AnimaliaSquamataElapidaeChenZe-NingShiSheng-ChaoVogelGernotDingLiShiJing-SongMultiple lines of evidence reveal a new species of Krait (Squamata, Elapidae, Bungarus) from Southwestern China and Northern MyanmarZookeys18320211025357110.3897/zookeys.1025.62305B28A12D4-F521-5F33-AC6F-609B5D59AEDA Bungarus candidus (Linnaeus, 1758)Figs 4C, D, 5C, D, 6C, D, 7C, D, 9D–F [English name: Blue Krait] [Chinese name: 马来环蛇] Coluber candidusLinnaeus 1758: 223. Bungarus candidus– Cantor 1847 Bungarus semifasciatusBoie 1827 Aspidoclonion semifasciatum– Wagler 1828 Bungarus candidus var. semifasciata– Werner 1900 Bungarus javanicusKopstein 1932 (fideSlowinski 1994) Bungarus candidus– Smith 1943: 416Type locality.

“Indiis” (in error). Holotype: NRM 37 (formerly ZIUS 89).

Typical B. candidus possesses following morphological characters based on the examination of 19 specimens from Sumatra and Java, Indonesia; Peninsular Malaysia (Appendix 1): (1) Dorsum of most specimens with 21.4 ± 1.8, (18–26) broad white crossbands, with each band covers 3.8 ± 0.6, (3.0–5.0) vertebral scales on midbody, (Figs 4C, D, 5C, D), uniform black in some populations (Kuch 2007); (2) ventral body immaculate white, without brown pigments (Figs 4C, D, 5C, D); (3) scales on temporal area and lateral neck stained white, contrast with neighbor scales on neck in adults, creamy white in juveniles (Fig. 6C, D); (4) black bands on body large, covering 3–5 vertebral scales on middle body, intruding to white ventral body, ventrals with narrow black edges 1–2 times of outer dorsal scales (Fig. 7C, D); (5) ventral tail with broad dark crossbands (Figs 4D, 5D); (6) posterior maxilla teeth four (n = 9), slightly curved behind, (Table 3); (7) prefrontal suture 1.4–2.4 (n = 17) times the length of internasals suture; (8) VEN = 209–224 (n = 18), NSC = 41–50 (n = 17).

10.3897/zookeys.1025.62305.figure7F9D4E8F2-1357-5A5A-8401-332BF5627952

Body patterns of adults (left) and juveniles (right) of the Bungarus candidus/multicinctus/wanghaotingi complex and B. suzhenae sp. nov. AB. multicinctus, adult male CIB DL2019051701 from Lishui, Zhejiang, China BB. multicinctus juvenile female, CIB DL18090209 from Fujian, China CB. candidus, adult male, NMW 27711:1 from Bandong, Java DB. candidus, juvenile male, RMNH 11416 from Pelambang, Java EB. wanghaotingi, adult male, CIB MLML20170801 from Mengla, Yunnan, China FB. wanghaotingi, unknown sex juvenile CIB JCR36 from Jiangcheng, Yunnan, China GB. suzhenae sp. nov. adult male, CIB 116089 from Yingjiang, Yunnan, China HB. suzhenae sp. nov. subadult male, CIB 116088 from Yingjiang, Yunnan, China.

https://binary.pensoft.net/fig/521375

The hemipenes of B. candidus is described based on photos of a male (Fig. 9D–F, collecting No. RH06153, total length 120 cm) from Phong Nha-Ke Bang National Park Administration, Quang Binh Province, Vietnam, by Ralf Hendrix. This specimen was tentatively identified as B. candidus by the presence of (1, 2, 4–8) characters in the former morphological description. The hemipenis was partially everted, with large spines present on the medial portion of the organ at the position of the first subcaudal scale; smaller spinous calyces present near the base and another spinous zone present posterior from the row of larger spines. Spinous calyces along organ all elongated, robust at bases and gradually tapering to a tip without distinct bordering. Tips of spines strongly keratinized, semitransparent when fresh, bent towards the base of hemipenes. Sulcus not shown in the photos.

Distribution. This species is known from following localities based on specimens examined and/or DNA sequences data: Java and Sumatra Island, Indonesia; Peninsular Malaysia; Cambodia; Central and Southern Vietnam.

10.3897/zookeys.1025.62305.figure4F20A595D-F4CF-51C1-98F8-E45815FFD493

Dorsal (left) and ventral (right) view of adults of the Bungarus candidus/multicinctus/wanghaotingi complex A, BBungarus multicinctus, adult male, CIB DL2019051701 from Lishui, Zhejiang, China C, DB. candidus, female, NMW 9486:1 from Pelambang, Java E, FB. wanghaotingi, male, CIB MLML20170801 from Mengla, Yunnan, China. Scale bars: 20 mm.

https://binary.pensoft.net/fig/52137210.3897/zookeys.1025.62305.figure588591F69-0455-5126-BDA9-3EB86BDBA638

Dorsal (left) and ventral (right) view of juveniles of the Bungarus candidus/multicinctus/wanghaotingi complex A, BB. multicinctus, female, CIB DL18090209 from Fujian, China C, DB. candidus, female, NMW 27730:4 from Tasikmalaya, Java E, FB. wanghaotingi, unknown sex, CIB JCR36 from Jiangcheng, Yunnan, China. Scale bars: 20 mm.

https://binary.pensoft.net/fig/52137310.3897/zookeys.1025.62305.figure630864D73-CECD-5B74-A231-44541879E280

Dorsolateral head view of adults (left) and juveniles (right) of the Bungarus candidus/multicinctus/wanghaotingi complex and B. suzhenae sp. nov. AB. multicinctus, adult male, CIB DL2019051701 from Lishui, Zhejiang, China BB. multicinctus, juvenile female, CIB DL18090209 from Fujian, China CB. candidus, adult female, NMW 9486:1 from Pelambang, Java DB. candidus, juvenile female, NMW 27730:4 from Tasikmalaya, Java EB. wanghaotingi, adult male, CIB MLML20170801 from Jiangcheng, Yunnan, China FB. wanghaotingi, unknown sex juvenile, CIB JCR36 from Jiangcheng, Yunnan, China GB. suzhenae sp. nov. adult female, CIB 116090 HB. suzhenae sp. nov. subadult male, CIB 116088.

https://binary.pensoft.net/fig/52137410.3897/zookeys.1025.62305.figure9AF808BB8-1BE0-509C-8958-4429574A82B8

Hemipenial morphology of the Bungarus candidus/multicinctus/wanghaotingi complex and B. suzhenae sp. nov. Sulcate view (left), asulcate view (middle), spines (right) A–CB. multicinctus, CIB DL2019051701 from Lishui, Zhejiang, China, body length 993 mm D–FB. candidus, RH06153 from Quang Binh Province, Vietnam, body length 1200 mm G–IB. wanghaotingi, CIB MLML20170801 from Mengla, Yunnan, China, body length 1170 mm J–LB. suzhenae sp. nov., CIB 116089 from Yingjiang, Yunnan, China, body length 1140 mm.

https://binary.pensoft.net/fig/521377Appendix 1

Specimens examined for measurements and morphology; localities as originally stated

B. caeruleus. (9 specimens). “Indes Orientalis” MNHN 7688. “Bengale” MNHN 3952, MNHN 7687. “Malabar” MNHN 7687. “Pondicheri” MNHN 7686. “Pakistan occ” MNHN 1962.239, MNHN 1962.236, MNHN 1962.238, MNHN 1962.237. “Birma” MNHN1893-413.

B. ceylonicus. (1 specimens). “Sri Lanka” MNHN 4259 (1872-32).

B. fasciatus. (1 specimens). “Bengkalis, Siak, Indonesia” RMNH 1667.

B. candidus. (20 specimens). “Palembang, Sumatra, Indonesia” RMNH 11416. “Java, Indonesia” NMW 27730:2, NMW 27730:7, NMW 27730:3, NMW 27730:4; NMW 27740:5, NMW 27740:4, NMW 9486:4, NMW 9486:3, NMW 27730:5, NMW 27730:1, NMW 9486:1, NMW 9486:6, NMW 9486:5, NMW 9486:2, NMW 27711:1, NMW 27711:2, NMW 27711:3. “Johor, Malaysia” SYNU R180411. “Quang Binh, Vietnam” RH06153.

B. multicinctus. China: (24 specimens). “Anhui, China” CIB 12209; “Chongqing, China” CIB 12215. “Fujian, China” CIB 12212, CIB 12204, CIB 12203, CIB 12207, CIB 12206, CIB 12205, CIB DL18090209, CIB DL18090210; “Guangdong, China” CIB 12191, CIB 12194; “Guangxi, China” CIB 12192, CIB 93924, CIB 93923, CIB 104228; “Zhejiang, China” CIB 12208; “Jiangxi, China” CIB 12210, CIB 12211; “Hainan, China” CIB 12197, SYNU R180305; “Guizhou, China” CIB 83793; “Hunan, China” CIB 12213, CIB 12214.

B. wanghaotingi. China: (16 specimens). “Guangxi, China” CIB FCDZ20170806, CIB 104227. “Yunnan, China” CIB 12216, CIB 12201, CIB ML20170801, CIB MLMY20170801, CIB JCR36, CIB JCR36-2, CIB DL2019051401, CIB DL20190525, CIB DL2019070301, CIB DL20190522, CIB JCR2019062003, CIB JCR2019062311, CIB JCR2019061703, CIB JCR2019061807.

B. suzhenae sp. nov. China. (4 specimens). “Yingjiang, Yunnan, China” CIB 116088–CIB 116091.

Specimens checked hemipenes:

B. multicinctus. “Guangdong, China” CIB 12191. “Shangyou, Jiangxi, China” CIB 12211. “Nanning, Guangxi, China” CIB 104228. “Lishui, Zhejiang, China” CIB DL2019051701.

B. wanghaotingi. China. “Luodian, Guizhou, China” CIB 83793. “Beiliu, Guangxi, China” CIB 104227. “Mengla, Yunnan, China” CIB MLML20170801.

B. suzhenae sp. nov. “Yingjiang, Yunnan, China” CIB 116089.

B. candidus “Phong Nha-Ke Bang National Park Administration, Quang Binh Province, Vietnam” RH06153.

Teeth count of some members of subfamily Elapidae in this study.

SpeciesMaxillaPalatinePterygoidDental
Bungarus suzhenae sp. nov. (n = 2) 1+310–119–1015–16
B. candidus 1+4///
B. wanghaotingi (n = 1) 1+412–131016–17
B. multicinctus (n = 1) 1+4121116
B. fasciatus (n = 2) 1+31311–1317
Ophiophagus hannah (n = 3) 1+38–910–1215–16
Naja melanoleuca (n = 3) 1+27–913–1615–16
N. atra (n = 3) 1+17–812–1514–16
Sinomicrurus kelloggi (n = 1) 1+18411
S. macclellandi (n = 1) 1+08915
SlowinskiJB (1994) A Phylogenetic Analysis of Bungarus (Elapidae) Based on Morphological Characters.Journal of Herpetology28: 440–446. https://doi.org/10.2307/1564956SmithMA (1943) The Fauna of British India, Ceylon and Burma, Including the Whole of the Indo-Chinese Sub-Region. Reptilia and Amphibia. 3 (Serpentes). Taylor and Francis, London, 387–390.KuchU (2007) The Effect of Cenozoic Global Change on the Evolution of a Clade of Asian Front-fanged Venomous Snakes: (Squamata: Elapidae: Bungarus). PhD thesis, Frankfurt am Main, Johann Wolfgang Goethe-Universität.