Its Impact on Evolutionary Biology. Science 294: 2310–2314.
  • Jablonski D, Kukushkin OV, Avcı A, Bunyatova S, Kumlutaş Y, Ilgaz Ç, Polyakova E, Shiryaev K, Tuniyev B, Jandzik D (2019) The biogeography of Elaphe sauromates (Pallas, 1814), with a description of a new rat snake species. PeerJ 7: e6944.
  • Jablonski D, Soltys K, Kukushkin OV, Simonov E (2019) Complete mitochondrial genome of the Blotched snake, Elaphe sauromates (Pallas, 1814). Mitochondrial DNA Part B 4(1): 468–469.
  • Kuriyama T, Brandley MC, Katayama A, Mori A, Honda M, Hasegawa M (2011) A time-calibrated phylogenetic approach to assessing the phylogeography, colonization history and phenotypic evolution of snakes in the Japanese Izu Islands. Journal of Biogeography 38: 259–271.
  • Lanfear R, Calcott B, Ho SY, Guindon S (2012) PartitionFinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution 29(6): 1695–1701.
  • Lenk P, Joger U, Wink M (2001) Phylogenetic relationships among European rat snakes of the genus Elaphe Fitzinger based on mitochondrial DNA sequence comparisons. Amphibia-Reptilia 22(3): 329–339.
  • Li JN, He C, Guo P, Zhang P, Liang D (2017) A workflow of massive identification and application of intron markers using snakes as a model. Ecology and Evolution 7: 10042–10055.
  • Li JN, Liang D, Wang YY, Guo P, Huang S, Zhang P (2020) A large-scale systematic framework of Chinese snakes based on a unified multilocus marker system. Molecular phylogenetics and evolution 148: e106807.
  • Moriyama J, Takeuchi H, Ogura-Katayama A, Hikida T (2018) Phylogeography of the Japanese rat snake, Elaphe climacophora (Serpentes: Colubridae): impacts of Pleistocene climatic oscillations and sea-level fluctuations on geographical range. Biological Journal of the Linnean Society 124: 174–187.
  • Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539–542.
  • Schulz K-D (1996) A monograph of the colubrid snakes of the genus Elaphe, Fitzinger. Koeltz Scientific Books, Hessen, 439 pp.
  • Shi JS, Jiang ZW, Zhao W, Shi Y (2019) Elaphe zoigeensis Found in Tewo Country, Gansu Province. Chinese Journal of Zoology 54(5): 769–770.
  • Shi JS (2020) Morphological diversity and character evolution of the maxillary teeth in Colubroidea (Reptilia: Serpentes). University of Chinese Academy of Sciences.
  • Tamura K, Stecher G, Peterson D, Filipski A, Kumar S (2013) MEGA6: molecular evolutionary genetics analysis, version 6.0. Molecular Biology and Evolution 30: 2725–2729.
  • Thanou E, Kornilios P, Lymberakis P, Leaché AD (2020) Genomic and mitochondrial evidence of ancient isolations and extreme introgression in the four-lined snake. Current Zoology 66(1): 99–111.
  • Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Higgins DG (1997) The CLUSTAL_X windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research 25: 4876–4882.
  • Utiger U, Helfenberger N, Schätti B, Schmidt C, Ruf M, Ziswiler V (2002) Molecular systematics and phylogeny of Old World and New World ratsnakes, Elaphe auct., and related genera (Reptilia, Squamata, Colubridae). Russian Journal of Herpetology 9(2): 105–124.
  • Wang K, Ren JL, Chen HM, Lyu ZT, Guo XG, Jiang K, Chen JM, Li JT, Guo P, Wang YY, Che J (2020) The updated checklists of amphibians and reptiles of China. Biodiversity Science 28(2): 189–218.
  • Wen SS, Ji DM (1997) The study of Elaphe davidi from China. Chinese Journal of Zoology 32(2): 16–19.
  • Wilcox TP, Zwickl DJ, Heath TA., Hillis DM (2002) Phylogenetic relationships of the Dwarf Boas and a comparison of Bayesian and bootstrap measures of phylogenetic support. Molecular Phylogenetics and Evolution 25: 361–371.
  • Xu CZ, Mu YS, Kong QR, Xie GL, Guo ZR, Zhao S (2016) Sequencing and analysis of the complete mitochondrial genome of Elaphe davidi (Squamata: Colubridae). Mitochondrial DNA Part A 27(4): 2383–2384.
  • Zhang RZ (1999) Zoogeography of China. Science Press, Beijing, 502 pp. [In Chinese]
  • Zhao EM (2006) Snakes of China. I. Anhui Science and Technology Publishing House, Hefei, 197–211. [In Chinese]

Appendix 1

Details of the osteological specimens examined for dentition comparisons in this study.



E. xiphodonta sp. nov. IVPP OV 2721, 3D model of impregnated specimens, IVPP.

E. zoigeensis IVPP OV 2672, 3D model of impregnated specimens, IVPP.

E. carinata IVPP OV 2296, osteological specimens, IVPP.

E. climacophora SH 530, lateral and dorsal views of skull (Helfenberger and Schätti 1998).

E. davidi BM 1916.1.15.17, lateral and dorsal views of skull, BM (Helfenberger and Schätti 1998).

E. dione IVPP OV 2302, osteological specimen, IVPP.

E. moellendorffi IVPP OV 2686, osteological specimen, IVPP.

E. schrenckii IVPP OV 2295, osteological specimen, IVPP.

E. taeniura IVPP OV 2298, osteological specimen, IVPP.


C. flavolineatus IVPP OV 2687, osteological specimen, IVPP.

C. radiatus IVPP OV 2411–2415, osteological specimens, IVPP.


Eu. mandarinus IVPP OV 2675, osteological specimen, IVPP.


Ol. ornatus SYS r001297, 3D model of Impregnated specimens, SYS.


Oo. Rufodorsatus IVPP OV 2691, osteological specimen, IVPP.


P. dhumnades IVPP OV 2686, osteological specimen, IVPP.



D. scabra MHNG 1362.78, lateral and dorsal views of skull (Helfenberger and Schätti 1998). MHNG; AMNH r-31638, 3D model of impregnated specimens, AMNH.


T. zhaoermii BFU R_Th_001. 3D model of impregnated specimen, BFU.