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Micrelapidae new family. Type genus: Micrelaps Boettger, 1880. Type species: Micrelaps muelleri Boettger, 1880. Etymology: Boettger (Bottger ¨) did not give the etymology for the generic nomen but was almost certainly from the Latin adjective micro-, derived from the Greek mikros (small), and elaps, the Latinised form of the Greek noun´ellops or´elaps (literally sea-fish or serpent, but here in reference to the snake genus Elaps, now a synonym of Homoroselaps). Micrelapidae fam. nov. is derived from Micrelaps by the taking the stem elap- of the root word of the nomen. Content: Micrelaps muelleri Boettger, 1880, Micrelaps bicoloratus Sternfeld, 1908, Micrelaps vaillanti Mocquard, 1888, Brachyophis revoili Mocquard, 1888. Diagnosis and definition: In the crania of Micrelaps and Brachyophis we examined the ectopterygoid was laterally and medially expanded at the point of contact with the pterygoid, with this expansion not being contiguous with the ectopterygoid anterolateral and anteromedial lobes (Fig. 4, Supplementary material fig. 53, 54). The lateral expansion is a posterolaterally and somewhat ventrally directed, very prominent protuberance continuous with a ridge on the ventral surface of the pterygoid. This character state was not present in any other cranium we examined and is very likely a synapomorphy of the family. Other common cranial characters include a premaxilla adapted for a fossorial lifestyle, premaxillary transverse processes closely approaching the maxilla, a short maxilla with ascending processes abutting the prefrontal, well-developed, grooved fangs below the orbit, preceded by a diastema and 2 – 3 teeth, an ectopterygoid deeply forked into anterolateral and anteromedial lobes that articulate with maxillary ectopterygoid processes leaving a foramen in the middle, prefrontal and parietal supraorbital processes laterally bordering the frontal and almost meeting each other, a tendency towards fusion of cranial bones (especially because the supratemporal is absent, very likely fused to the quadrate in Brachyophis and to posterior chondrocranial elements in Micrelaps), and a short quadrate. Brachyophis, however, differs from the type genus in possessing a postorbital (versus postorbital absent in Micrelaps), dorsolateral adductor ridges on the parietal (versus a single sagittal ridge in Micrelaps), only a faint pseudocoronoid ridge on the dentary (versus a prominent process in Micrelaps). Scalation characters that are common in both genera include 1 nasal, 7 supralabials, 15 smooth dorsal scale rows, absence of a loreal, 2 anal shields. Ventrals range from 170 to 280 and subcaudals (paired) 16 – 32 in Micrelaps (Boulenger, 1896; De Witte and Laurent, 1947; Rasmussen, 2002; Werner et al., 2006; Spawls et al., 2018). In Brachyophis, ventrals range from 103 to 123 and subcaudals (single) 8 – 14 (De Witte and Laurent, 1947; Lanza, 1966). Brachyophis has a large, azygous occipital shield (Boulenger, 1896). Micrelaps and Brachyophis possess a rectal caecum and a short genital sinus in the female, two soft tissue traits used to cluster these two genera by Underwood and Kochva (1993). Distribution: Micrelaps spp. is distributed in eastern and northeastern Africa and western Asia. Brachyophis is limited to Somalia in north-eastern Africa. Distribution: Micrelaps spp. is distributed in eastern and northeastern Africa and western Asia. Brachyophis is limited to Somalia in north-eastern Africa. Remarks: Geniez (2018) commented that Micrelaps “could constitute a separate family within its own right, that of Micrelapsidae”. Bar et al. (2021) likewise wrote that “. The actual placement of the genus [Micrelaps] is often poorly supported within studies and inconsistent across them. We suspect it will soon be placed in its own family — as is the norm in taxonomy these days. We predict this family, containing a single genus (Micrelaps), will be called Micrelapidae.”. However, these authors did not explicitly express that they are erecting a new family for these snakes. Rather, it was a suggestion about what should/could be done. It therefore is not in accordance with Article 16.1 and Recommendation 16A of The Code (ICZN, 1999). They also did not also provide characters for the express purpose of differentiating or diagnosing “Micrelapsidae” or “ Micrelapidae ”, nor did they cite a work containing the same (again, very likely because a nomenclatural act presumably was not the intention of Geniez [2018] and it was not the intention of SM, who wrote this in Bar et al. [2021] either) and this contravenes Articles 13.1.1, 13.1.2 and Recommendation 13A of The Code. Hence, we regard the nomen “Micrelapsidae” as unavailable. The ZooBank LSID for this taxonomic action is urn:lsid:zoobank.org:pub:D8475246-AD8E-4886-AB55- 12F6F242E9C4., Published as part of Das, Sunandan, Greenbaum, Eli, Meiri, Shai, Bauer, Aaron M., Burbrink, Frank T., Raxworthy, Christopher J., Weinell, Jeffrey L., Brown, Rafe M., Brecko, Jonathan, Pauwels, Olivier S. G., Rabibisoa, Nirhy, Raselimanana, Achille P. & Merila, Juha, 2023, Ultraconserved elements-based phylogenomic systematics of the snake superfamily Elapoidea, with the description of a new Afro-Asian family, pp. 1-11 in Molecular Phylogenetics and Evolution 180 on pages 8-9, DOI: 10.1016/j.ympev.2022.107700, http://zenodo.org/record/7746501, {"references":["Sternfeld, R., 1908. Zur Schlangenfauna Ostafrikas. I. Schlangen aus Sud-Abessinien. Mitt. Zool. Mus. Berlin 4, 239 - 247.","Mocquard, F., 1888. Sur une collection de Reptiles et de Batraciens rapport´es des Pays Comalis et de Zanzibar par M. G. R´evoil. M´emoires Publies par la Soci´ete´ Philomathique a l' occasion du Centenaire de sa fondation 1788 - 1888, 109 - 134.","Boulenger, G. A., 1896. Catalogue of the Snakes in the British Museum (Natural History). Volume III, containing the Colubridae (Opisthoglyphae and Proteroglyphae), Amblycephalidae, and Viperidae. Trustees of the British Museum, London.","de Witte, G. - F., Laurent, R., 1947. R´evision d' un groupe de Colubridae africains: genres Calamelaps, Miodon, Aparallactus et formes affines. M´em. Mus. R. His. Nat. Belg. 29, 1 - 134.","Rasmussen, J. B., 2002. A review of the African members of the genus Micrelaps Boettger 1880 (Serpentes Atractaspididae). Tropical Zoology 15, 71 - 87.","Werner, Y. L., Babocsay, G., Carmely, H., Thuna, M., 2006. Micrelaps in the southern Levant: variation, sexual dimorphism, and a new species (Serpentes: Atractaspididae). Zool. Middle East 38, 29 - 48.","Spawls, S., Howell, K., Hinkel, H., Menegon, M., 2018. Field Guide to East African Reptiles, second edition. Bloomsbury Wildlife, London.","Lanza, B., 1966. Il genere Brachyophis e descrizione di una nuova forma (Reptilia, Serpentes, Colubridae). Monit. Zool. Ital. 74, 30 - 48.","Underwood, G., Kochva, E., 1993. On the affinities of the burrowing asps Atractaspis (Serpentes: Atractaspididae). Zool. J. Linn. Soc. 107, 3 - 64.","Geniez, P., 2018. Snakes of Europe, North Africa and the Middle East: A Photographic Guide. Princeton University Press, Princeton and Oxford.","Bar, A., Haimovitch, G., Meiri, S., 2021. Field guide to the amphibians and reptiles of, Israel. Edition. Chimaira, Frankfurt Am Main.","ICZN (International Commission on Zoological Nomenclature), 1999. International Code of Zoological Nomenclature, fourth edition. International Trust for Zoological Nomenclature, London."]}

Yang, Dian-Cheng, Huang, Ru-Yi, Jiang, Ke, Burbrink, Frank T., Yu, Yan-An Gong Jing, Zhang, Yi, Huang, Tian-Qi, Huang, Song (2022): A new species of the genus Achalinus (Squamata: Xenodermidae) from Ningshan County, Shaanxi Province, China. Zootaxa 5190 (1): 127-140, DOI: https://doi.org/10.11646/zootaxa.5190.1.5

Achalinus ningshanensis sp. nov. D.C. Yang, R.Y Huang, K. JIANG, F.T. BURBRINK & S. Huang (Figs. 3–5) Suggested English common name: Ningshan Odd-scaled Snake. Suggested Chinese common name: Ťkḛffi (Bopomofo: Níng Shǎn Jǐ Shé). Holotype. ANU20220001 (field number JK0917, Fig. 3), an adult female, collected in summer of 2008 from Xunyangba Town (33.5434 N, 108.5439 E, 1372 m a. s. l.), Ningshan County, Shaanxi Province, China by the team of Ke Jiang. Paratypes. Six females, JK0918, JK0889, JK0890, JK0891, HSR19131, and HSR19132 (Voucher number: ANU20220002–7), all with the same collecting information as the holotype. Diagnosis. Achalinus ningshanensis sp. nov. can be differentiated from all other members of the genus Achalinus by the following combination of morphological characters:(1) dorsum iridescent and uniformly dark brown in preservative, longitudinal vertebral line absent; (2) brown beneath; (3) dotted black streak in the middle of the subcaudals absent; (4) tail length relatively short, TaL/ToL 12–16% in females; (5) fewer subcaudals, 41–46, in females; (6) dorsal scales 23 rows throughout, strongly keeled, and the outer-most rows on both sides of the body also keeled and slightly enlarged; (7) one loreal; (8) internasal not fused to prefrontal; (9) suture between internasals is similar size when compared to the suture between prefrontals; (10) preocular and postocular absent; (11) 6 supralabials; (12) 5 infralabials, the first 3 (rarely 2) touching the first pair of chin shields; (13) 3 pairs of chin shields. Description of the holotype. An adult female in excellent condition (Figs. 3 & 4). Body slender and subcylindrical, total length 436 mm (SVL 374 mm and Tal 62 mm); tail relatively shorter, TaL/ToL 14%; head slightly distinct from neck dorsally covered with large head scales, head length 13.72 mm; head width 7.16 mm; eyes small with subround pupils. Rostral small, subtriangular in frontal view, barely visible from above. Internasals paired, the suture between internasals (1.94 mm) subequal to that between prefrontals (1.63 mm); prefrontals paired, subrectangular. Frontal shield-shaped, nearly straight anteriorly, pointed backwards, slightly broader than it is long, and much shorter than the parietals. Parietals paired, the parietal suture longer than frontal. Nostril in the anterior part of the nasal. One loreal, rectangular, LorH: 0.97 mm, LorL: 1.90 mm (LorH/LorL: 0.51), extending from the nasal to the eye. One supraocular, in contact with loreal, prefrontals, frontal, parietals, and superior anterior temporals. No preocular or postocular. Temporals 2+3+4, two anterior temporals elongated, the upper one smaller in contact with parietal, the lower one in contact with fifth and sixth supralabials, both in contact with the eye; three elongated middle temporals; four elongated posterior temporals, the uppermost (super-temporal) significantly enlarged, the supertemporal split into two parts on the left, surrounding the parietal; the super-temporals from two sides contacting at the midline. Supralabials six, the first very small, 1–3 contacting the nasal, fourth and fifth contacting the orbit, fourth in narrow contact, fifth in broad contact, and sixth the longest and largest. One mental. Five infralabials, the first pair in contact blocking the mental from contacting an anterior pair of chin shields; first infralabial smallest, increasing in size until the fifth infralabial that is the longest. Three pairs of chin shields followed by ventrals. First three infralabials touching the first pair of chin shields on both sides. Dorsal scale rows 23–23–23, dorsal scales fusiform, not overlapping, distinctly keeled posterior to nape, and outermost dorsal scale row on both sides slightly keeled and enlarged. Ventrals 167; cloacal plate entire; subcaudals 43, uniserial. Coloration of holotype in preservative. Dorsum uniformly dark brown with metallic luster under light, the anterior portion of the head ash black, interstitial skin of dorsal dark brown. Ventral ground color brown, free margins of ventral scales grayish white. Variation. Measurements, body proportions, and scale counts are listed in Table 1. All paratypes are very similar to the holotype except in the following: paratypes JK0889, JK0891, and HSR19132 have two middle temporals; paratypes JK0890 and HSR19131 have two middle temporals and three posterior temporals; paratypes JK0890 have the first two infralabials touching the first pair of chin shields on both sides. The number of anterior temporals that contact the eyes is not a stable trait. Comparisons. Achalinus ningshanensis sp. nov. can be differentiated from all other known species of snakes in the genus Achalinus by a combination of the following morphological characters: infralabials five, three pairs of chin shields, relatively shorter tail length (TaL/ToL 12–16% in females), longitudinal vertebral line absent, and missing a dotted black streak in the middle of the subcaudals. Achalinus ningshanensis sp. nov. differs from all other species in Achalinus except A. spinalis Peters, 1869, A. hainanus Huang, 1975, A. emilyae Ziegler, Nguyen, Pham, Nguyen, Pham, Van Schingen, Nguyen & Le, 2019, A. huangjietangi Huang, Peng, & Huang, 2021, and A. dehuaensis Li, Wu, Xu, Zhu, Ren, Guo & Dong, 2021 by having fewer infralabials (five vs. six). Furthermore, it differs from A. spinalis and A. huangjietangi by having the following characters: absent longitudinal vertebral line, and missing a dotted black streak in the middle of the subcaudals; from A. hainanus by having two anterior temporals (vs. only one), three pairs of chin shields (vs. two pairs), fewer subcaudals in females (41–46 vs. 67–69); from A. emilyae by having fewer subcaudals in females (41–46 vs. 63), three pairs of chin shields (vs. two pairs), outermost dorsal scales rows keeled (vs. outmost rows smooth); from A. dehuaensis by having a relatively shorter tail length (TaL/ToL 12–16% vs. 21–22% in females), fewer subcaudals in females (41–46 vs. 63–65), uniformly dark brown above, light brown beneath (vs. greyish brown above, pale yellow beneath). Achalinus ningshanensis sp. nov. differs from A. werneri Van denburgh, 1912 by having the following characters: absent longitudinal vertebral line, and missing a dotted black streak in the middle of the subcaudals. Achalinus ningshanensis sp. nov. differs from A. yangdatongi by having fewer infralabials (5 vs. 6), different dorsal scale row counts (23–23–23 vs. 23–23–19) and having three pairs of chin shields (vs. two pairs). Achalinus ningshanensis sp. nov. differs from A. ater by the suture between internasals subequal to that between prefrontals (vs. suture between internasals distinctly longer than between prefrontals) and having fewer subcaudals (41–46 vs. 56–63). Achalinus ningshanensis sp. nov. differs from A. juliani by the suture between internasals subequal to that between prefrontals (vs. the suture between internasals distinctly longer than between prefrontals) and having a relatively shorter tail length (TaL/ToL 12–16% vs. 22–37%), fewer subcaudals (41–46 vs. 77–91), different dorsal scale row counts (23–23–23 vs. 25–23–23), and having three pairs of chin shields (vs. two pairs). Achalinus ningshanensis sp. nov. differs from A. rufescens by having six supralabials, the fourth and fifth widely in contact with the eye (vs. five supralabials, third and fourth entering the eye) and being light brown beneath (vs. uniform yellowish beneath). Achalinus ningshanensis sp. nov. differs from A. formosanus Boulenger, 1908, A. jinggangensis (Zong & Ma, 1983), A. timi Ziegler, Nguyen, Pham, Nguyen, Pham, Van Schingen, Nguyen & Le, 2019, A. pingbianensis Li, Yu, Wu, Liao, Tang, Liu & Guo, 2020, A. zugorum Miller, Davis, Luong, Do, Pham, Ziegler, Lee, De Queiroz, Reynolds & Nguyen, 2020, by having a separated loreal (vs. absence). Further it is different from A. formosanus by having fewer midbody dorsal scale rows (23 vs. 27), three pairs of chin shields (vs. two pairs), fewer subcaudals in females (41–46 vs. 61–70); from A. jinggangensis by having fewer infralabials (5 vs. 6), three pairs of chin shields (vs. two pairs), fewer subcaudals in females (41–46 vs. 51); from A. timi by different dorsal scale row counts (23–23–23 vs. 25–25–23), fewer infralabials (5 vs. 6), three pairs of chin shields (vs. two pairs), and the suture between internasals subequal to that between prefrontals (vs. the suture between internasals distinctly longer than between prefrontals); from A. pingbianensis by having fewer infralabials (5 vs. 6), fewer supralabials (6 vs. 7), 3 pairs of chin shields (vs. 2 pairs), and being uniformly dark brown above, light brown beneath (vs. uniform black above and beneath), and from A. zugorum by having fewer infralabials (5 vs. 6), 3 pairs of chin shields (vs. 2 pairs). Achalinus ningshanensis sp. nov. differs from A. meiguensis Hu & Zhao, 1966 and A. panzhihuaensis by having divided internasals (vs. absent), absence of postocular (vs. presence), fewer infralabials (5 vs. 6), mental separated from anterior chin shields (vs. in contact), and the first pair of infralabials in contact (vs. separated). Furthermore, it differs from A. panzhihuaensis by different dorsal scale row counts (23–23–23 vs. 23–23–19). Achalinus ningshanensis sp. nov. differs from A. niger Maki, 1931 by having fewer infralabials (five vs. six), different dorsal scale row counts (23–23–23 vs. 25–25–23), and fewer subcaudals in females (41–46 vs. 52–58). Achalinus ningshanensis sp. nov. differs from A. tranganensis Luu, Ziegler, Ha, Lo, Hoang, Ngo, Le, Tran & Nguyen, 2020 by having fewer infralabials (five vs. six), three pairs of chin shields (vs. two pairs), a relatively shorter tail length (TaL/ToL 12–16% vs. 25%), and fewer subcaudals in females (41–46 vs. 73+). Achalinus ningshanensis sp. nov. differs from A. yunkaiensis Wang, Li & Wang, 2019 by having fewer infralabials (five vs. six), three pairs of chin shields (vs. two pairs), and the outer-most dorsal scales rows keeled (vs. outmost rows smooth). Etymology. The specific epithet refers to the location of type specimens, Ningshan County, Shaanxi Province, China. We suggest Níng Shǎn Jǐ Shé (Ťkḛffi) as Chinese common name, and Ningshan Odd-scaled Snake as English common name. Distribution and habits. Achalinus ningshanensis sp. nov. is currently known only from the type locality Ningshan County, Shaanxi Province, China. All the specimens were found in farmland. The surrounding habitat was composed of secondary conifer/broad-leaved mixed forest. The new species is sympatric with A. cf. spinalis, A. cf. rufescens, Stichophanes ningshaanensis (Yuan, 1983), Euprepiophis mandarinus (Cantor, 1842), Elaphe taeniura (Cope, 1861), E. carinata (Günther, 1864), E. xiphodonta Qi, Shi, Ma, Gao, Bu, Grismer, Li, Wang 2021, Oreocryptophis porphyraceus (Cantor, 1839), Cyclophiops major (Günther, 1858), Lycodon ruhstrati (Fischer, 1886), L. liuchengchaoi Zhang, Jiang, Vogel and Rao, 2011, Pseudoxenodon macrops (Blyth, 1855), Rhabdophis nuchalis (Boulenger, 1891), Zaocys dhumnades (Cantor, 1842), Azemiops kharini Orlov, Ryabov and Nguyen, 2013, Gloydius qinlingensis (Song and Chen, 1985), Protobothrops jerdonii (Günther, 1875), Published as part of Yang, Dian-Cheng, Huang, Ru-Yi, Jiang, Ke, Burbrink, Frank T., Yu, Yan-An Gong Jing, Zhang, Yi, Huang, Tian-Qi & Huang, Song, 2022, A new species of the genus Achalinus (Squamata: Xenodermidae) from Ningshan County, Shaanxi Province, China, pp. 127-140 in Zootaxa 5190 (1) on pages 133-136, DOI: 10.11646/zootaxa.5190.1.5, http://zenodo.org/record/7119993, {"references":["Peters, W. C. H. (1869) Nachtrag zu Uber neue Gattungen und neue oder weniger bekannte Arten von Amphibien (Eremias, Dicrodon, Euprepes, Lygosoma, Typhlops, Eryx, Rhynchonyx, Elapomorphus, Achalinus, Coronella, Dromicus, Xenopholis, Anoplodipsas, Spilotes, Tropidonotus). Monatsberichte der koniglichen Akademie der Wissenschaften zu Berlin, 1869, 445 - 447.","Huang, R. Y., Peng, L. F., Yu, L., Huang, T. Q., Jiang, K., Ding, L., Chang, J. K., Yang, D. C., Xu, Y. H. & Huang, S. (2021) A New Species of the Genus Achalinus from Huangshan, Anhui, China (Squamata: Xenodermidae). Asian Herpetological Research, 12 (2), 178 - 187. https: // doi. org / 10.16373 / j. cnki. ahr. 200075","Li, K., Wu, Y. Y., Xu, R. Y., Zhu, F., Ren, J. L., Guo, P. & Dong, B. J. (2021) A new species of the Achalinus rufescens complex (Xenodermidae: Achalinus) from Fujian Province, China. Zootaxa, 5026 (2), 239 - 254. https: // doi. org / 10.11646 / zootaxa. 5026.2.5","Van Denburgh, J. (1912) Concerning certain species of reptiles and amphibians from China, Japan, the Loo Choo Islands, and Formosa. Proceedings of The California Academy of Sciences, 3, 187 - 258.","Boulenger, G. A. (1908) Description of a new frog and a new snake from Formosa. Annals & Magazine of Natural History, 8 (2), 221 - 222. https: // doi. org / 10.1080 / 00222930808692472","Zong, Y. & Ma, J. (1983) A new species of the genus Achalinopsis from Jiangxi and the restoration of this genus. Acta Herpetologica Sinica, 2 (2), 61 - 63. [in Chinese]","Li, K., Yu, M., Wu, Y. Y., Liao, L. H., Tang, K., Liu, Q. & Guo, P. (2020) A new species of the genus Achalinus (Squamata: Xenodermatidae) from southeastern Yunnan Province, China. Zootaxa, 4860 (1), 116 - 128. https: // doi. org / 10.11646 / zootaxa. 4860.1.6","Hu, S. Q. & Zhao, E. M. (1966) Three new species of reptiles from Szechwan. Acta Zootaxon Sinica, 3 (2), 158 - 164. [In Chinese with English abstract]","Luu, V. Q., Ziegler, T., Van Ha, N., Van Lo, O., Hoang, T. T., Ngo, H. T., Le, M. D., Tran, D. H. & Nguyen, T. Q. (2020) A new species of Achalinus (Squamata: Xenodermidae) from Trang An Landscape Complex, Ninh Binh Province, Vietnam. Zootaxa, 4877 (1), 174 - 184. https: // doi. org / 10.11646 / zootaxa. 4877.1.8","Wang, J., Li, Y., Zeng, Z. C., Lyu, Z. T., Sung, Y. H., Li, Y. Y., Lin, C. Y. & Wang, Y. Y. (2019) A new species of the genus Achalinus from southwestern Guangdong Province, China (Squamata: Xenodermatidae). Zootaxa, 4674 (4), 471 - 481. https: // doi. org / 10.11646 / zootaxa. 4674.4.6","Qi, S., Shi, J. S., Ma, Y. B., Gao, Y. F., Bu, S. H., Grismer, L. L., Li, P. P. & Wang, Y. Y. (2021) A sheep in wolf's clothing: Elaphe xiphodonta sp. nov. (Squamata, Colubridae) and its possible mimicry to Protobothrops jerdonii. ZooKeys 1048, 23 - 47. https: // doi. org / 10.3897 / zookeys. 1048.65650"]}

Herpetoreas burbrinki Guo, Zhu, Liu, Zhang, Li, Huang & Pyron, 2014 Description of female ( ANU20210006); Figs 1–3. Body elongated and thin, cylindrical; head elongate, rather narrow, distinct from the neck; nostril lateral, round, piercing in middle of divided nasal; eye large, pupil round; tail cylindrical, long and tapering; dorsal scale rows 19-19-17, all distinctly keeled; scales on posterior part of head and temporal region keeled; reduction from 19 to 17 dorsal scale rows occurs at the level of the 3 rd and 4 th dorsal scale rows and between the 105 th and 107 th ventrals; 2 preventrals and ventrals 169; paired subcaudals 94; anal divided. The scale rows reduction formula on the tail: Rostral broad, approximately 1.5 times as wide as high, curved onto the dorsal rostral surface; nasal rectangular, longer than high; internasals subtriangular, slightly longer than wide, anteriorly narrowed but truncated, with the fore width about 0.5 times as great as the hind width; prefrontals subrectangular, barely longer than internasal, reaching loreal; frontal subpentagonal, 1.8 times as long as wide, flatted front, pointed backwards, two long sides approximately parallel; parietals in contact for a length of 0.9 times the frontal length; supraoculars broadly in contact with prefrontal, with supraocular two-thirds as broad as the frontal; loreals 2/2, small, rectangular, in broad contact with nasal; preocular 1/1; postoculars 3/3, the upper one larger than two lower ones; temporals 2+2 on both sides; supralabials 9/8, the 3 rd to 5 th bordering the orbit, the left 6 th to 8 th occupying the same position as the right 6 th and 7 th; infralabials 10/10, first five contacting anterior chin shields; nostrils lateral, nasals are undivided; two pairs of chin shields. Maxillary teeth are 21 in continuous series, last 15 enlarged. Morphometric measurements are as follows: SVL: 462 mm; TaL: 184 mm; TL: 646 mm; Tal/TL: 0.285; LsI: 1.6 mm; LsP: 1.7 mm; HW: 9.2 mm; HL: 16.5 mm; HH: 6.1 mm; DE: 5.5 mm; EL: 3.8 mm; EW: 3.8 mm; RH: 1.1 mm; RW: 2.9 mm; MH: 0.9 mm; MW: 1.8 mm. Coloration in life (Figs 1–2). Dorsal ground color grey-olive green with most scales densely speckled with dark spots, turning into reddish on its posterior portion of the body. Lateral dorsum ornate with an alternating brickred and white stripe on DSR 5-6, and caudal dorsum with five regular black longitudinal stripes. Ventral scales tangerine (the prior part coloration lighter than the posterior part), with a row of well-defined dark spots at their outer border, extending to the tip of the tail. Dorsal surface of head grey-olive green with irregular dark stripes, turning into blackish-brown on the posterior part of the body. Paler on the snout and sides. Two small, faint off-white spots on the parietals and a faint off-white sagittal line just behind the parietals. Paired, pale ivory cream occipital spots, from the uppermost Po, to the 9 th (left) and 8 th (right) SL bent upwards and distinctly connected with the thin pale brown or canary dorsolateral stripe of the neck (“Y” shaped). SL off-white; each of the first five having a black speckle with brick red rim; in the middle of 6 th having a brick red speckle mixed with small black spots (“blusher”); paired, black brown lateral spots, from the middle Po, to the 7 th to 9 th (left) and 7 th to 8 th (right) SL, extending backwards more than one head length. The head ventrally off-white. Variation. Where this specimen differs from the holotype (Guo et al. 2014), features of the holotype follow in brackets. An adult female ANU20210006 (holotype an adult male YBU071128); preocular 1/1 (2/1); temporals 2+2/2+2 (3+2/3+2); supralabials 9/8 (8/8); ventrals 169 (172); subcaudals 94 pairs (96) (Figs 1–2). Distribution. The new specimen was collected in a broad-leaved evergreen forest (28.7141°N, 96.7820°E, elevation 1938 m a. s. l.) approximately ten kilometers from the type locality (Guo et al. (2014) speculated that the species might occur in northern Burma and/or extreme eastern India., Published as part of Peng, Lifang, Huang, Song, Burbrink, Frank T., Zhang, Yong, Guo, Peng & Wu, Hailong, 2021, Morphological description of a new specimen of Herpetoreas burbrinki Guo et al 2014 (Serpentes: Colubridae), pp. 433-439 in Zootaxa 5039 (3) on pages 434-437, DOI: 10.11646/zootaxa.5039.3.8, http://zenodo.org/record/5511564, {"references":["Guo, P., Zhu, F., Liu, Q., Zhang, L., Li, J. X., Huang, Y. Y. & Pyron, R. A. (2014) A taxonomic revision of the Asian keelback snakes, genus Amphiesma (Serpentes: Colubridae: Natricinae), with description of a new species. Zootaxa, 3873 (4), 425 - 440. https: // doi. org / 10.11646 / zootaxa. 3873.4.5"]}

Weinell, Jeffrey L, Barley, Anthony J, Siler, Cameron D, Orlov, Nikolai L, Ananjeva, Natalia B, Oaks, Jamie R, Burbrink, Frank T, Brown, Rafe M (2021): Phylogenetic relationships and biogeographic range evolution in cat-eyed snakes, Boiga (Serpentes: Colubridae). Zoological Journal of the Linnean Society 192 (1): 169-184, DOI: 10.1093/zoolinnean/zlaa090, URL: https://academic.oup.com/zoolinnean/article/192/1/169/5892978

Madagascarophis lolo, new species urn:lsid:zoobank.org:act:E94496B3-AC04-4FE2-AFA6- 8CC7DBE3A63A Figures 1�� 2�� 3�� 4�� 5; Table 1 Holotype.��� AMNH 176422 (RAX 12626), adult male (Fig. 1), Madagascar, Antsiranana Province, Diana Region, Ankarana National Park, ~ 4 km northwest of the village of Mahamasina, tsingy karst trail, 102 m elevation, 49.11507��E, 12.94210��S, 9 February 2014, 1930 hours, B. Randriamahatantsoa, C. Raxworthy, S. Ruane. Diagnosis.��� A new species of Madagascarophis than can be diagnosed from its congeners by the following combination of characters: an overall gray body color with a black vertebral stripe and alternating light gray blotches down the dorsum, 25 scale rows at midbody, 189 ventral scales and 56 divided subcaudal scales, with extended contact of the posterior inframaxillary scales. Madagascarophis lolo �� new species, differs from all other species of Madagascarophis having a gray body color combined with an alternating pattern of pale gray blotches along the vertebral column and the presence of black scales on the vertebral row scales, giving the appearance of a thin black dorsal stripe (Figs. 1, 3, 4). This overall combination of color and pattern is unique among species of Madagascarophis (see Domergue, 1987:fig. 4 for comparison). Madagascarophis lolo �� new species, differs from all other species with the exception of M. fuchsi by having extended contact of the posterior inframaxillary scales (Fig. 5). We note the specimen of M. lolo �� new species, has a single gular scale that infringes on the posterior end of the posterior inframaxillaries. However, with the exception of M. fuchsi, the posterior inframaxillary contact of M. lolo �� new species, is still much greater than for the other species of Madagascarophis (Fig. 5; see Glaw et al., 2013a for additional examples). Madagascarophis lolo �� new species, differs from M. fuchsi by having a lower number of infralabial scales (10 M. lolo vs. 12��� 13 M. fuchsi) and a higher ventral scale count (171���174 M. fuchsi vs. 1 8 9 M. lolo). However, this 1 5 ventral scale difference falls within the intraspecific range of other species (e.g., 35 ventral scales in M. meridionalis). It differs from the other species of Madagascarophis except M. colubrinus by having a lower ventral scale count (189 M. lolo): 183���209 in M. colubrinus, 205���224 in M. ocellatus, and 197���232 in M. meridionalis. A general difference between M. lolo �� new species, and most other Madagascarophis is the dorsal scale count at midbody. Madagascarophis lolo �� new species, has 25 dorsal scale rows as does M. fuchsi, in contrast to the 27���29 typically found in M. colubrinus (rarely 25, and not syntopically), 29���33 in M. meridionalis, and 29���31 in M. ocellatus (Glaw and Vences, 2007; Glaw et al., 2013a). It also differs genetically from all other species in the genus, e.g., M. lolo vs. M. fuchsi, COI uncorrected pairwise distance =9.6% (Table 1). For specimens not examined here (Appendix 1), additional data were used from Domergue (1987) and Glaw et al. (2013a) for the diagnosis. Description.��� Adult male in excellent state of preservation, tail complete, short ventral slit midbody for DNA tissue sample, lower body slit for assessing gonad development (fully formed mature testes; 10 mm length, 2 mm width). Snout���vent length 426 mm, tail length 65 mm, tail short (13% of total body length). Head length 20 mm, width 12 mm. Head distinct from neck. Eyes large, 3 mm horizontal diameter, pupil vertically elliptical. Supralabials 8, not in contact with the eye. Infralabials 10, first pair in contact behind mental, infralabials 1���5 in contact with inframaxillaries. Rostral broader than high, 3 mm wide/1.5 mm high, visible from above. Nasal divided below nostril, in contact with 1st and 2nd supralabials. Single loreal, in contact with nasal, preoculars, prefrontal, and supralabials 2 and 3. Circumoculars 9, 1 supraocular, 2 preoculars, 3 suboculars, and 3 postoculars. Temporals 4 + 4/4 + 5. Dorsal surface of head includes pair of internasals (width 1.7 mm/length of suture 2 mm), pair of prefrontals (width 2 mm/length of suture 2.1 mm), pair of supraoculars (width 2.8 mm/length 4.8 mm), frontal longer than wide (length 5.7 mm/anterior width 2.9 mm), pair of parietals (length of suture 4.6 mm). Two pairs of inframaxillaries (anterior inframaxillary length 4.9 mm, posterior inframaxillary length 3.1 mm), posterior inframaxillaries substantially in contact with each other excepting small gular scale at posterior end (Fig. 5). Dorsal scale rows 23-25-19 at 10th ventral from anterior, midbody, and 10th ventral anterior to cloaca. Coloration and pattern.��� Dorsal ground color gray in life, alternating lighter gray blotches/squares alongside vertebral column, many vertebral row scales black in coloration, giving general appearance of black dorsal line interrupted occasionally by gray scales (Fig. 1). At roughly the posterior 1/3 of the body, dorsal scale rows 7 and 8 occasionally black, giving spotted appearance in lateral view. Overall coloration pale gray in preservation. Color of iris in life gray/silver with gold flecking (Fig. 1), opaque gray in preservation. Dorsum of head, including rostral, internasals, prefrontals, frontal, supraoculars, and parietals gray. Supralabials 4���8 with gray and white mottling, infralabials with gray and white mottling. Slightly darker diffuse brown line runs from the posterior of eye to posterior margin of mouth. Tail gray with black mottling, darker and more contrasting compared to body, with slight flush of pale orange towards tail tip. Ventral scales cream with no pattern anteriorly, small amounts of gray flecking on ventral scales beginning at the posterior 1/3 of the body, continuing and increasing in intensity onto the subcaudal scales. Natural history.��� Similar to other species of Madagascarophis, M. lolo appears to be crepuscular/nocturnal; the specimen was found active on the ground at 1930 hours on tsingy karst rocks, in an exposed area with low scrub habitat. This is very similar to what has been described for M. fuchsi (Glaw et al., 2013a) �� and our own observation of the M. fuchsi sample included here, which we found outside a small cave in the karst system of the Montagne des Fran��ais massif, approximately 7 0 km away. By contrast, the other species of Madagascarophis found at Ankarana, M. colubrinus, was common in canyon forests and surrounding relict forests, as well as in anthropogenically disturbed habitat. We suspect the reason that M. lolo has gone undetected for so long at Ankarana is that the exposed tsingy plateau has been poorly surveyed at night in previous expeditions due to problems of gaining safe access to these areas. Madagascarophis lolo may be endemic to the karst habitats of Ankarana, and possibly Analamerana, which is the closest karst system to the east. Etymology.��� The species name, lolo, is taken from the Malagasy word for ghost; it is a noun in apposition to the genus name. This name refers to 1) the ghostly pale gray color of the holotype, and 2) that M. lolo has eluded discovery for so long at Ankarana, arguably one of the better surveyed sites in Madagascar. DISCUSSION High levels of microendemicity may be common for certain regions in Madagascar (Brown et al., 2016), such as Ankarana, where several reptile genera have endemic representatives (e.g., Alluaudina, Brookesia, Lygodactylus; Glaw and Vences, 2007). The new species of Madagascarophis described here is, as far as we currently know, restricted to the tsingy habitat of Ankarana. It seems unlikely that this snake has never been recorded previously, given that this particular area of Ankarana is one of the most accessible areas in the national park and a popular ecotourist destination in Madagascar. However, the trail where we collected this snake was only created in the last ten years and traverses a previously inaccessible area of exposed tsingy plateau which is otherwise difficult to access, especially at night. Because this new snake is found in a national park, and its habitat is naturally well-protected from anthropogenic degradation, we do not consider this species to be vulnerable to extinction. However more survey work is needed to establish the population size and distribution limits of this snake. Both the morphological and genetic results indicate a close/sister taxa relationship of M. lolo with M. fuchsi (Fig. 4). With just a single specimen, more individuals are needed to describe the intraspecific variation of M. lolo, which is also true for the recently described M. fuchsi, known from only four specimens. There is the possibility that these two sister taxa represent populations of the same species; however, the genetic differentiation between M. lolo and M. fuchsi (e.g., COI = 9.6%; Table 1) is higher than or similar to the mtDNA divergence found among many species of snakes, e.g., Lachesis (Zamudio and Greene, 1997), Naja (Slowinski and W��ster, 2000), and Pantherophis (Burbrink et al., 2000), and is beyond what is found within other species of Madagascarophis, including the highly variable M. colubrinus (Table 1; Nagy et al., 2007). Madagascarophis lolo can also be readily identified from all other species in the genus, including M. fuchsi, based on morphology alone (see Key to the species of Madagascarophis). Our coalescent species delimitation analyses also indicate M. lolo is a distinct species, although this is best supported under small ancestral population sizes and shallow divergences (Pp = 100%). This scenario may be the most realistic for the Madagascarophis, where populations of the range-restricted M. lolo as well as M. fuchsi are likely small. The new species described here is unique with respect to coloration. Although species of Madagascarophis (with the exception of M. ocellatus) have extremely variable intraspecific color patterns, we have not observed any species or individuals with the same coloration seen in the specimen of M. lolo; it possesses a pattern that appears to be well matched to the tsingy rock habitat with varying and alternating shades of gray (Figs. 1, 2). By contrast, no other species of Madagascarophis has a predominantly gray dorsal ground coloration���they are either brown, blackish brown, orange, or yellowish brown. Although it is beyond the scope of this study to validate the status of previously described subspecies of Madagascarophis colubrinus, we discuss them here to avoid problems with synonymy with respect to M. lolo. In the most complete examination of the genus, Domergue (1987) recognized five subspecies of M. colubrinus and an additional full species, M. citrinus (as well as M. meridionalis and M. ocellatus). More recent work on these snakes (Nagy et al., 2007; Glaw et al., 2013a) proposed: 1) M. meridionalis and M. ocellatus remain distinct species; 2) M. c. occidentalis is a junior synonym of M. c. colubrinus; 3) M. c. insularis is a junior synonym of M. citrinus, but M. citrinus is a subspecies of M. colubrinus; 4) There is some genetic structure and corresponding morphological variation indicating that M. c. septentrionalis and M. c. citrinus are distinct from the nominant M. c. colubrinus; and 5) M. fuchsi is a distinct species that occurs sympatrically with M. colubrinus at Montagne des Fran��ais. Importantly, in our genetic analyses and in the aforementioned studies, all of the subspecies of M. colubrinus form a separate and distinct M. colubrinus clade, which is the sister taxon to M. meridionalis. Our study, like Glaw et al. (2013a), finds M. fuchsi falling outside the M. colubrinus + M. meridionalis clade, with the addition that M. lolo is the sister taxon to M. fuchsi. Therefore, all taxa within M. colubrinus are distinct from M. lolo and M. fuchsi. One subspecies, M. c. pastoriensis�� has not been included in any molecular phylogenetic studies (due to a lack of genetic material); however, this taxon is restricted to the Antananarivo region in central Madagascar and is characterized by having a nearly black body and yellow eyes (Domergue, 1987), which does not correspond to M. lolo. The other Madagascarophis taxon also found at Ankarana, M. c. septentrionalis�� is distinct from M. lolo in that it does not typically have 25 dorsal scale rows (rather 27, or 29 from sites further north) or contact of the posterior inframaxillaries (see Glaw et al., 2013a). We included M. colubrinus (which would correspond to M. c. septentrionalis) from the collecting locality of M. lolo in our genetic analyses and demonstrate that these are not sister taxa. This study is the first to include genetic data for M. ocellatus and all other described species of Madagascarophis in a coalescent-based species tree (Fig. 4). This tree was well supported at almost all nodes (Pp �� 0.99), and importantly, it provides information on the sister taxa relationship of the two most recently described species, M. lolo and M. fuchsi. Each of these species occurs sympatrically with the widespread M. colubrinus (which is the sister taxon to the more southerly distributed M. meridionalis), but our tree indicates that no sister species of Madagascarophis occur sympatrically. These findings are similar to other studies that suggest that recently diverged sister taxa are typically allopatric and that similarity in niche may limit sympatry (e.g., Peterson, 1999; McCormack et al., 2 0 10; Pigot and Tobias, 2 0 1 3). The possible karst specialization of M. lolo and M. fuchsi may allow it to occur in the same area with the more distantly related generalist M. colubrinus. The exception to the generally high support values across the species tree is the placement of M. ocellatus as the sister taxon to M. colubrinus + M. meridionalis (Pp = 0.62). Unlike the more widespread M. colubrinus and M. meridionalis, M. ocellatus is poorly known and is found only in the dry regions of southwestern Madagascar. Our results suggest M. ocellatus is the sister taxon to M. colubrinus + M. meridionalis (Fig. 4), though we expect future studies using larger genetic datasets may be able to provide more robust support for the phylogenetic placement of M. ocellatus. Finally, this study demonstrates that snake species new to science are likely waiting to be discovered in Madagascar, even among commonly encountered taxa, and we expect there are still high numbers of endemic, undescribed squamates in Madagascar. As many of the currently recognized squamate species in Madagascar have very large ranges (e.g., Geckolepis maculata, Zonosaurus madagascariensis, Phelsuma lineata; Glaw and Vences, 200 7), phylogeographic studies using modern genomic techniques in an integrative context with morphology and ecology will likely discover additional taxa. Future work in Madagascar and other tropical regions worldwide should focus not only on the discovery of obvious, morphologically differentiated species, but also consider widespread taxa with potential cryptic diversity as well., Published as part of Sara Ruane, Frank T. Burbrink, Bernard Randriamahatantsoa & Christopher J. Raxworthy, 2016, The Cat-eyed Snakes of Madagascar: Phylogeny and Description of a New Species of Madagascarophis (Serpentes: Lamprophiidae) from the Tsingy of Ankarana, pp. 711-721 in Copeia 104 (3) on pages 716-719, DOI: 10.1643/Ch-15-346, http://zenodo.org/record/269699, {"references":["Domergue, C. A. 1987. Notes sur les serpents de la region malgache. VII. Revision du genre Madagascarophis Mertens, 1952. Bulletin du Museum national d'Histoire Naturelle, Paris 9: 455 - 489.","Glaw, F., C. Kucharzewski, J. Koehler, M. Vences, and Z. T. Nagy. 2013 a. Resolving an enigma by integrative taxonomy: Madagascarophis fuchsi (Serpentes: Lamprophiidae), a new opisthoglyphous and microendemic snake from northern Madagascar. Zootaxa 3630: 317 - 332.","Glaw, F., and M. Vences. 2007. Amphibians and Reptiles of Madagascar. Third edition. Verlag, Cologne, Germany.","Brown, J. L., N. Sillero, F. Glaw, P. Bora, D. R. Vieites, and M. Vences. 2016. Spatial biodiversity patterns of Madagascar's amphibians and reptiles. PLOS ONE 11: e 0144076.","Zamudio, K. R., and H. W. Greene. 1997. Phylogeography of the bushmaster (Lachesis muta: Viperidae): implications for neotropical biogeography, systematics, and conservation. Biological Journal of the Linnean Society 62: 421 - 442.","Burbrink, F. T., R. Lawson, and J. B. Slowinski. 2000. Mitochondrial DNA phylogeography of the polytypic North American rat snake (Elaphe obsoleta): a critique of the subspecies concept. Evolution 54: 2107 - 2118.","Nagy, Z. T., F. Glaw, F. Andreone, M. Wink, and M. Vences. 2007. Species boundaries in Malagasy snakes of the genus Madagascarophis (Serpentes: Colubridae sensu lato) assessed by nuclear and mitochondrial markers. Organisms Diversity & Evolution 7: 241 - 251.","Peterson, A. T. 1999. Conservatism of ecological niches in evolutionary time. Science 285: 1265 - 1267."]}

KEY TO THE SPECIES OF MADAGASCAROPHIS Information is based on samples in Appendix 1 and Glaw et al. (2013). 1a. Dorsal pattern consists of distinct brown/black ocel- lae, eye color red _ _ _ _ _ _ _ _ _ _ MadagascarophIs ocellatus 1b. Dorsal pattern consists of anything other than brown/black ocellae, eye color not red _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 2 2a. Extended contact of the posterior inframaxillary (genial) scales, 25 dorsal scale rows _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 3 2b. Posterior inframaxillary scales not in contact, 25 or more dorsal scale rows _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 4 3a. Ventral scales ~ 171���174, infralabials 12 or 13, body color variable _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ MadagascarophIs fuchsI 3b. Ventral scales ~ 189, infralabials 10, gray body color with a thin black dorsal stripe _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ MadagascarophIs lolo, new species 4a. Dorsal scale rows usually �� 29, ventral scale count 187���232 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ MadagascarophIs merIdIonalIs 4b. Dorsal scale rows Ś 29, ventral scale count 180��� 209 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ MadagascarophIs colubrInus, Published as part of Sara Ruane, Frank T. Burbrink, Bernard Randriamahatantsoa & Christopher J. Raxworthy, 2016, The Cat-eyed Snakes of Madagascar: Phylogeny and Description of a New Species of Madagascarophis (Serpentes: Lamprophiidae) from the Tsingy of Ankarana, pp. 711-721 in Copeia 104 (3) on page 719, DOI: 10.1643/Ch-15-346, http://zenodo.org/record/269699, {"references":["Glaw, F., C. Kucharzewski, J. Koehler, M. Vences, and Z. T. Nagy. 2013 a. Resolving an enigma by integrative taxonomy: Madagascarophis fuchsi (Serpentes: Lamprophiidae), a new opisthoglyphous and microendemic snake from northern Madagascar. Zootaxa 3630: 317 - 332."]}

Family Charinidae Gray 1849 Type genus. Charina Gray 1849 Genus content. Subfamily Charininae Gray 1849 (Charina, Lichanura), subfamily Ungaliophiinae McDowell 1987 (Exiliboa, Ungaliophis) Diagnosis. Charinids can be distinguished from all other similar or related taxa by the following combination of characters: in all species, a distinct lateral muscular bundle (M.) in the jaw known as the adductor mandibulae externus medialis pars anterior (M. aem 1 of Zaher 1994), with loss of the left lung and presence of a tracheal lung in Exiliboa and Ungaliophis (Ungaliophiinae), and in Charina and Lichanura (Charininae), an M. aem 1 comprised of elongated and undifferentiated fibers, as well as lack of a postorbital, anterior end of ectopterygoid narrow and pointed, crista circumfenestris small or absent between the fenestra pseudorotunda and foramen of the vagus cranial nerve, and coronoid absent on the prearticular portion of the compound bone (see Kluge 1993; Zaher 1994). Phylogenetic definition. Includes the MRCA of included Charina and Lichanura (Charininae) and Exiliboa and Ungaliophis (Ungaliophiinae) and all descendants thereof, and all species more closely related to Cha. bottae than to Boa constrictor (stem of Charinidae), all species more closely related to Cha. bottae than to U. continentalis (stem of Charininae), and all species more closely related to U. continentalis than to Cha. bottae (stem of Ungaliophiinae). Etymology. From the Greek charieis for ���graceful��� or ���delightful.��� Distribution. North and Central America, including southern Canada, the western United States and northwestern Mexico for Charininae, and southwestern Mexico, Central America, and extreme northwestern South America for Ungaliophiinae. Remarks. This group has had a turbulent taxonomic history, as many morphological characters are poorly known and apparently exhibit strong convergence (see Kluge 1991; Zaher 1994; Wilcox et al. 2002). However, recent molecular results are unambiguous in supporting the monophyly of this group and uniting it with Booidea (Wilcox et al. 2002; Noonan and Chippindale 2006; Wiens et al. 2008; Pyron et al. 2013; Reynolds et al. 2014)., Published as part of Pyron, R. Alexander, Reynolds, R. Graham & Burbrink, Frank T., 2014, A Taxonomic Revision of Boas (Serpentes: Boidae), pp. 249-260 in Zootaxa 3846 (2) on page 254, DOI: 10.11646/zootaxa.3846.2.5, http://zenodo.org/record/230379, {"references":["Gray, J. E. (1849) Catalogue of the Specimens of Snakes in the Collection of the British Museum. Newman, London, 125 pp.","McDowell, S. B. (1987) Systematics. In: Siegel, R. A., Collins, J. T. & Novak, S. S. (Eds.), Snakes: Ecology and Evolutionary Biology. Macmillan, Toronto, pp. 3 - 50.","Zaher, H. (1994) Les Tropidopheoidea (Serpentes; Alethinophidea) sont-ils reellement monophyletiques? Arguments en faveur de leur polyphyleetisme. Comptes Rendus de L Academie Des Sciences, Serie III - Sciences de La Vie - Life Sciences, 317 (5), 471 - 478.","Kluge, A. G. (1993) Calabaria and the phylogeny of erycine snakes. Zoological Journal of the Linnaean Society, 107, 293 - 351.","Kluge, A. G. (1991) Boine snake phylogeny and research cycles. Miscellaneous Publications of the Museum of Zoology of the University of Michigan, 178, 1 - 58.","Wilcox, T. P., Zwickl, D. J., Heath, T. A. & Hillis, D. M. (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. http: // dx. doi. org / 10.1016 / s 1055 - 7903 (02) 00244 - 0","Noonan, B. P. & Chippindale, P. T. (2006) Dispersal and vicariance: the complex evolutionary history of boid snakes. Molecular Phylogenetics and Evolution, 40, 347 - 358. http: // dx. doi. org / 10.1016 / j. ympev. 2006.03.010","Pyron, R. A., Burbrink, F. T. & Wiens, J. J. (2013) A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology, 13, 93. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 13 - 93","Reynolds, R. G., Niemiler, M. L. & Revell, L. J. (2014) Toward a Tree-of-Life for the boas and pythons: multi locus species-level phylogeny with unprecedented taxon sampling. Molecular Phylogenetics and Evolution, 71, 201 - 213. http: // dx. doi. org / 10.1016 / j. ympev. 2013.11.011"]}

Gonyosoma Wagler Gonyosoma Wagler, 1828. Type species Gonyosoma oxycephalum (Boie) Gonyophis Boulenger, 1891. Type species Gonyophis margaritatus (Peters), by monotypy Syn.n. Rhadinophis Vogt, 1922. Type species Rhadinophis frenatus (Gray) Syn. n. Rhynchophis Mocquard, 1897. Type species Rhynchophis boulengeri Mocquard, by monotypy Syn.n. Diagnosis. The genus Gonyosoma is comprised of six currently recognized species (Table 3), which form a monophyletic group based on Maximum Likelihood and Bayesian inference using six genes. Morphological characters for this genus are listed as follows: body size from medium to large (80 cm to 240 cm); body slender, compressed, head elongated and distinct from neck; relatively large eyes with round pupil; tail relatively long (20 % to 30 % of total length) and slender; various color patterns represented but generally displaying some green, yellowish or tan ground color; single preocular; postoculars two; mid-body dorsal scales rows 19 or 23; mid-dorsal scales smooth or weakly keeled; 186���257 ventral scales with well-developed lateral keels; 91���149 paired subcaudals; precloacal plate usually divided (some Gonyosoma prasinum populations remain undivided); primarily arboreal or semi-arboreal and mainly found in tropical rain forest or subtropical montane forest habitats; distribution extends from Southern China, Southeast Asia and South Asia; diurnal; oviparous (Schulz 1996; Zhao 2006; Das 2010; Schulz & Gumprecht 2013). Finally, our reconstructed phylogeny indicates that the two tropical clades comprising the genera Gonyosoma and Coelognathus are not closely related to the Old World and New World ratsnake assemblages as typically considered in the literature (Schultz 1996; Utiger et al. 2002, Burbrink & Lawson 2007), which currently includes the New World tribe Lampropeltini and the Old World genera Oreocryptophis, Archelaphe, Euprepiophis, Orthriophis, Elaphe, Rhinechis, Zamenis, Coronella and Oocatochus. Although Gonyosoma, Coelognathus and the ratsnake radiation group all originated around the early Miocene (~ 25 ��� 20 Ma), comparison of the species richness of the three groups shows obvious asymmetries in biodiversity (Gonyosoma = 6 taxa, Coelognathus = 5 taxa, and the ratsnakes = ~ 70 taxa), implying that the tropical Asian environment has not provided more opportunities for species divergence within these two genera or extinction has been elevated relative to the temperate groups., Published as part of Chen, Xin, Grismer, L. Lee, Matsui, Masafumi, Nishikawa, Kanto & Burbrink, Frank T., 2014, The phylogenetIc posItIon and taxonomIc status of the RaInbow Tree Snake Gonyophis margaritatus (Peters, 1871) (Squamata: ColubrIdae), pp. 532-548 in Zootaxa 3881 (6) on page 537, DOI: 10.11646/zootaxa.3881.6.3, http://zenodo.org/record/231067, {"references":["Boulenger, G. A. (1891) On new or little-known Indian and Malayan reptiles and Batrachians. The Annals and Magazine of Natural History, 8, 288 - 290. http: // dx. doi. org / 10.1080 / 00222939109460437","Vogt, T. (1922) Zur Reptilien-und Amphibienfauna Sudchinas. Archiv fur Naturgeschichte, 88 (A), 135 - 146.","Mocquard, F. (1897) Notes herpetologiques. Bulletin du Museum National d'Histoire Naturelle, Paris, 3, 211 - 217. [Series. 1]","Schulz, K. D. (1996) A monograph of the colubrid snakes of the genus Elaphe Fitzinge r. Koeltz Scientific Books, Wurselen, 439 pp.","Zhao, E. (2006) Snakes of China. Anhui Science and Technology Publishing House, China, 369 pp.","Das, I. (2010) A Field Guide to the Reptiles of South-East Asia. New Holland Publishers Ltd, London, 376 pp.","Schulz, K. D. & Gumprecht, A. (2013) Undercover in the rainforest canopy: The rainbow tree snake Gonyophis margaritatu s. In: Schulz, K. D. (Ed.), Old World Ratsnakes. A collection of paper s. Koeltz Scientific Books, Koenigstein, pp. 395 - 402."]}

Pyron, R. Alexander, Reynolds, R. Graham, Burbrink, Frank T. (2014): A Taxonomic Revision of Boas (Serpentes: Boidae). Zootaxa 3846 (2): 249-260, DOI: http://dx.doi.org/10.11646/zootaxa.3846.2.5

Family Boidae Type genus. Boa Linnaeus 1758 Genus content. Boa, Chilabothrus, Corallus, Epicrates, Eunectes Diagnosis. Boids can be distinguished from all other similar or related taxa by the following combination of characters: internarial septum with large fenestra, anterior margin of the ventral lamina of the nasal indented in lateral view, anterolateral margin of horizontal lamina of nasal noticeably indented viewed dorsally, horizontal lamina of the nasal does not overlap dorsal surface of frontal, most of palatine process of maxilla occurs posteriorly within the orbit, anterior end of ectopterygoid consists of indistinct lateral and medial heads, supratemporal inclined slightly in lateral view, posterior end of supratemporal rounded but not dilated, parasphenoid wing large and without pedicellate ventral surface, dorsal margin of prearticular noticeably curved upward near attachment of adductor posterior muscle, cornua of hyobranchium discontinuous anteriorly, and shallow labial pits (see Kluge 1991). Phylogenetic definition. Includes the MRCA of included Boa, Chilabothrus, Corallus, Epicrates, and Eunectes species and all descendants thereof, and all species more closely related to Boa constrictor than to Candoia carinata or Eryx jaculus. Etymology. From the Latin boa for ���large snake,��� after an animal mentioned in the Natural History of Pliny the Elder. Distribution. New World tropics, from northern Mexico to Argentina, and the West Indies. Remarks. Several species in this group contain significant intra-specific genetic variation (Colston et al. 2013; Reynolds et al. 2014), and more species may be recognized in the future. This arrangement (family-level rank of the former boid subfamilies) provides a robust taxonomy for booid snakes. All families are strongly supported as monophyletic by both molecular and morphological data, and their nomenclature is thus unlikely to be affected in the future by the sampling of additional taxa or characters in phylogenetic analyses. The recognition of these ecomorphologically and biogeographically distinct groups as families mirrors other squamate and amphibian taxa such as Iguania (Frost & Etheridge 1989; Frost et al. 2001) and Ranoidea (Frost et al. 2006; Pyron & Wiens 2011), where large cosmopolitan families (Iguanidae and Ranidae) have been broken up into more manageable and geographically localized units. This classification should provide a stable basis for future revision and description of new species, and other comparative analyses., Published as part of Pyron, R. Alexander, Reynolds, R. Graham & Burbrink, Frank T., 2014, A Taxonomic Revision of Boas (Serpentes: Boidae), pp. 249-260 in Zootaxa 3846 (2) on page 256, DOI: 10.11646/zootaxa.3846.2.5, http://zenodo.org/record/230379, {"references":["Linnaeus, C. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii, Holmiae, 824 pp.","Kluge, A. G. (1991) Boine snake phylogeny and research cycles. Miscellaneous Publications of the Museum of Zoology of the University of Michigan, 178, 1 - 58.","Colston, T. J., Grazziotin, F. G., Shepard, D. B., Vitt, L. J., Colli, G. R., Henderson, R. W., Hedges, S. B., Bonatto, S., Zaher, H., Noonan, B. P. & Burbrink, F. T. (2013) Molecular systematics and historical biogeography of tree boas (Corallus spp.). Molecular Phylogenetics and Evolution, 66, 953 - 959.","Reynolds, R. G., Niemiler, M. L. & Revell, L. J. (2014) Toward a Tree-of-Life for the boas and pythons: multi locus species-level phylogeny with unprecedented taxon sampling. Molecular Phylogenetics and Evolution, 71, 201 - 213. http: // dx. doi. org / 10.1016 / j. ympev. 2013.11.011","Frost, D. R. & Etheridge, R. (1989) A phylogenetic analysis and taxonomy of iguanian lizards (Reptilia: Squamata). University of Kansas, Lawrence, 65 pp.","Frost, D. R., Etheridge, R. E., Janies, D. & Titus, T. A. (2001) Total evidence, sequence alignment, evolution of polychrotid lizards, and a reclassification of the Iguania (Squamata: Iguania). American Museum Novitates, 3343, 1 - 38. http: // dx. doi. org / 10.1206 / 0003 - 0082 (2001) 343 2.0. co; 2","Frost, D. R., Grant, T., Faivovich, J., Bain, R. H., Haas, A., Haddad, C. F. B., De Sa, R. O., Channing, A., Wilkinson, M., Donnellan, S. C., Raxworthy, C. J., Campbell, J. A., Blotto, B. L., Moler, P., Drewes, R. C., Nussbaum, R. A., Lynch, J. D., Green, D. M. & Wheeler, W. C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 8 - 370. http: // dx. doi. org / 10.1206 / 0003 - 0090 (2006) 297 [0001: tatol] 2.0. co; 2","Pyron, R. A. & Wiens, J. J. (2011) A large-scale phylogeny of Amphibia including over 2800 species, and a revised classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution, 61, 543 - 583. http: // dx. doi. org / 10.1016 / j. ympev. 2011.06.012"]}

Family Erycidae Bonaparte 1831 Type genus. Eryx Daudin 1803 Genus content. Eryx Diagnosis. Erycids can be distinguished from all other similar or related taxa by the following combination of characters: transverse process of the premaxilla long, internarial septum of the premaxilla absent, vomerine process of the premaxilla long and narrow, anterior one-third to one-half of the ventral lamina of the nasal decreases anteriorly or is absent, anterior end of the horizontal lamina of the nasal expands slightly to markedly viewed dorsally, anterolateral edge of the horizontal lamina of the nasal bulges outward viewed dorsolaterally, lateroposterior margin of the horizontal lamina of the nasal adjacent to the prefrontal oriented vertically as a large wall, dorsal lappet of the prefrontal approximately straight on the dorsolateral margin contacting the frontal and nasal, anterolateral and medial laminae of the prefrontal form a gradually rounded corner, frontonasal facet where ventral lamina of the nasal contacts the frontal lies between and below the olfactory canal with long horizontalventral portion, facet present on the preorbital process of the frontal, anterolateral corner of the frontal projects anteriorly and partially exposes the preorbital process, floor of the optic foramen formed by the parietal, postorbital process of the parietal large, midsaggital crest markedly developed on the anterior portion of the parietal, choanal process of the palatine tiny and discontinuous, maxillary process of the palatine narrow, medioventral margin of the posterior third of the pterygoid folded into deep groove, parasphenoid shallow anteriorly deepening posteriorly with groove absent anteriorly and present posteriorly, interparietal area of the parasphenoid of modest width with parallel sides, anteriorly projecting process present on the neural spine, distal tip of the posterior diapophysis of caudal vertebrae oriented nearly vertically into a thin and occasionally convoluted blade, distal tip of anterior diapophysis of caudal vertebrae elaborated into a horizontal blade originating from centrum, transverse process of caudal vertebrae forms a large and rounded anteroposteriorly oriented blade, caudal haemopophysis flattened terminally and curved towards opposite-side counterpart but widely separated therefrom, first infralabial scale small and separated from counterpart by one or more scales, dorsal head scales small and usually asymmetrically arranged, and 34 diploid chromosomes (see Kluge 1993). Phylogenetic definition. Includes the MRCA of the included Eryx species and all descendants thereof, and all species more closely related to E. jaculus than to Boa constrictor. Etymology. Named, apparently without special significance, after a son of Aphrodite, king of the mountain city Eryx in Sicily, who was defeated by Heracles in a boxing match. Distribution. Southeastern Europe, northern Africa, Middle East, and southwestern Asia. Remarks. This taxon has been recognized as a distinct family in the past, as well as by some current authors (see Wallach et al. 2014)., Published as part of Pyron, R. Alexander, Reynolds, R. Graham & Burbrink, Frank T., 2014, A Taxonomic Revision of Boas (Serpentes: Boidae), pp. 249-260 in Zootaxa 3846 (2) on page 255, DOI: 10.11646/zootaxa.3846.2.5, http://zenodo.org/record/230379, {"references":["Bonaparte, C. L. J. L. (1831) Saggio di una distribuzione metodia degli animali vertebrati. Giornale Arcadico di Scienze Lettere ed Arti, 49, 3 - 77. http: // dx. doi. org / 10.5962 / bhl. title. 48624","Daudin, F. M. (1803) Histoire Naturelle, Generale et Particuliere des Reptiles. Vol. VII. Dufart, Paris, 436 pp.","Kluge, A. G. (1993) Calabaria and the phylogeny of erycine snakes. Zoological Journal of the Linnaean Society, 107, 293 - 351.","Wallach, V., Williams, K. L. & Boundy, J. (2014) Snakes of the World: a Catalogue of Living and Extinct Species. CRC Press, Boca Raton, 1209 pp."]}

Burbrink, Frank T., Guiher, Timothy J. (2015): Considering gene flow when using coalescent methods to delimit lineages of North American pitvipers of the genus Agkistrodon. Zoological Journal of the Linnean Society 173 (2): 505-526, DOI: 10.1111/zoj.12211, URL: http://dx.doi.org/10.1111/zoj.12211

Lampropeltis californiae (Blainville 1835) (Figs. 1���3) California Kingsnake Holotype: Unknown, collected by M. Botta. Type Locality: ���California��� (Blainville 1835). Restricted to Fresno, California by Schmidt (1953). Etymology: Specific epithet refers to the type locality. Synonymy: This species is essentially co-terminous with the previously recognized subspecies L. g. californiae, and L. g. nigrita. Diagnosis: The California Kingsnake (L. californiae) is a medium- to large-bodied constrictor, with a mean adult size range of 76���122cm, and a maximum size of ~ 200cm (Stebbins 2003). In both sexes, ventral scale counts range from 213���255, with 46���63 subcaudal scales in males and 44���57 in females (Blaney 1977). Scales are smooth and anal plate single, with 23���25 dorsal scale rows at midbody. The California Kingsnake exhibits one of the broadest ranges of any kingsnake species, occupying most habitats west of the continental divide at the Cochise filter barrier (Fig. 2). The species ranges from Oregon in the north, through the Great Basin in Nevada and Utah, east to extreme southwestern Colorado, south through the majority of California, Arizona and the Mexican states of Sonora, Baja California Norte, and Baja Sur (Fig. 2). The California Kingsnake can be distinguished from other species on the basis of color pattern, possibly the most distinct of the group. Throughout the majority of their range, California Kingsnakes exhibit a black or dark brown ground color, with 21���44 broad crossbands of white or light yellow, which typically widen laterally. Along the Pacific coast from Los Angeles to San Diego counties, individuals can be found possessing a black or dark brown ground color and a single thin, white dorsal stripe beginning at the neck and continuing to to the tail (Fig. 3). Finally, populations in the Mexican states of Sonora and Sinaloa may exhibit considerable ontogenetic darkening, with adults, and occasionally subadults and even juveniles turning jet black, with almost no trace of pattern (Blaney 1977; Stebbins 2003). To which species the Sonora populations belong is unclear; while we group them with L. californiae on the basis of geography, some authors have suggested that they resemble and hybridize with both L. g. ���splendida��� and L. g. ���californiae��� (Blanchard 1921; Blaney 1977)., Published as part of Pyron, Alexander & Burbrink, Frank T., 2009, Systematics of the Common Kingsnake (Lampropeltis getula; Serpentes: Colubridae) and the burden of heritage in taxonomy, pp. 22-32 in Zootaxa 2241 on pages 27-29, DOI: 10.5281/zenodo.190597, {"references":["Blainville, H. D. (1835) Description de quelques especes de reptiles de la Californie. Nouvelles Annales Museum National D'Histoire Naturelle Paris, 4,233 - 296.","Schmidt, K. P. (1953) A check list of North American. amphibians and reptiles. 6 th ed. University Chicago Press, Chicago, IL, 280 pp.","Stebbins, R. C. (2003) A Field guide to western reptiles and amphibians. 3 rd ed. Houghton Mifflin, Boston, 572 pp.","Blaney, R. M. (1977) Systematics of the Common Kingsnake, Lampropeltis getulus (Linnaeus). Tulane Studies in Zoology and Botany, 19, 47 - 104.","Blanchard, F. N. (1921) A revision of the king snakes, genus Lampropeltis. Bulletin of the United States National Museum, 114, 1 - 260."]}

Lampropeltis nigra (Yarrow 1882) (Figs. 1���3) Black Kingsnake Holotype: USNM 12149, collected by Robert Ridgway. Type Locality: Wheatland, Knox Co. Indiana. Etymology: Specific epithet refers to the predominantly black dorsal coloration of many specimens. Synonymy: This species comprises the previously recognized subspecies L. g. nigra and L. g. holbrooki (part). Diagnosis: The Black Kingsnake (L. nigra) is a large- to medium-bodied constrictor with an average adult size of 90���122cm, with larger individuals attaining maximum lengths of 147���183cm (Conant & Collins 1998). Scales are smooth, anal plate single, and individuals typically exhibit 19���25 scale rows at midbody. Ventral scale counts range from 197���222 in both sexes (fewer in the north), while subcaudals range from 45��� 59 in males and 37���51 in females (Blaney 1977). The Black Kingsnake can be distinguished from other species in the genus based on a combination of geography and color pattern. The Black Kingsnake ranges from southern Illinois to the Gulf coast along the Mississippi River, and east to the Appalachian mountain and the Alabama River drainage in south Alabama (Fig. 2). Black Kingsnakes all exhibit a black ground color, typically with a black-and-white checkered venter, and rarely faint traces of dorsal crossbands (Blanchard 1921; Blaney 1977; Conant & Collins 1998). Each dorsal scale is punctuated by a yellow or white speckle near the center of the scale; this is strongest in the southern portion of their range and fades considerably in the north, where many adults may be almost completely black (Conant & Collins 1998; Fig. 3). The Black Kingsnake can be distinguished from the morphologically similar Central lineage on the basis of geography, as the Black Kingsnake is only found east of the Mississippi River (Fig. 2)., Published as part of Pyron, Alexander & Burbrink, Frank T., 2009, Systematics of the Common Kingsnake (Lampropeltis getula; Serpentes: Colubridae) and the burden of heritage in taxonomy, pp. 22-32 in Zootaxa 2241 on page 26, DOI: 10.5281/zenodo.190597, {"references":["Yarrow, H. C. (1882) Check list of North American reptilia and batrachia. Bulletin of the United States National Museum, 24, 1 - 249.","Conant, R. & Collins, J. T. (1998) Reptiles and Amphibians of Eastern and Central North America, 3 rd ed. Houghton Mifflin, Boston, 616 pp.","Blaney, R. M. (1977) Systematics of the Common Kingsnake, Lampropeltis getulus (Linnaeus). Tulane Studies in Zoology and Botany, 19, 47 - 104.","Blanchard, F. N. (1921) A revision of the king snakes, genus Lampropeltis. Bulletin of the United States National Museum, 114, 1 - 260."]}

Lampropeltis holbrooki (Stejneger 1903) (Figs. 1���3) Speckled Kingsnake Holotype: Unknown Type Locality: ���Valley of the Mississippi��� (Holbrook 1842), restricted to Hot Springs, Arkansas (Schmidt 1953). Etymology: Specific epithet is a patronym honoring John Edwards Holbrook, a prominent American herpetologist of the 19 th century, known as the ���father of North American Herpetology���. Synonymy: This species is essentially co-terminous with populations of the previously recognized subspecies L. g. holbrooki occurring west of the Mississippi River. Diagnosis: The Speckled Kingsnake (L. holbrooki) is a medium- to large-bodied constrictor with a maximum adult size of 183cm and a mean adult range of 90���122cm (Conant & Collins 1998). Scales are smooth, anal plate single, and midbody scale rows number 19���25 (Blaney 1977). Ventral scales number 197��� 222 in both sexes, with subcaudals ranging from 46���59 in males and 37���51 in females (Blanchard 1921; Blaney 1977). The Speckled Kingsnake occurs west of the Mississippi River, from Iowa and Nebraska in the north to the Gulf Coast, and west to west-central Texas (Fig. 2). The majority of the range of L. holbrooki is characterized by the ���speckled��� pattern, which consists of a black ground color, with a white or yellow speckle in the center of each scale, and very occasionally a faint trace of dorsal crossbanding (Fig. 3). Large geographical areas harboring at least superficial morphological intermediacy between the Speckled Kingsnake and the Desert lineage in west central Texas are apparently inhabited only by the Speckled Kingsnake, suggesting that such color pattern variation may be due to phenotypic responses to ecological gradation, rather than hybridization or introgression (Pyron & Burbrink 2009 c). The precise western extent of the range of L. holbrooki is unclear, but ecological niche modeling predicts that the range extends approximately to the Pecos and Rio Grande River drainages (see Fig. 4 in Pyron & Burbrink 2009 c; Fig. 2)., Published as part of Pyron, Alexander & Burbrink, Frank T., 2009, Systematics of the Common Kingsnake (Lampropeltis getula; Serpentes: Colubridae) and the burden of heritage in taxonomy, pp. 22-32 in Zootaxa 2241 on page 26, DOI: 10.5281/zenodo.190597, {"references":["Stejneger, L. H. (1903) The reptiles of the Huachuca Mountains, Arizona. Proceedings of the United States National Museum, 25, 149 - 158.","Schmidt, K. P. (1953) A check list of North American. amphibians and reptiles. 6 th ed. University Chicago Press, Chicago, IL, 280 pp.","Conant, R. & Collins, J. T. (1998) Reptiles and Amphibians of Eastern and Central North America, 3 rd ed. Houghton Mifflin, Boston, 616 pp.","Blaney, R. M. (1977) Systematics of the Common Kingsnake, Lampropeltis getulus (Linnaeus). Tulane Studies in Zoology and Botany, 19, 47 - 104.","Blanchard, F. N. (1921) A revision of the king snakes, genus Lampropeltis. Bulletin of the United States National Museum, 114, 1 - 260.","Pyron, R. A. & Burbrink, F. T. (2009 c) Lineage diversification in a widespread species: roles for niche divergence and conservatism in the Common Kingsnake, Lampropeltis getula. Molecular Ecology, 18, 3443 - 3457."]}

Lampropeltis splendida (Baird & Girard 1853) (Figs. 1���3) Desert Kingsnake Holotype: USNM 1726, collected by Col. J.D. Graham. Type Locality: Sonora, Mexico (no further locality given). Etymology: Specific epithet refers to the ���splendid��� visage of the dorsal coloration. Synonymy: This species is essentially co-terminous with the formerly recognized subspecies L. g. splendida. Diagnosis: The Desert Kingsnake (L. splendida) is a medium- to large-bodied constrictor with an average adult size range of 90���114cm and a maximum size of 152cm (Conant & Collins 1998). Scales are smooth, anal plate single, with midbody scale rows typically numbering 23���25 (Blaney 1977). Ventral scales number 199���227 in males and 203���237 in females, while subcaudals range from 45���62 in males and 40���52 in females (Blaney 1977). The Desert Kingsnake can be distinguished from related species primarily on the basis of color pattern. The pattern of the Desert Kingsnake is characterized by a black or dark brown ground color with heavy yellow lateral and dorsolateral stippling. The remnant crossbands formed by this stippling yield a row of black or brown dorsal blotches or saddles, numbering 42���97. The head is typically black or dark brown, and the onset of the yellow dorsal patterning sometimes gives the appearance of a collar (Fig. 3; Blanchard 1921; Blaney 1977; Conant & Collins 1998). The Desert Kingsnake inhabits the Chihuahuan desert east of the Cochise Filter Barrier, from western Texas to extreme southeastern Arizona, north from central New Mexico in the Rio Grande River valley south to the south central portion of the Mexican Plateau (Fig. 2). Additionally, the ecological niche modeling results from Pyron & Burbrink (2009 c) predict an area of habitat in northcentral Arizona as suitable for L. splendida which is not predicted as suitable for the geographically adjacent California lineage (Fig. 2; Pyron & Burbrink 2009 c). While kingsnakes are known from this region of Arizona (Stebbins 2003), it is not known to which species this population belongs. The Desert Kingsnakes may hybridize with the Western lineage in a narrow area in extreme southeastern Arizona and extreme southwestern New Mexico, where haplotypes co-occur and some apparent hybrids have been found (Fig. 2; R.A. Pyron, pers. obs.), though morphological intermediacy is apparently not widespread (Conant & Collins 1998)., Published as part of Pyron, Alexander & Burbrink, Frank T., 2009, Systematics of the Common Kingsnake (Lampropeltis getula; Serpentes: Colubridae) and the burden of heritage in taxonomy, pp. 22-32 in Zootaxa 2241 on page 27, DOI: 10.5281/zenodo.190597, {"references":["Conant, R. & Collins, J. T. (1998) Reptiles and Amphibians of Eastern and Central North America, 3 rd ed. Houghton Mifflin, Boston, 616 pp.","Blaney, R. M. (1977) Systematics of the Common Kingsnake, Lampropeltis getulus (Linnaeus). Tulane Studies in Zoology and Botany, 19, 47 - 104.","Blanchard, F. N. (1921) A revision of the king snakes, genus Lampropeltis. Bulletin of the United States National Museum, 114, 1 - 260.","Pyron, R. A. & Burbrink, F. T. (2009 c) Lineage diversification in a widespread species: roles for niche divergence and conservatism in the Common Kingsnake, Lampropeltis getula. Molecular Ecology, 18, 3443 - 3457.","Stebbins, R. C. (2003) A Field guide to western reptiles and amphibians. 3 rd ed. Houghton Mifflin, Boston, 572 pp."]}

Pyron, Alexander, Burbrink, Frank T. (2009): Systematics of the Common Kingsnake (Lampropeltis getula; Serpentes: Colubridae) and the burden of heritage in taxonomy. Zootaxa 2241: 22-32, DOI: 10.5281/zenodo.190597

Lampropeltis getula (Linnaeus 1766) (Figs. 1���3) Eastern Kingsnake Holotype: Unknown. Type Locality: ���Carolina��� (Linnaeus 1766), restricted to Charleston, SC by Klauber (1948) Etymology: Specific epithet refers to the Getulian people of northern Morocco, whose tribal insignia bears a resemblance to the ���chain��� pattern of the kingsnakes of the Eastern Seaboard of the United States. Synonymy: This species comprises the previously recognized subspecies L. g getula, L. g. floridana and L. g. meansi, as well as the historically recognized subspecies L. g. goini and L. g. sticticeps. The nominate subspecies was first designated by Cope (1875). Diagnosis: The Eastern Kingsnake (L. getula) is a medium- to large-bodied constrictor, the largest in the genus Lampropeltis with a maximum total length of 208.3cm, though the average adult size range is 90��� 122cm (Conant & Collins 1998). Scales are smooth, anal plate single, and individuals typically exhibit 19���25 scale rows at midbody. Ventral scales number 200 to 223 in both sexes, while subcaudals number 45���58 in males and 37���55 in females (Blaney 1977). The Eastern Kingsnake ranges from New Jersey to the Florida keys in the east, and west to the western panhandle of Florida and southeastern Alabama (Fig. 2). The species L. getula can be distinguished from all other related snakes primarily on the basis of color pattern, which can be divided into two primary variants. From northern Florida to New Jersey, individuals typically exhibit a dark brown or black ground color, which is punctuated by 17���36 narrow crossbands of white, yellow, or reddish yellow (Blaney 1977), giving the appearance of a ���chain��� pattern (Fig. 3). In peninsular Florida, the bands increase in both number (22���54) and width, and the ground color lightens considerably to a light brown color with yellow stippling (Blaney 1977; Fig. 3). Isolated populations of other aberrant color pattern variants can be found in the panhandle of Florida (Krysko & Judd 2006). The shift between the Eastern Kingsnake and the Mississippi lineage is fairly abrupt, and previous authors have noted the narrow transition zone and apparent lack of morphological intermediacy in southern Alabama and central Georgia (Fig. 2, 3; Blanchard 1921; Blaney 1977; Mount 1980)., Published as part of Pyron, Alexander & Burbrink, Frank T., 2009, Systematics of the Common Kingsnake (Lampropeltis getula; Serpentes: Colubridae) and the burden of heritage in taxonomy, pp. 22-32 in Zootaxa 2241 on pages 24-25, DOI: 10.5281/zenodo.190597, {"references":["Linnaeus, C. (1766) Systema natur per regna tria natur, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio duodecima, reformata. Laurentii Salvii, Stockholm, Holmiae, 532 pp.","Klauber, L. M. (1948) Some misapplications of the Linnaean names applied to American snakes. Copeia, 1, 1 - 14.","Cope, E. D. (1875) The herpetology of Florida. Proceedings of the Academy of Natural Sciences of Philadelphia, 1875, 10 - 11.","Conant, R. & Collins, J. T. (1998) Reptiles and Amphibians of Eastern and Central North America, 3 rd ed. Houghton Mifflin, Boston, 616 pp.","Blaney, R. M. (1977) Systematics of the Common Kingsnake, Lampropeltis getulus (Linnaeus). Tulane Studies in Zoology and Botany, 19, 47 - 104.","Krysko, K. L. & Judd, W. S. (2006) Morphological systematics of kingsnakes, Lampropeltis getula complex (Serpentes: Colubridae), in the eastern United States. Zootaxa, 1193, 1 - 39.","Blanchard, F. N. (1921) A revision of the king snakes, genus Lampropeltis. Bulletin of the United States National Museum, 114, 1 - 260.","Mount, R. H. (1980) The reptiles and amphibians of Alabama. Auburn University Press, Auburn, 368 pp."]}