Venomous Snakes of Southern China

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In cooperation with Chinese scien-tists, we were able to study the her-petofauna of southern China, with aspecial focus on the venomoussnakes. More than 30 species of ter-restrial venomous snakes of thefamilies Elapidae and Viperidae arefound in China (ZHAO andADLER, 1993; ZHAO et al., 1998;ZHAO et al., 2000), and most ofthem (except the genera Gloydius,Vipera, and Ermia) are found insouthern China.

The family Elapidae is represent-ed in southern China by members ofthe subfamily Bungarinae: Naja atraCantor, 1842; Naja kaouthia Lesson,1831; Ophiophagus hannah (Cantor,1836); Bungarus fasciatus (Schneider,1801); Bungarus multicinctus Blyth,1860 — and members of the subfamilyElapinae: Hemibungarus macclellandi(Peters, 1862); Hemibungarus kelloggiPope, 1928; Hemibungarus sauteri(Steindachner, 1913).

The family Viperidae is representedin southern China by the single mem-ber of the subfamily Azemiopinae:Azemiops feae Boulenger, 1888 —one member of the subfamilyViperinae: Daboia russelli (Shawand Nodder, 1797) — and manymembers of the subfamily Crotalinae:Deinagkistrodon acutus (Günther,1888); Ovophis gracilis (Oshima,1920); Ovophis monticola (Günther,1864); Ovophis zayuensis (Jiang,1977); Protobothrops jerdoni

The diversity of herpeto-logical fauna in the forestsof eastern and southeast-ern Asia is comparableonly with that of Neo-

tropical forests. Especially rich andextraordinary in reptile and amphib-ian species is an area covering theeastern Himalayas, southern China,and northern Indochina. This in-cludes territories of Assam, India;northern and north-central Myanmar(Burma); southern Xizang (Tibet),southern Sichuan, Yunnan, Guangxi,Guangdong, Hainan, and Taiwanprovinces of China; northern Laos;and northern Vietnam.

Especially interesting in China isthe South Yunnan MountainousSubregion of the South Chinazoogeographical region (ZHAO andADLER, 1993). This is the westernpart of the Tonkin region, compris-ing the Hoang Lien range and theparallel Ailao Shan range, whichtransect Yunnan Province fromnorthwest to southeast. Primarymonsoon and rain forests once cov-ered this territory, but today remainonly in the high mountains and alongsteep slopes. These tropical mon-tane forests support a unique anddiverse herpetofauna, and are ofincreasing interest to zoologists,botanists, and ethnographers.

(Günther, 1875); Protobothropsmucrosquamatus (Cantor, 1839);Protobothrops xiangchengensis(Zhao, Jiang, and Huang, 1978);Trimeresurus albolabris (Gray,1842); Trimeresurus medoensisZhao, 1977; Trimeresurus tibetanusHuang, 1982; Trimeresurus stejnegeriSchmidt, 1925; and Trimeresurusyunnanensis Schmidt, 1925 (ORLOVet al., 2002, 2002a).

In this article, we will present dataon the distribution and biology ofsome of these venomous snakes,based partly on our own field obser-vations, and on captive-maintenanceand breeding experience at TulaExotarium (Breeding Center ofAsian snakes).

Family ElapidaeSSuubbffaammiillyy BBuunnggaarriinnaaeeOOpphhiioopphhaagguuss Günther, 1864, is amonotypic genus distributed in Asia.

OOpphhiioopphhaagguuss hhaannnnaahh (Cantor,1836), the king cobra or hamadryad,was originally classified as thegenus Hamadrias from theSundarbans region and nearCalcutta, India. This is one of themost fascinating venomous snakesin the world. It is the largest ven-omous snake of the Old World, witha maximum total length of 5.5meters. Most are smaller than this,but king cobras of 4–4.5 meters arenot rare in southern and southeast-ern Asia.

Hemibungarus macclellandi. Photo: N. Orlov

Naja atra. Photo: N. Orlov

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Myanmar, Thailand, Laos, Cambodia,Vietnam, southern China includingHong Kong and Hainan, and islandsof Indonesia (Sumatra, Java, andBorneo) (GOLAY et al., 1993;ZHAO and ADLER, 1993; DAVIDand INEICH, 1999).

Bungarus fasciatus is one of themost common and abundant ven-omous snakes. It lives in a variety ofhabitats, including dense jungle, pri-mary stratified forest, secondarybamboo forest at elevations of up to2,500 meters, agricultural land, the

vicinity of human settlements,and even city parks. It does notavoid steep forest slopes or therocky canyons of mountainstreams.Being strongly nocturnal,this snake hides during theday in a variety of shelters,and emerges as darknessfalls. A very active hunter, itis a good swimmer and oftenfound in forest streams atnight. Prey items recordedinclude the buff-striped keel-back, Amphiesma stolata; thebig-tooth snake, Dinodonflavozonatus; the Malayanbanded wolf snake, Lycodonsubcinctus; and an Asiaticwater snake, Sinonatrix sp.We also observed a 1-meterfemale that had eaten two

adult spadefoot toads,Megophrys lateralis.

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Ophiophagus hannah is widelydistributed, from India to Vietnamand from southern China throughIndochina, the Philippines andIndonesia. In southern China itoccurs from Hong Kong and Hainanwestward to Yunnan, southwesternSichuan, and southeastern Xizang(GOLAY et al., 1993; ZHAO andADLER, 1993; MANTHEY andGROSSMAN, 1997; ZHAO et al.,2000).

We have observed Ophiophagushannah in different types of moun-tain forests from foothills to eleva-tions of up to 2,000 meters above sealevel. This species has been recordedin anthropogenic landscapes (vege-tation influenced by human activi-ties) and often inhabits relativelyopen and sunlit woodland. In thickrain and monsoon forests it prefersclearings and riverbanks where thecanopy cover is not fully closed.

The genus name Ophiophagusmeans “snake-eating,” and the kingcobra diet indeed consists largely ofother snakes, as well as other typesof reptiles. In specimens with bodylengths of 3–3.2 meters we foundprey species of snakes: the Indianrat snake, Ptyas korros; the Asiaticwater snake, Sinonatrix trianguligera;the yellowbelly water snake,Enhydris plumbea; the bandedkrait, Bungarus fasciatus; and oth-ers — and large lizards: the EastIndian brown mabuya, Mabuya

multifasciata; and the green prickle-nape, Acanthosaura capra. In cap-tivity, cannibalism was seen.

A gravid female was recorded on22 April in the southern part of theAnnam mountain range. More than30 large females examined at zoos innorthern Vietnam in September andOctober did not have enlarged folli-cles; they were either barren, or hadalready laid eggs. In October andNovember, large king cobras wereseen baskingduringt h e

daytime. In May and Junethis species is crepuscular andnocturnal.

BBuunnggaarruuss Daudin, 1803, is agenus of 13 species inhabitingsouthern and southeastern Asia.

BBuunnggaarruuss ffaasscciiaattuuss (Schneider,1801), the banded krait orpama, was originally classi-fied as the genus Pseudoboa.It attains a maximum totallength of 2 meters or more,although most specimens donot exceed 1.5 meters. Thetail is short and blunt.

Bungarus fasciatus is distrib-uted in the central and north-eastern regions of India,

Mountainous terrain in southern Yunnan, China. Photo: N. OrlovMountain stream on Hainan Island, China. Photo: N. Orlov

Ophiophagus hannah.Photo: N. Orlov


We saw gravid females from the middle of May to themiddle of July. Clutches consist of 3–12 eggs.

BBuunnggaarruuss mmuullttiicciinnccttuuss Blyth, 1860, the many-bandedkrait, is distributed in the southern mainland and TaiwanProvince of China, Laos, Myanmar, and northernVietnam (GOLAY et al., 1993; ZHAO and ADLER,1993; DAVID and INEICH, 1999; MCDIARMID et al.,1999; ORLOV, 1998a; ZHAO et al., 2000).

This species is found near water in plains and hills,especially near rice paddies. It is nocturnal and eatsrats, lizards, other snakes, frogs, fishes (mainly loaches),and eels. In June, the female lays 3–12 eggs, which hatchin about 48 days.

Bungarus multicinctus has highly neurotoxic venom,which is used in traditional medicine, as are the gallbladder, and dried embryos. The meat of this species isalso eaten.

NNaajjaa Laurenti, 1768, is an Afro-Asian genus including morethan 10 species of large,extremely venomous snakes.

NNaajjaa aattrraa Cantor, 1842,the Chinese cobra, is dis-tributed in southern andsoutheastern China in-cluding the islands ofHainan and Taiwan, north-ern Vietnam, and Laos(GOLAY et al., 1993; ZHAOand ADLER, 1993; ORLOV,1998a; DAVID and INEICH,1999).

Naja atra occurs in a wide varietyof habitats, from rice paddies in maritimelowlands to various types of mountain forests. It canlive at elevations of more than 2,000 meters above sealevel. It avoids dark forest with closed canopy. In pri-mary monsoon and rain forests Naja atra inhabits clear-ings and riverbanks. Higher population density is

observed in the vicinity of human settlements, in sec-ondary forests, and in rice paddies adjacent to forest.

Naja atra is diurnal and crepuscular. We saw huntingcobras during all daylight periods and as late as 2–3hours after sunset from March to October, with ambienttemperatures of 20–32°C (68–90°F).

The diet of Naja atra is highly variable. It preys on anyvertebrates from fishes to mammals. Juveniles eat moslyamphibians, whereas adults usually prefer reptiles andmammals — during amphibian breeding periods, how-ever, adult cobras eat mostly frogs: e.g., the paddy frog,Fejervarya limnocharis; the Java whipping frog,Polypedates leucomystax; and the black-spined toad,Bufo melanostictus.

Mating and egg-laying periods are very extended. Weobserved cobras mating in the mountains of the westernTonkin region, at elevations of 400–2,000 meters, in

March through May; gravid females with6–23 eggs were recorded from May to

the end of July.

SSuubbffaammiillyy EEllaappiinnaaeeHHeemmiibbuunnggaarruuss Peters, 1862,is a genus of four species

distributed in India,Nepal, Indochina, south-ern China including theislands of Hainan and

Taiwan, the RyukyuIslands of Japan, and the

Philippines.HHeemmiibbuunnggaarruuss mmaacccclleellllaannddii

(Reinhardt, 1844), the orientalcoral snake, was originally classified

as the genus Elaps, from Assam, India. Itcan grow to a total length of up to 69 centimeters.

The oriental coral snake is distributed in the states ofAssam and Sikkim in India, Myanmar, Thailand,Vietnam, Laos, and southern China including Taiwan(GOLAY et al., 1993; ZHAO and ADLER, 1993;

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Ophiophagus hannah basking. Photo: N. Orlov Hemibungarus kelloggi. Photo: N. Orlov

Bungarus fasciatus. Photo: N. Orlov


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ORLOV, 1998a; DAVIDand INEICH, 1999).

This species inhab-its mountain slopes(sometimes verysteep) of mon-soon and rainforests. We haveobserved it at ele-vations from 400 to2,500 meters. In themountains of the west-ern Tonkin region it prefersareas of karst covered with well-drained, layered forests (treespecies of differing heights andsunlight requirements).

Hemibungarus macclellandi is crepuscular and noc-turnal, being especially active at night. It emerges onrainy nights in the beginning of March, at temperaturesas low as 14°C (57°F). Gravid females were recorded inMay and June at elevations of 800–1,000 meters.Clutches consist of 7–12 eggs. Recorded prey itemsincluded the dwarf snake Calamaria pavimentata; aworm snake, Typhlops sp.; and a ground skink, Scincella sp.

HHeemmiibbuunnggaarruuss kkeellllooggggii Pope, 1928, Kellogg’s coralsnake, is distributed in northern Laos, northernVietnam, and the provinces of Sichuan, Guizhou,Zhejiang, Jiangxi, Fujian, Hainan, and Guangxi in China(GOLAY et al., 1993; ZHAO and ADLER, 1993;ORLOV, 1998a; DAVID and INEICH, 1999).

This nocturnal species is found in mountain forests.We have observed it at elevations from 600 to 1,500meters. It preys on other species of snakes. In May,females lay 5–8 elongate eggs.

Family ViperidaeSSuubbffaammiillyy AAzzeemmiiooppiinnaaeeAAzzeemmiiooppss Boulenger, 1888, is a monotypic genus.

AAzzeemmiiooppss ffeeaaee Boulenger, 1888, Fea’s viper, is one ofthe most mysterious viperid snakes. Until the lastdecade, only a few specimens were known. The specieswas named in honor of prominent Italian naturalistLeonardo Fea of the Natural History Museum inGenoa. During a 4-year field study (1885–1889) inBurma (now Myanmar) he collected insects and otherinvertebrates, amphibians, reptiles, and birds(HALLERMAN et al., 2002). He found Azemiops feaein the Kakhien (=Kakhyen) Hills, several kilometersfrom Bhamo, on the border with Yunnan, China(ORLOV and RYABOV, 2002).

Azemiops feae is distributed in northern Myanmar;the provinces of southeast Xizang, Yunnan, Guangxi,Guizhou, Sichuan, Fujian, Zhejiang, Jiangxi, andShaanxi in China; and the provinces of Vinh-phu, Bac-thai, Cao-bang, and Lang-son of northern Vietnam(GOLAY et al., 1993; ZHAO and ADLER, 1993;ORLOV, 1995, 1997, 1998b; ZHAO et al., 1998; DAVID

and INEICH, 1999;MCDIARMID et al.,

1999; ORLOV et al.,2002; ORLOV andRYABOV, 2002).

We observedthis cryptozoicspecies in themountains ofnorthern Viet-

nam and south-ern China at eleva-

tions from 600 to 1,100meters. POPE (1935) reported the species at elevationsof 600–1,500 meters in China. In addition to our ownobservations, we used a questionnaire and color photo-graphs to survey the local people, especially hunters andforesters. Virtually everyone working regularly in theforest recognized this snake and said it was very rarelyseen on forested mountain slopes. Sightings are proba-bly so rare partly because of the extremely secretivelifestyle of the species. It is interesting to note that mostrecorded sightings are of subadult specimens.

Our observations and data received from the localpeople indicate that in northern Vietnam Azemiops feaeoccurs in habitats characterized by the presence of bam-boo and tree-fern thickets interspersed with sunlit openareas, a thick layer of leaf litter and decomposing treeferns, and karst outcrops, well-drained by numerous sur-face and underground streams. The snake probably

Azemiops feae. Photo: N. Orlov

Deinagkistrodon acutus at the edge of a mountain stream. Photo: N. Orlov


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spends most of its life in the crevicesand galleries of the karst fields,where it can find prey withoutappearing above ground. Limiteddata on feeding in nature show thatthe diet of Azemiops feae includesrodents and shrews (GREEN, 1992)associated with the karst outcropscut by swift mountain streams.

We recorded Azemiops feae inmid May and early June from 3–4a.m. Activity was observed duringlight rainfall, with air temperaturesof 18–20°C (64–68°F) near the soilsurface. The snakes moved slowly,sometimes appearing on the surfaceentirely, and sometimes disappear-ing in the forest litter. The juvenilespecimens found in October andNovember were active on the sur-face at air temperatures of 16–19°C(61–66°F) in the evening, from justafter dark at 7 p.m. until 10–11 p.m.Their appearance on the surface wasalways during light rainfall.

We also have long-term experi-ence with Azemiops feae in captivi-ty. It has been found that this vipercannot tolerate dry conditions.Even with relative air humidityapproaching 100 percent, the sub-strate must be kept very wet (butloose). Otherwise, the snake willlose mobility and its skin will quick-ly become dry and wrinkled.During the active period (Februarythrough November), daytime tem-perature should be kept at 23–27°C(73–81°F); nighttime temperature,at 17–19°C (63–66°F). This snakefeeds at temperatures as low as16–17°C (61–63°F), and is able todigest food at temperatures as lowas 19–20°C (66–68°F). In the loosewet substrate of the terrarium, cap-tive vipers quickly burrow a net-work of underground tunnels andchambers.

During the active period, sheddingin captive specimens occured aboutonce every 6 weeks. Over a periodof 6 years in a terrarium, one femalegrew from 32 centimeters to 98centimeters in total length. Over thesame time period, a male grew from30 centimeters to 92 centimeters.Specimens of such a large size havenot been recorded in the wild.

Hibernation in captivity was set upin very moist substrate at tempera-tures of 13–14°C (55–57°F) for 60days from the end of December untilthe end of February.

During the active period, Fea’svipers ate once every 8–12 days.They were fed mice, newborn rats,lizards (Hemidactylus sp.), andpieces of chicken prepared with thescent of rat or mouse.

Unlike many other terrestrialvipers, Azemiops feae is relativelysluggish and unaggresive. Whenmating, the male aligns himself par-allel with a female, twitches againsther, and turns his tail and cloacaunder hers. The female then lifts hertail, the cloacas are drawn together,and copulation takes place, lastingfor about 10 minutes. Copulationmay be repeated several times over aperiod of 10–12 days (ORLOV andRYABOV, 2002).

The female is gravid for about 90days before laying eggs. We haveobserved two clutches of 5 eggs eachlaid by this species.

SSuubbffaammiillyy CCrroottaalliinnaaeeDDeeiinnaaggkkiissttrrooddoonn Gloyd, 1979, is amonotypic genus.

DDeeiinnaaggkkiissttrrooddoonn aaccuuttuuss (Günther,1888), the Chinese copperhead, wasfirst classified as the genus Halys,from the mountains north ofKinkiang (=Jinjiang Shi), JiangxiProvince, China.

Deinagkistrodon acutus is distrib-uted in Taiwan and central mainlandChina from Zhejiang and Fujianwestward to Guizhou and southeast-ern Sichuan, and southward tonorthern Vietnam, and possiblyLaos (GOLAY et al., 1993; ZHAOand ADLER, 1993; DAVID andINEICH, 1999; MCDIARMID et al.,1999; ZHAO et al., 2000; ORLOVet al., 2002).

This species lives in forested areasof mountains. It is often found onrocks in mountain streams, fromlower hills and mountains to eleva-tions up to 2,000 meters. It is veryactive in the evenings and on rainyor cloudy days. It feeds on frogs,lizards, birds, and rats. On themountain Fan Si Pan (Sa Pa District,


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Lao Cai Province) it has been seenfeeding on bamboo rats (genusRhizomys). Local people indicatethat the adult snakes live in associa-tion with bamboo rat colonies, usingthe rat tunnels for shelter andambush. Snake hunters visit thesecolonies at the proper time, andlocate snakes by listening for thepeeping sound emitted by rats beingattacked in their underground nestchambers. The hunters then dig upthe tunnels and catch the snakes.Deinagkistrodon acutus isused in Vietnam and Chinafor food and in traditionalmedicine.

A female measur-ing about 1 meter intotal length wascaptured at theend of June inMau Son (LangSon Province), atan elevation of 600meters. In captivity, thissnake feeds on frogs, small rats, andother rodents. After eating, it oftenremains immobile for up to 19 daysand 8 hours.

This species is oviparous. Copula-tion takes place in May, and again inSeptember and October. Clutchesof 15–35 eggs are laid between Julyand September (sperm from theautumn copulations is stored untilthe following year). Eggs measure45–55 x 25–30 millimeters, weigh

about 15 grams, and already containdeveloped embryos at the time oflaying. The female guards and incu-bates the eggs. Newly hatchedyoung measure 21.5–24 centimetersin total length. The female aban-dons her young about 24–30 daysafter they hatch. The young snakeseat mostly frogs, but shift to mice asthey grow. ■

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Deinagkistrodon acutus in the forest. Photo: N. Orlov

Bungarus multicinctus. Photo: N. Orlov


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Venomous Snakes of Southern China