Lycodon rufozonatus Cantor, 1842

Lycodon rufozonatus Cantor, 1842 View in CoL

Tables 2 View Table 2 , Fig. 3 View Figure 3 ; Suppl. material 1: table S 1, figs S 4 – S 11

Lycodon rufo-zonatus Cantor (1842: 483) — Holotype: NHMUK [The Natural History Museum, London, UK] 1843.7.21.36 donated by T. Cantor. Type locality: Chusan (now Zhoushan) Islands, Zhejiang Province, China. View in CoL

Dinodon cancellatum Duméril & Bibron in Duméril et al. 1854: 447. Holotype: not traced. Type locality: unknown, probably from China (see Stejneger 1907). View in CoL

Coronella striata Hallowell, 1856: 152. Syntypes: ANSP 3477–78. Type locality: Ningpo (now Ningbo), Zhejiang Province, China. View in CoL

Dinodon rufozonatus var. formosana Boettger, 1885: 125. Holotype: SMF 18045. Type locality: Formosa, now Taiwan. View in CoL

Dinodon rufozonatum williamsi Schmidt, 1925: 2. Holotype: AMNH 17453. Type locality: Changsha City, Hunan Province, China. View in CoL

Dinodon rufozonatum yunnanense Mell, 1931: 2007. Syntypes: ZMB 52629 –31, ZMB 27711. Type locality: Talifu (now Dali City), Yunnan Province, China. View in CoL

Material examined.

A total of 61 specimens ( 32 males and 29 females) were examined; see in Suppl. material 1: table S 1.

Referred materials.

A total of 14 specimens were used for reference ( seven males, six females and one sex unknown) and were reported by Duméril et al. (1854), Maki (1931), and Yang and Rao (2008); see in Suppl. material 1: table S 1.

Diagnosis.

Larger-sized species have a maximum snout-vent length of up to 1122 mm; a loreal slight entering the eye (rarely not); dorsal scales in 17 (19 or 21) – 17 (19) – 15 (16 or 17) rows, smooth throughout (rarely very faintly keeled posteriorly); 186–216 ventrals; 60–88 subcaudals, paired; cloacal plate undivided; eight supralabials with 3–5 touching the eye; one preocular, two postoculars; temporals 2 + 3; ground colour back with 60–106 red narrow crossbands on body and tail; ventral surface of body uniform cream, ventral surface of tail heavily dark speckled, not banded (based on Cantor 1842; Duméril et al. 1854; Hallowell 1856; Boettger 1885; Schmidt 1925; Mell 1931; Maki 1931; Yang and Rao 2008; this study).

Description of the holotype

( Fig. 3 View Figure 3 ): The body is robust and slightly laterally compressed. The tail is relatively long, thin, and tapering. The head is elongated, longer than wide, and moderately flattened, with a distinct separation from the neck. The snout is elongated, flattened, and slightly projects beyond the lower jaw. The nostrils are relatively large, positioned dorsolaterally, and round in shape. The eyes are relatively large, with vertical pupils.

Body size. SVL 370 mm, TaL 92 mm; ratio TaL / TL 0.199.

Body scalation. Dorsal scale rows 17–17 – 15, all smooth; scales of the vertebral row not enlarged; no apical pits; 198 ventrals; 74 subcaudals, all paired; cloacal plate undivided.

Head scalation. Rostral heptagonal, wider than high, slightly visible from above; nasal single, elongated; nasal surrounded by the first two supralabials, rostral, internasal, and prefrontal; internasals two, curved, slightly wider than longer, in contact with rostral anteriorly, nasal, and prefrontal; two prefrontals, large, subrectangular, prefrontal slightly shorter than frontal; prefrontals in contact with internasals, nasals, preoculars, and frontal; frontal rather small, pentagonal, tapering posteriorly, shorter than the distance from tip of snout to frontal; parietals longer than wide, in contact approximately the length of the frontal; 1 / 1 supraocular, distinctly wider than high, in contact with prefrontal; 1 / 1 loreal, contacting eye; 1 / 1 preocular, large, higher than wide, in broad contact with prefrontal; subocular absent; 1 / 2 postoculars; 2 + 3 temporals; 8 / 8 supralabials, first and second in contact with nasal, second and third contact with loreal, third and fifth in contact with eye, six and seven largest; infralabials 10 / 10, first pair in broad contact with each other, first to fifth in contact with anterior pair of chin shields; posterior chin shields smaller than anterior ones, separated from each other by a pair of small scales.

Colouration in preservative. The dorsal surface is blackish-brown, with 60 pale transverse crossbands on the body and 20 on the tail. The head is black, featuring a distinct inverted V-shaped marking on the nape. The ventral surface is cream-coloured, gradually becoming darker toward the cloaca, while the ventral surface of the tail is entirely dark.

General description and variation

(see Table 2 View Table 2 ; Suppl. material 1: table S 1, figs S 4 – S 11). Morphology variation based on 61 examined specimens as well as data morphology of 14 specimens were reported by Duméril et al. (1854), Maki (1931), and Yang and Rao (2008). The longest known specimen is 1,349 mm long (adult female; SVL 1145 mm, TaL 204 mm, ZMB 24830 A). The longest known male is 1,323 mm long ( SVL 1122 mm, TaL 201 mm; ZMB 19329 ). Body elongated; head distinct from neck, markedly flattened; eye medium; pupil vertically oval; rostral triangular, broader than high, clear visible from above; internasals as broad as long, approximately half the length of the prefrontal; prefrontal shorter than frontal; frontal hexagonal; parietals large, longer than wide; nasal divided; one loreal, nearly rectangular, narrowing posteriorly, protruding somewhat beneath preocular, usually entering the eye or not, not in entering with internasals; one (rarely absent or two) preocular; two (single) postoculars; two (single) anterior temporals; three posterior temporals (two); eight (seven or nice) supralabials, 1 st and 2 nd SL in contact with the nasal, 2 nd and 3 rd SL in contact with the loreal, 3 rd – 5 th SL entering orbit, 6 th and 7 th SL largest; ten (nice or 11) infralabials; first pair in contact each with other, 1 st – 5 th IL in contact with anterior chin shields, 5 th and 6 th IL largest; 17 (18, 19 or 21) dorsal scale rows at head, 17 (19) dorsal scale rows at midbody, 15 (16 or 17) dorsal scale rows at vent, the upper dorsal and vertebral scale rows entirely smooth or very faintly keeled posteriorly; ventrals 186–216 (199.60 ± 7.84, n = 75), without sexual dimorphism, vertebral scale slightly enlarged, distinctly angulate laterally; cloacal plate undivided; subcaudals 60–88 (74.30 ± 7.83, n = 63), without sexual dimorphism; relative tail length 0.151 –0.237 (0.196 ± 0.021, n = 63), without sexual dimorphism.

Colouration. The dorsal surface of the body and tail is blackish, with 42–78 narrow red or orange-pink crossbands on the body and 15–30 on the tail. Each pale crossband is ~ 1–3 dorsal scales wide, interconnecting to divide the ground colour into elliptical patches. The ventral surface of the body is uniformly cream, while the ventral surface of the tail is heavily speckled with dark markings. The head is black, with conspicuously red-margined plates and a distinct inverted V-shaped marking on the nape. Pale stripes extend downward from the top of the temporal scales to the last supralabial scale.

Etymology.

The species name consists of two Latin adjectives, rufus (meaning red) and zonatus (meaning banded), literally meaning “ red-banded ”. We recommend the following common names for this species: Red-banded Wolf Snake (in English); Grosszahnnatter (in German); Северный краснопоясный волкозуб “ Severyni krasnopoyasnyi volkozub ” (in Russian); 赤链蛇 “ Chì liàn shé ” (in Chinese); 능구렁이 “ Neung-guleong-I ” (in South Korean); アカマダラ “ Akamadara ” (in Japanese); R ắn khuy ết d ải thân đ ỏ (in Vietnamese).

Distribution

(Fig. 1 View Figure 1 ). China: This species is widely distributed across the country, occurring in the provinces of Anhui, Beijing, Chongqing, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hainan, Henan, Hebei, Hubei, Hunan, Jilin, Jiangsu, Jiangxi, Liaoning, Sichuan, Shandong, Shanghai, Shanxi, Shaanxi, Tianjin, Yunnan, and Zhejiang ( Zhao and Adler 1993; Wang et al. 2021; this study). Taiwan: This species is very common and widely distributed throughout the island (C. W. You, pers. obs.). South Korea: It is likely to be common in the country and has been recorded in Incheon City, Gangwon Province, Seoul City, and Busan City ( Shin et al. 2024; this study). Japan: Reported from Uotsuri and Tsushima Islands, Nagasaki Prefecture ( Li et al. 2017; Morikawa and Inoue 2025). Russia: Recorded in Nezhino, Chernigovka, and the Posyet districts, all within the southern part of Primorsky Krai ( Maslov and Kotlobay 1998; Li et al. 2017; Sundukov 2025). Vietnam: We confirm the occurrence of this species in northeastern Vietnam, including Tuyen Quang Province (Na Hang NR, based on specimen ROM 30814 ; see Suppl. material 1: fig. S 9 A), Vinh Phuc Province (Tam Dao NP, based on specimen ROM 34615 ; see Suppl. material 1: fig. S 9 B), and an individual observed in Tay Yen Tu NR, Bac Giang Province (see Suppl. material 1: fig. S 11 D). Additional records from previously reported locations are discussed below. Laos: The record from Xiengkhouang Province ( Deuve 1970) was later revised as Lycodon meridionalis Bourret ( Orlov and Ryabov 2004; this study).

Natural history notes.

This species is common in China, Taiwan, and South Korea but is rare in Russia, Japan, and Vietnam ( Ananjeva et al. 2006; this study). It is an oviparous species occurring in a wide range of habitats, including plains, hills, and montane areas, from boreal to tropical forests. It is also found in villages and other rural areas, typically near water bodies, at elevations of ca 850–1,170 m asl. This species is primarily nocturnal and terrestrial but has been occasionally observed swimming. Its diet includes a wide range of vertebrates. Prey items include fish and toads, such as Duttaphrynus cf. gymnauchen (Bleeker) and Bufo gargarizans Cantor ; frogs, including Pelophylax nigromaculatus (Hallowell) , Fejervarya limnocharis (Gravenhorst) , F. kawamurai Tjong, Matsui, Kuramoto, Nishioka & Sumida , Kaloula cf. pulchra Gray , Limnonectes cf. fujianensis Ye & Fei , and Polypedates braueri (Vogt) ; and lizards such as Diploderma polygonatum Hallowell , Plestiodon elegans (Boulenger) , tail fragments of Scincella vandenburghi (Schmidt) , and Gekko japonicus (Duméril & Bibron) . It also preys on other snakes, including Oocatochus rufodorsatus (Cantor) , eggs of Elaphe climacophora (Boie) , and Gloydius tsushimaensis (Isogawa, Moriya & Mitsui) , as well as young birds, rats, and even domestic animals such as a guinea pig ( Pope 1935; Morikawa and Inoue 2025; TVN, pers. obs.). Females lay clutches of more than ten eggs ( Pope 1935; Dieckmann et al. 2010).