Expanded description & range extension of Lycodon sidiki Wostl, Hamidy, Kurniawan & Smith, 2017 (Squamata: Colubridae) Author Nguyen, Tan Van Institute for Research and Training in Medicine, Biology and Pharmacy, Duy Tan University, Da Nang, 550000, Vietnam. & College of Medicine and Pharmacy, Duy Tan University, 120 Hoang Minh Thao, Lien Chieu, Da Nang, 550000, Vietnam Author David, Patrick Reptiles & Amphibiens, UMR 7205 OSEB, Département évolution et Systématique, CP 30, Muséum National d’Histoire Naturelle. 57 rue Cuvier, F- 75231 Paris Cedex 05, France. Author Vogel, Gernot Society for South East Asian Herpetology, Im Sand- 3, D- 69115 Heidelberg, Germany. text Zootaxa 2024 2024-08-07 5492 4 563 581 http://dx.doi.org/10.11646/zootaxa.5492.4.5 journal article 10.11646/zootaxa.5492.4.5 1175-5326 13269088 87906811-8BB8-4AFC-A50A-8F27B8F9DF50 Redescription of Lycodon sidiki Wostl, Hamidy, Kurniawan & Smith, 2017 ( Figures 1–2 , 3A–G , 4A ; Tables 1–2 ) Holotype : MZB .Ophi.598, adult male . Type locality : Between Takengon and Isaq , Aceh Province , Sumatra, Indonesia ( 04.50611°N , 96.86061°E ; altitude 1,614 m asl ). Additional specimens examined (n=8), all from West Sumatra Province , Indonesia : NHMW 23146 :1 ( adult female ) and NHMW 23146 :2 ( subadult female ) from Padang , collected by F. Schild in 1900 ; NHMW 23146 : 7 ( adult males ) , NHMW 23146 : 4 ( subadult male ) , NHMW 23146 : 5 & 6 ( two adult females ) , NHMW 23146 : 8 ( subadult female ), from Padang , collected by F. Schild in 1899 ; NHMW 23149 :1 ( one adult male ) no specific location, donated by Steindachner in 1900 . Revised diagnosis . Medium-sized species, maximum total length 830 mm ; loreal present (sometimes fused with the prefrontal), usually in contact with eye; dorsal scale rows 17–17–15; 5–7 upper dorsal scale rows feebly keeled at midbody; 195–212 ventrals; 77–91 subcaudals, paired; cloacal plate undivided; 8 supralabials with 3 rd –5 th SL touching the eye; 1 preocular (rarely fused with prefrontal), 2 postoculars; temporals 2+2; dorsal crossbands narrow, cream colored, 25–33 crossbands on dorsum, 10–16 crossbands on tail; venter with discreet bands throughout (based on Wostl et al . 2017 and this study). Expanded description of the species, including all known specimens (n=9): Morphology . Body elongate, somewhat laterally compressed; longest known specimen 830 mm long (SVL 650 mm , TaL 180 mm , male; NHMW 23149 :1); longest known female 718 mm long (SVL 555 mm , TaL 163 mm ; NHMW 23146 :1)]; tail comparatively long, thin and tapering (Ratio TaL/TL 0.213 –0.239 in males (n=3) and 0.193 –0.227 in females (n=5)). Head subrectangular, elongate, flattened, well-distinct from the neck; snout elongate, strongly flattened, oblique in profile, projecting over the lower jaw; nostril oval, large; eye moderate in size, pupil vertically elliptic. Body scalation . DSR 17–17–15, the 5–7 upper rows feebly keeled; VEN 195–210 in males (n=4), 207–212 in females (n=5); SC 80–91 in males (n=4), 77–84 in females (n=5); paired; single cloacal plate. Head scalation . Rostral triangular, hardly visible from above; nasal vertically divided by a furrow below and above the nostril; two small internasals, widely in contact with each other behind the rostral and with prefrontals; two large prefrontals, longer and wider than internasals; rather small, more or less triangular frontal; 2 large parietals, each bordered by three large scales, i.e., 2 upper temporals and one larger paraparietal posteriorly; 1/1 supraocular; 1/1 preocular, above posterior part of loreal; 2/2 subequal postoculars; 1/1 loreal (6/ 9 specimens ), sometime fused with prefrontal (3/ 9 specimens ), rather elongate and narrow, entering orbit, in contact with SL 2 and 3, preocular, prefrontal and posterior part of nasal; 8/8 SL in all known specimens, SL 1–2 in contact with nasal, SL 3–5 entering orbit, 6 th SL largest; 2+2 (8/ 9 specimens ) or rarely 2+3 (1/ 9 specimens ) temporals, lower anterior temporal slightly broader than upper; 9 (5/ 9 specimens ), less frequently 8 (2/ 9 specimens ) or 10 (2/ 9 specimens ) infralabials, IL 1–5 in contact with first pair of chin shields anterior and posterior chin shields about same length on right side, on left side anterior larger than posterior, anterior pair wider. FIGURE 1 . Lycodon sidiki in preservative, specimen NHMW 23149:1 (male) in West Sumatra, Indonesia—(A) General dorsal view; (B) General ventral view; (C) Lateral view of the head, right side; (D) Lateral view of the head, left side; (E) Dorsal view of the head; (F) Ventral view of the head. Photos by G. Vogel. Coloration in preservative (see Figs. 1 , 2 , 3A–G ): Body and tail dark brown, with 25–33 cream crossbands on the body and 10–14 crossbands on tail, anteriorly wider, becoming narrow posteriorly; crossbands about 1 dorsal scale broad middorsally, widening ventrolaterally to about the length of 2–2.5 dorsals. Head uniformly blackishbrown; no collar. Venter dark with clear, rather irregular, cream bands, 6–10 ventrals broad, with 3–5 ventrals between each band. FIGURE 2. Lycodon sidiki in preservative, from Sumatra, Indonesia—(A) NHMW 23146:1 (adult female); (B) NHMW 23146:2 (subadult female); (C): NHMW 23146:4 (adult male); (D) NHMW 23146:5 (adult female); (E) NHMW 23146:6 (adult female); (F) NHMW 23146:7 (adult male). Photos by G. Vogel. Comparisons. Lycodon sidiki is morphologically and phylogenetically placed within the Lycodon fasciatus group (eg. Wostl et al . 2017 ; Wang et al . 2021 ; Janssen et al 2019 ; Shu et al . 2024 ; Nguyen et al . in review; this study) and is overall most similar to other species from the Lycodon fasciatus group, of which we gave the list of included species above. Therefore, the comparisons with these 14 species appear to be the most pertinent. The main diagnostic characters separating Lycodon sidiki from these 14 species are summarized in Table 2 and Figs. 4–5. L . sidiki is distinguished from L . butleri by having: lower number of VEN in both sexes (195–210 [avg. 204.3] in males, 207–212 [avg. 209.8] in females vs. 216–227 [avg. 223.0] in males, 221–228 [avg. 224.5] in females); lower number of SC in females (77–84 [avg. 81.3] vs. 88–92 [avg. 90.5]; lower number of crossbands on the tail in females (10–14 [avg. 11.2] vs. 15–23 [avg. 18.9]. L . sidiki is distinguished from L . davidi by having: lower number of VEN in males (195–210 [avg. 204.3] vs. 224); lower number of SC in males (80–91 [avg. 84.8] vs. 99); lower number of bands on body in males (25–32 [avg. 28.3] vs. 84); lower number of crossbands on tail in males (13–16 [avg. 14.5] vs. 38); loreal in contact with eye (vs. not in contact); bands on venter present (vs. absent). FIGURE 3. Lateral view of the head, right side, of Lycodon sidiki and L . subcinctus from Padang, West Sumatra, Indonesia— L . sidiki : (A) NHMW 23146:1 (adult male); (B) NHMW 23146:2 (subadult female); (C) NHMW 23146:4 (subadult male); (D) NHMW 23146:5 (adult female); (E) NHMW 23146:6 (adult female); (F) NHMW 23146:7 (adult male); (G) NHMW 23146:8 (subadult female).— L . subcinctus : (H) NHMW 23146:9 (subadult female); (I) SMF 22442 (subadult male). Photos by G. Vogel. L . sidiki is distinguished from L . cavernicolus by having: larger size in males (max SVL 650 mm vs. 406 mm ); lower number of VEN in both sexes (195–210 [avg. 204.3] in males, 207–212 [avg. 209.8] in females vs. 245 in male, 232 in female); lower number of SC in both sexes (80–91 [avg. 84.8] in males, 77–84 [avg. 81.3] in females vs. 113 in male, 92 in female); lower number of crossbands on body in both sexes (25–32 [avg. 28.3] in males, 25–33 in females vs. 45 in male, 36 in female); lower number of crossbands on tail in both sexes (13–16 [avg. 14.5] in males, 10–14 [avg. 11.2] in females vs. 41 in male, 29 in female). L . sidiki is distinguished from the morphologically similar L . fasciatus by having: slightly smaller size in both sexes (max SVL 650 mm in males, 555 mm in females vs. 706 mm in males, 604 mm in females); slightly higher ratio TaL/TL in males (0.21–0.24 [avg. 0.23] vs. 0.20–0.22 [avg. 0.21]); colour of crossbands (whitish or cream vs. brown or grayish-brown). In L . sidiki there is a tendency to a fusion of the loreal with the prefrontal (3/ 9 specimens on both sides), a character which was not observed in the 44 specimens of L . fasciatus that were examined (see Appendix I). Furthermore, L . sidiki is widely separated from L . fasciatus on a geographical basis, as the latter species inhabits China , Myanmar , India , Bhutan , Thailand , Northern Laos , and Northwest Vietnam . Moreover, L . butleri and L . cavernicolus separate the ranges of both species. L . sidiki is distinguished from L . gammiei by having: smaller size in females (max SVL 555 mm vs. 1,110 mm ); lower number of SC in both sexes (80–91 [avg. 84.8] in males, 77–84 [avg. 81.3] in females vs. 101–110 [avg. 105.5] in males, 101–112 [avg. 107.0] in females); lower number of crossbands on the body in both sexes (25–32 [avg. 28.3] in males, 25–33 [avg. 27.4] in females vs. 42–57 [avg. 49.5] in males, 43–58 [avg. 49.5] in females); loreal in contact with eye (vs. not in contact); colour of crossbands (whitish or cream vs. yellow). L . sidiki is distinguished from L . gongshan by having: smaller size in both sexes (max SVL 650 mm in males, 555 mm in females vs. 740 mm in males, 798 mm in females); lower number of SC in females (77–84 [avg. 81.3] vs. 92–94 [avg. 93.0]); lower number of crossbands on the body in females (25–33 [avg. 27.4] vs. 32–36 [avg. 34.3]). FIGURE 4 . Photos in life of Lycodon sidiki compared with other species of the L . fasciatus group (Part 1). (A) L . sidiki in Aceh, Sumatra, Indonesia; (B) L . butleri in Fraser’s Hill, Selangor, Malaysia; (C) L . cavernicolus in Gua Wang Burma, Perlis, Malaysia; (D) L . davidi in Vang Vieng, Vientiane, Laos; (E) L . fasciatus in Aizawl, Mizoram, India; (F) L . gammiei in Eaglenest WS, Arunachal Pradesh, India; (G) L . gongshan in Nujiang, Yunnan, China; (H) L . liuchengchaoi in Lishui, Zhejiang, China. Photos by: (A) reproduced from Wostl et al. (2017) ; (B) R. Grassby-Lewis; (C) M.A. Muin; (D) reproduced from Vogel et al. (2012) ; (E) P. Shinde; (F) C. Sheth; (G) S.C. Shi; and (H) F. Gao. FIGURE 5. Photos in life of Lycodon sidiki compared with other species of the L . fasciatus group (Part 2). (A) L . multifasciatus in Ishigaki, Okinawa, Japan; (B) L . multizonatus in Yingjing, Sichuan, China; (C) L . obvelatus in Panzhihua, Sichuan, China; (D) L . pictus in Ba Be NP, Bac Kan, Vietnam; (E) L . ruhstrati ruhstrati in Baling, Taiwan; (F) L . ruhstrati abditus in Dong Chau-Ke Nuoc Trong NR, Quang Tri, Vietnam; (G) L . serratus in Litang, Sichuan, China; (H) L . synaptor in Kunming, Yunnan, China. Photos by: (A) D. By; (B) H. Lagoon; (C) reproduced from Wang et al. (2021) ; (D) L.A. Neymark; (E) P. Freed; (F) T.V. Nguyen; (G) Y. Cui; and (H) Y.H. Pu. FIGURE 6 . Map showing distributions of the Lycodon fasciatus group in Malay Peninsula and Sumatra including: L . butleri , L . cavernicolus , and L . sidiki . Notes: numbers indicate the different localities where these species have been recorded (see Appendix II for the details of localities). TABLE 1. Main measurements and meristic characters of the examined specimens of Lycodon sidiki from Sumatra, Indonesia. Notes : N/a: not available, * : preoculars fused with prefrontals; ** : loreal fused with prefrontals.
No. Collection Locality Sex SVL TaL TL TaL/ VEN SC ASR MSR DSR
number (mm) (mm) (mm) TL
1 MZB.Ophi.5980 Aceh M 548 167 715 0.23 195 85 17 17 15
2 NHMW 23146:4 Padang SM 378 102 480 0.21 210 80 17 17 15
3 NHMW 23146:7 Padang M 405 127 532 0.24 202 91 17 17 15
4 NHMW 23149:1 No M 650 180 830 0.22 210 83 17 17 15
specified
5 NHMW 23146:1 Padang F 555 163 718 0.23 207 82 17 17 15
6 NHMW 23146:5 Padang F 499 143 642 0.22 209 84 17 17 15
7 NHMW 23146:6 Padang F 426 118 544 0.22 211 82 17 17 15
8 NHMW 23146:2 Padang SF 371 N/a N/a N/a 212 N/a 17 17 15
9 NHMW 23146:8 Padang SF 201 48 249 0.19 210 77 17 17 15
TABLE 1. (Continued)
Collection Locality Sex SL IL Lor Lor-E PrO PoO AT PT BB TB Source
No. number
MZB.Ophi.5980 Aceh M 8/8 10/9 1/1 1/1 0/0* 2/2 2/2 2/2 25 16 Wostl et al .
1 (2017)
2 NHMW 23146:4 Padang SM 8/8 9/9 1/1 1/1 1/1 2/2 2/2 2/2 32 13 This study
3 NHMW 23146:7 Padang M 8/8 8/8 1/1 1/1 1/1 1/1 2/2 2/2 25 16 This study
4 NHMW 23149:1 No specified M 8/8 9/9 0/0** 1/1 1/1 2/2 2/2 2/2 31 13 This study
5 NHMW 23146:1 Padang F 8/8 9/8 0/0** 1/1 1/1 2/2 2/2 2/2 25 10 This study
6 NHMW 23146:5 Padang F 8/8 8/8 0/0** 1/1 1/1 2/2 2/2 3/3 33 14 This study
7 NHMW 23146:6 Padang F 8/8 8/10 1/1 1/1 1/1 2/2 2/2 2/2 29 12 This study
8 NHMW 23146:2 Padang SF 8/8 9/9 1/1 1/1 1/1 2/2 2/2 2/2 25 10 This study
9 NHMW 23146:8 Padang SF 8/8 9/9 1/1 1/1 1/1 2/2 2/2 2/2 25 10 This study
L . sidiki is distinguished from L . liuchengchaoi by having: smaller size in females (max SVL 555 mm vs. 670 mm ); lower number of SC in males (80–91 [avg. 84.8] vs. 68–81 [72.7]); lower number of crossbands on the body in males [25–32 [avg. 28.3] vs. 40–45 [avg. 43.3]); colour of crossbands (whitish or cream vs. yellow). L . sidiki is distinguished from L . multifasciatus by having: larger size in males (max SVL 650 mm vs. 490 mm ); lower number of VEN in both sexes (195–210 [avg. 204.3] in males, 207–212 [avg. 209.8] in females vs. 237 in males, 227–232 [avg. 229.8] in females]; lower number of SC in both sexes (80–91 [avg. 84.8] in males, 77–84 [avg. 81.3] in females vs. 117 in males, 106–108 [avg. 107.0] in females); lower number of crossbands on body in both sexes (25–32 [avg. 28.3] in males, 25–33 in females vs. 72 in male, 51–60 [avg. 56.8] in females); lower number of crossbands on tail in both sexes (13–16 [avg. 14.5] in males, 10–14 [avg. 11.2] in females vs. 35 in male, 25–32 [avg. 28.7] in females); loreal in contact with eye (vs. not in contact); ventral pattern (banded vs. uniform). L . sidiki is distinguished from L . multizonatus by having: larger size in males (max SVL 650 mm vs. 505 mm ); higher ratio TaL/TL in males (0.21–0.24 [avg. 0.23] vs. 0.16–0.18 [avg. 0.17]); higher number of SC in males (80–91 [avg. 84.8] vs. 56–75 [avg. 65.5]); lower number of crossbands on the body in males (25–32 [avg. 28.3] vs. 55–62 [57.8]); temporals 2+2 (vs. 2+3); cloacal plate (undivided vs. divided); ventral pattern (banded vs. uniform); colour of crossbands (whitish or cream vs. pink). TABLE 2. Comparison of morphological characters of Lycodon sidiki with those of the Lycodon fasciatus group. Characters used are : (1) : max SVL (mm) in males; ( 2 ): max SVL (mm) in females; ( 3 ): ratio TaL/TL in males ; (4) : ratio TaL/TL in females; (5) : VEN in males; (6) : VEN in females. Diagnostic differences from Lycodon sidiki are marked in bold. N/a: not available.
Species 1 2 3 4 5 6
L . sidiki 650 555 0.21–0.24 0.19–0.23 195–210 207–212
(0.23±0.01, n=4) (0.21±0.02, n=5) (204.3±7.2, n=4) (209.8±1.9, n=5)
L . butleri 648 563 0.19–0.24 0.20–0.22 216–227 221–228
(0.21±0.01, n=9) (0.21±0.01, n=3) (223.0±3.6, n=9) (224.5±2.9, n=4)
L . davidi 308 N/a 0.21 N/a 224 N/a
(n=1) (n=1)
L . cavernicolus 406 N/a 0.20 0.21 245 232
(n=1) (n=1) (n=1) (n=1)
L . fasciatus 706 604 0.20–0.22 0.19–0.22 197–213 201–214
(0.21±0.01, n=21) (0.20±0.01, n=18) (203.8±4.2, n=26) (208.0±3.4, n=24)
L . gammiei 632 1110 0.20–0.24 0.19–0.25 205–217 205–228
(0.22±0.03, n=2) (0.22±0.03, n=6) (211.00±8.49, n=2) (216.14±7.13, n=7)
L . gongshan 740 798 0.21–0.27 0.22–0.23 203–216 210–215
(0.23±0.02, n=8) (0.22±0.01, n=2) (210.3±3.6, n=10) (212±2.6, n=3)
L . liuchengchaoi 676 670 0.20–0.22 0.16–0.22 202–204 200–228
(0.21±0.02, n=2) (0.19±0.02, n=6) (203.0±1.0, n=3) (207.3±9.7, n=7)
L . multifasciatus 490 570 0.24 0.23–0.26 237 227–232
(n=1) (0.25±0.01, n=5) (n=1) (229.8±2.2, n=4)
L . multizonatus 505 N/a 0.16–0.18 N/a 191–202 N/a
(0.17±0.01, n=4) (195.5±4.7, n=4)
L . obvelatus 447 N/a 0.19 N/a 199 N/a
(n=1) (n=1)
L . pictus 500 543 0.18–0.23 0.21–0.23 209–215 213–218
(0.21±0.04, n=2) (0.21±0.02, n=4) (212.0±3.0, n=3) (215.0±2.1, n=5)
L . ruhstrati abditus 761 762 0.19–0.24 0.19–0.24 206–224 220–229
(0.22±0.01, n=15) (0.22±0.01, n=19) (217.2±5.5, n=20) (224.2±3.3, n=19)
L . ruhstrati ruhstrati 810 672 0.22–0.25 0.21–0.24 211–228 216–228
(0.23±0.01, n=7) (0.23±0.01, n=11) (217.7±4.9, n=9) (220.4±3.2, n=12)
L . serratus 480 N/a 0.24 N/a 198 N/a
(n=1) (n=1)
L . synaptor 550 525 0.20–0.21 0.18–0.21 186–199 190–203
(0.21±0.01, n=6) (0.19±0.01, n=9) (191.8±4.3, n=6) (194.9±4.3, n=9)
......continued on the next page TABLE 2. (Continued). Characters used are : (7) : SC in males; (8) : SC in females; (9) : Bands on the body in males; (10) : Bands on the body in females; (11) : Bands on the tail in males; (12) : Bands on the tail in females. Remark : Diagnostic differences from Lycodon sidiki are marked in bold. N/a: not available.
Species 7 8 9 10 11 12
L . sidiki 80–91 77–84 25–32 25–33 13–16 10–14
(84.8±4.6, n=4) (81.3±3.0, n=4) (28.3±3.8, n=4) (27.4±3.6, n=5) (14.5±1.7, n=4) (11.2±1.8, n=5)
L . butleri 81–97 88–92 28–37 29–33 14–25 15–23
(90.5±5.2, n=8) (90.5±5.2, n=4) (33.0±2.6, n=9) (30.5±1.7, n=4) (18.9±3.5, n=8) (18.9±3.5, n=4)
L . davidi 99 (n=1) N/a 84 (n=1) N/a 38 (n=1) N/a
L . cavernicolus 113 92 45 36 41 29
(n=1) (n=1) (n=1) (n=1) (n=1) (n=1)
L . fasciatus 76–88 74–90 24–43 26–37 10–20 10–18
(82.3±3.8, n=22) (80.8±3.7, n=19) (32.5±3.8, n=26) (32.5±3.0, n=24) (16.0±2.6, n=26) (15.3±2.1, n=23)
L . gammiei 101–110 101–112 42–57 43–58 18–23 16–21
(105.5±6.4, n=2) (107.0±4.9, n=5) (49.5±10.6, n=2) (49.9±5.2, n=7) (20.5±3.5, n=2) (18.8±2.2, n=5)
L . gongshan 79–96 92–94 26–46 32–36 9–16 12–15
(85.7±6.9, n=7) (93.0±1.4, n=3) (36.4±6.4, n=9) (34.3±2.1, n=3) (12.8±3.1, n=8) (14.0±1.7, n=3)
L . liuchengchaoi 68–81 75–81 40–45 18–47 10–14 7–14
(72.7±7.2, n=3) (78.0±2.4, n=5) (43.3±2.9, n=3) (34.5±10.6, n=6) (12.0±2.8, n=3) (11.2±2.5, n=6)
L . multifasciatus 117 106–108 72 51–60 35 25–32
(n=1) (107.0±1.0, n=3) (n=1) (56.8±4.3, n=4) (n=1) (28.7±3.5, n=3)
L . multizonatus 56–75 N/a 55–62 N/a 11–19 N/a
(65.5±8.0, n=4) (57.8±3.4, n=4) (14.0±3.6, n=4)
L . obvelatus 76 (n=1) N/a 31 (n=1) N/a 13 (n=1) N/a
L . pictus 82–90 85–91 29–32 25–30 8–14 10–16
(86.8±5.7, n=2) (89.0±2.7, n=4) (30.0±1.7, n=3) (28.4±1.9, n=5) (11.3±3.1, n=3) (13.4±2.3, n=5)
L . ruhstrati 85–102 82–106 31–46 17–44 14–23 9–19
abditus (93.9±4.4, n=16) (97.1±6.0, n=19) (36.9±4.6, n=16) (30.4±8.1, n=16) (16.7±2.3, n=13) (14.8±3.0, n=14)
L . ruhstrati 105–114 97–108 37–46 33–40 16–25 14–22
ruhstrati (107.4±3.3, n=7) (103.0±3.6, (41.6±3.0, n=9) (36.4±1.8, n=12) (21.3±2.5, n=8) (18.5±2.4, n=11)
n=10)
L . serratus 84 (n=1) N/a 66 (n=1) N/a 26 (n=1) N/a
L . synaptor 74–79 67–77 22–36 20–31 8–12 7–11
(76.2±2.0, n=6) (70.9±3.7, n=9) (25.8±5.1, n=6) (24.7±3.8, n=9) (10.2±1.5, n=6) (8.6±1.2, n=9)
TABLE 2. (Continued). Characters used are : (13) : Dorsal scales; ( 14) : Anal; ( 15) : Loreal in contact with eye; (16): Temporals; (17): Belly pattern; (18) Colour of bands. Remark : Diagnostic differences from Lycodon sidiki are marked in bold. N/a: not available.
Species 13 14 15 16 17 18 Distributions Sources
L . sidiki Keeled Entire Yes 2+2 Banded Whitish or cream Indonesia Wostl et al . (2017) ; our data
L . butleri Entire Yes 2+2 Banded & Whitish Thailand, our data
Keeled spotted Peninsular
Malaysia
L . davidi Keeled Entire No 2+3 Banded & spotted Whitish Laos Vogel et al . (2012)
L . cavernicolus Keeled Entire Yes 2+3 Banded Whitish Peninsular Grismer et al . (2004)
Malaysia
......continued on the next page TABLE 2. (Continued).
Species 13 14 15 16 17 18 Distributions Sources
L . fasciatus Keeled Entire Yes 2+2 Banded Brown or China, Myanmar, our data
grayish-brown India, Bhutan,
Thailand
L . gammiei Keeled Entire No 2+2 Banded Yellow India, Bhutan, Mistry et al . (2007) ;
China Chettri & Bhupathy
(2009); Shu et al .
(2024)
L . gongshan Keeled Entire Yes 2+2 Banded Whitish China, Myanmar our data
L . liuchengchaoi Keeled Divided Yes 2+2 Banded Yellow China our data
or Entire
L . multifasciatus Keeled Entire No 2+3 None Whitish Japan Maki (1931) ; Vogel et
al . (2009); our data
L . multizonatus Keeled Divided Yes 2+3 None Pink China Lei et al . (2014) ; our
data
L . obvelatus Smooth Entire Yes 2+2 None Salmon China Wang et al . (2021) ;
(2+3) our data
L . pictus Smooth Entire Yes or 2+3 None Whitish Vietnam, China Janssen et al . (2019 ,
No (1+3) 2020); Xu et al .
(2023); our data
L . ruhstrati abditus Keeled Entire No 2+3 Banded & spotted Whitish China, Vietnam, Laos Vogel et al . (2009) ; our data
L . ruhstrati ruhstrati Keeled Entire No 2+3 Banded & spotted Whitish Taiwan Vogel et al . (2009) ; our data
L . serratus Smooth Divided Yes 2+2 None Tawny olive China Wang et al . (2021)
L . synaptor Keeled Entire Yes 2+2 Banded Whitish China, Vietnam our data
L . sidiki is distinguished from L . obvelatus by having: larger size in males (max SVL 650 mm vs. 447 mm ); higher ratio TaL/TL in males (0.21–0.24 [avg. 0.23] vs. 0.19]); higher number of SC in males (80–91 [avg. 84.8] vs. 76]; dorsal scales keeled vs. smooth; ventral pattern (banded vs. uniform); colour of body crossbands (whitish or cream vs. salmon). L . sidiki is distinguished from L . pictus by having: larger size in males (max SVL 650 mm vs. 500 mm ); lower number of VEN in males (195–210 [avg. 204.3] vs. 209–215 [avg. 212.0]); lower number of SC in females (77–84 [avg. 81.3] vs. 85–91 [avg. 89.0]); dorsal scales keeled (vs. smooth); ventral pattern (banded vs. uniform). L . sidiki is distinguished from L . ruhstrati abditus by having: smaller size in both sexes (max SVL 650 mm in males, 555 mm in females vs. 761 mm in males, 762 mm in females); lower number of VEN in both sexes (195–210 [avg. 204.3] in males, 207–212 [avg. 209.8] in females vs. 206–224 [avg. 217.2] in males, 220–229 [avg. 224.2] in females); lower number of SC in both sexes (80–91 [avg. 84.8] in males, 77–84 [avg. 81.3] in females vs. 85–102 [avg. 93.9] in males, 82–106 [avg. 97.1] in females); lower number of crossbands on body in males (25–32 [avg. 28.3] vs. 31–46 [avg. 36.9]); loreal in contact with eye (vs. not in contact). L . sidiki is distinguished from L . ruhstrati ruhstrati by having: smaller size in both sexes (max SVL 650 mm in males, 555 mm in females vs. 810 mm in males, 672 mm in females); lower number of VEN in both sexes (195–210 [avg. 204.3] in males, 207–212 [avg. 209.8] in females vs. 211–228 [avg. 217.7] in males, 216–228 [avg. 220.4] in females); lower number of SC in both sexes (80–91 [avg. 84.8] in males, 77–84 [avg. 81.3] in females vs. 105–114 [avg. 107.4] in males, 97–108 [avg. 103.0] in females); lower number of crossbands on body in both sexes (25–32 [avg. 28.3] in males, 25–33 [avg. 27.4] in females vs. 37–46 [avg. 41.6] in males, 33–40 [avg. 36.4] in females]; lower number of crossbands on tail in both sexes (13–16 [avg. 14.5] in males, 10–14 [avg. 11.2] in females vs. 16–25 [avg. 21.3] in males; 14–22 [avg. 18.5] in females); loreal in contact with eye (vs. not in contact). L . sidiki is distinguished from L . serratus by having: larger size in males (max SVL 650 mm vs. 480 mm ); lower number of crossbands on body in males (25–32 [avg. 28.3] vs. 66]); lower number of crossbands on tail in males (13–16 [avg. 14.5] vs. 26); dorsal scales keeled (vs. smooth); ventral pattern (banded vs. uniform); colour of crossbands (whitish or cream vs. tawny olive). L . sidiki is distinguished from L . synaptor by having: slightly larger size in males (max SVL 650 mm vs. 550 mm ); slightly higher ratio TaL/TL in females (0.19–0.23 [avg. 0.21] vs. 0.18–0.21 [avg. 0.19]); higher number of VEN in both sexes (195–210 [avg. 204.3] in males, 207–212 [avg. 209.8] in females vs. 186–199 [avg. 191.8] in males, 190–203 [avg. 194.9] in females); slightly higher number of SC in females (77–84 [avg. 81.3] vs. 67–77 [avg. 70.9]).
Distribution ( Fig. 6 ). Lycodon sidiki was previously known only from Aceh Province , Sumatra , Indonesia ( Wostl et al . 2017 ). We here add a second locality from Padang, West Sumatra Province . The new location in Padang is ca. 710 airline kilometers southeast of the type locality. It also represents the southernmost locality of the Lycodon fasciatus species group. Given its geographic proximity, it likely occurs in several forests of Sumatra ; in particular, records in the highlands of the provinces of North Sumatra , Riau , and Jambi are anticipated.