Description of a New Cobra (Naja Laurenti, 1768; Squamata, Elapidae) from China with Designation of a Neotype for Naja atra

Abstract

Taxonomic frameworks for medically important species such as cobras (genus Naja Laurenti, 1768; Squamata, Elapidae) are essential for the medical treatment of snake bites and accurate antivenin development. In this paper, we described the former N. kaouthia populations recorded from China as a new species and designated a neotype for N. atra-based morphological and mitochondrial phylogenetic analysis. The new species N. fuxisp. nov. was morphologically diagnosed from N. kaouthia by (1) regular single narrow crossband present on the middle and posterior parts of the dorsum (3-15, 7.9 ± 2.7, n = 32) and the dorsal surface of the tail (1-6, 4.2 ± 1.1, n = 32) of both adults and juveniles, buff-colored with dark fringes on both edges, vs. South Asian populations (n = 39) and Southeast Asian populations (n = 35) without cross bands, with irregular cross bands or multiple light-colored crossbands pairs, or densely woven lines; (2) small scales between the posterior chin shields, usually three (40%) or two (37%), rarely four (13%), or one (10%) (n = 30) vs. mostly one (81%) and rarely two (19%) (n = 28); (3) ventrals 179-205 (195.4 ± 6.7, n = 33) vs. South Asian populations 179-199 (188.7 ± 5.9, n = 12); Southeast Asian populations 168-186 (177.8 ± 4.9, n = 18). Phylogenetically, the new species forms an independent sister clade to the clade including N. atra, N. kaouthia, N. oxiana and N. sagittifera. Furthermore, the subspecies N. naja polyocellata should be resurrected and recognized as a full species, N. polyocellatacomb. nov., and the subspecies N. sumatrana miolepis should be resurrected.

Keywords: Asian cobras; Naja atra; Naja kaouthia; taxonomy.

Figure 1

Distributions of cobras in Asia. Distribution data were adjusted from IUCN https://www.iucnredlist.org (accessed on 26 October 2022).

Figure 2

Phylogenetic trees of the genus Naja inferred by Bayesian analyses (BI) based on concatenated mitochondrial gene alignment for 1028 bp cyt b and 659 bp ND4. Both bootstrap supports (BS) and Bayesian posterior probabilities (BPP) are indicated on each of the corresponding node. Support values for weekly supported (BS < 50, BPP < 0.50) nodes are indicated as “-”.

Figure 3

Phylogenetic trees of the genus Naja inferred by Bayesian analyses (BI) based on 627 bp mitochondrial gene COI. Both bootstrap supports (BS) and Bayesian posterior probabilities (BPP) are indicated on each of the corresponding node. Support values for weekly supported (BS < 50, BPP < 0.50) nodes are indicated as “-”.

Figure 4

Body and head morphological comparisons between Naja atra (A), N. kaouthia South Asian clade (B), N. kaouthia Southeastern Asian clade (C) and N. fuxi sp. nov. (D). (A) subadult female (body length 730 mm) neotype of N. atra CIB12273 from Zhoushan Island, Zhejiang Province; (B) adult female N. kaouthia ZMH R04803 from Port Canning, West Bengal, India; (C) adult male ZMH R02885 of N. kaouthia from Bangkok, Thailand; (D) adult male holotype of N. fuxi sp. nov. CIB DL2018053147 from Panzhihua, Sichuan, China. 1: dorsal body; 2: ventral body; 3: dorsal head; 4: ventral head; 5: lateral head. (A,D) photographed by Sheng-Chao Shi; (B,C) by Gernot Vogel.

Figure 5

Naja atra in life. (A) Adult from Qingtian, Zhejiang, China; (B–D) one adult (B,C) and another (D) from Lishui, Zhejiang, China; (E) juvenile from Wuyi, Fujian, China; (F,G) one adult from Guangzhou, Guangdong, China; (H) adult from Guangzhou, Guangdong, China. (A–E) Photographed by Bin-Qing Zhu; (F–H) photographed by Liang Zhang.

Figure 6

Hemipenis of Naja atra ((A) CIB LJT-FJ22020060; (B) CIB LJT-GD2020103) and N. fuxi sp. nov. ((C) KIZ 030071; (D) CIB DL2018053147). Sulcate, asulcate, lateral and top view from left to right. Photographed by Jun-Jie Huang and Jin-Long Ren.

Figure 7

South Asian Naja kaouthia in life. (A,B) Two adults from Jhargram, West Bengal, India; (C) Individual from Kolkata outskirts of West Bengal, India; (D) Adult from Siliguri, North Bengal region of West Bengal, India; (E) Individual from Bhuvneshwar, Odisha, India; (F,G) two juveniles from Mizoram, India; (H) Individual from Samtse town, Bhutan. ((A–E) photographed by Vivek R Sharma; (F,G) photographed by Gernot Vogel; (H) photographed by Choggyal Taz).

Figure 8

Southeastern Asian Naja kaouthia in life. (A,B) a juvenile from Selangor, Peninsular Malaysia; (C) an adult from Chang Wat Nakhon Ratchasima, Thailand; (D) a juvenile from Samut Prakan, Thailand; (E) an adult from Surat Thani, Thailand; (F) an adult from Ranong, Thailand; (G,H) two adults from Duc Trong, Lam Dong, Vietnam. (A) was photographed by Dr. Teo Eng Wah. Other photos were cited from iNaturalist.org occurrence dataset https://doi.org/10.15468/ab3s5x via GBIF.org (licensed under http://creativecommons.org/licenses/by-nc/4.0/) by following photographer: Gregory Greene ((C) record No. 2006052432), Wich’yanan L ((D) record No. 3090707753), Mintkhaosok ((E) record No. 3325726349), Knotsnake ((F) record No. 3384192379), Herpingvietnam ((G) record No. 2557801804), Leonid A. Neymark ((H) record No. 2366151765).

Figure 9

Paratypes of Naja fuxi sp. nov. (A,B) Dorsal and ventral view of adult female paratype CIB DL2018081005 from type locality Panzhihua, Sichuan before preservation; (C,D) Dorsolateral and ventral view of adult male KIZ030071 from Simao, Pu’er, Yunnan in preservative; (E,F) Dorsal and ventral view of juvenile CIB YNJC0068 from Jiangcheng, Yunnan, China before preservation. (Photographed by Sheng-Chao Shi).

Figure 10

Naja fuxisp. nov. in life and habitats. (A,B) Dorsolateral view and hood pattern of adult female paratype CIB DL2018081005 from Panzhihua, Sichuan, China; (C,D) Dorsolateral and front view of an unvouchered adult from Jiangcheng, Pu’er, Yunnan, China; (E,F), two adults from Wenshan, Yunnan, China; (G) Subtropical Forest at type locality Panzhihua, Sichuan, China; (H) Tropical monsoon forest and farmland at Jiangcheng, Pu’er, Yunnan, China. (Photos (A,B,H) by Sheng-Chao Shi; (C,D,G) by Li Ding; (E,F) by Liang Zhang).