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  • Asiatic Cobras Systematics and Snakebite

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    John Gamesby
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    Asiatic Cobras Systematics and Snakebite

    Direitos autorais:

    © All Rights Reserved
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    21 visualizações5 páginas

    Asiatic Cobras Systematics and Snakebite

    Enviado por

    John Gamesby
    Asiatic Cobras Systematics and Snakebite

    Direitos autorais:

    © All Rights Reserved
    Baixe no formato PDF ou leia online no Scribd
    Sinalizar o conteúdo como inadequado
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    Asiatic cobras: Systematics and snakebite W. Wiister and R. S, Thorpe Department of Zoology, University of Aberdeen, Tillydrone Avenue, Aberdeen AB9 2TN (Scotland, UK) Received 25 April 1990; accepted 5 July 1990 ‘Summary. The population affinities of the Asiatic cobras of the genus Naja are investigated, using multivariate analysis of a range of morphological characters. This complex, which was formerly thought to be monospecific, Consists of at least eight full species. In some cases, species whose bites require different antivenoms occur sympatri- cally. The new understanding of the systematics of the Asiatic cobra complex calls for a reappraisal of cobra antivenom use in Asia, and for more research into venom composition, Key words, Naja; cobra; Serpentes; systematics; snakebite; antivenom, ‘The Asiatic cobras of the genus Naja are of considerable ‘medical, social and economic importance in many parts of Asia, killing thousands of people every year. The complex is usually thought to consist of a single species, Naja naja, with ten generally recognized subspecies?~* (ig. 1). Many of these are poorly defined and heteroge- neous, and the population systematics of the group have been a source of confusion for many years, Our study has already shown that some of the conventional subspecies are in fact full species =” ‘The poor understanding of the population systematics of these snakes is regrettable in view of their medical impor- tance: as has been demonstrated in the carpet viper (Eckis carinatus) species complex, closely related and morphologically similar species can have venoms with different antigenic qualities. Consequently, the antiven- ‘om against one species may not neutralize the venom of another®, which can lead to greatly increased fatality rates®!°. The aim of this paper is to summarize the population affinities of the Asiatic cobra complex, and to relate these findings to the literature on the treatment of bite victims and to venom research. This study is based on the multivariate analysis of mor- phological characters recorded from approximately 700 preserved specimens loaned from 29 museums in Europe, the United States and India *#, 66 morphological charac- tet, listed in table 1, relating to scalation, colour pattern, dentition, internal anatomy and body proportions were recorded from each specimen. Specimens were grouped into operational taxonomic units (OTUs) on the basis of collecting gaps and potential physiographic barriers. The homogeneity of the OTUs was confirmed by principal components analysis. Canonical variate analysis (CVA), cone of the most widely used techniques in the analysis of population affinities !?~19, was used for the investiga- tion of the population systematics of the group. ‘The four CVAs, presented in figure 2, show that the Asiatic cobra populations comprise eight distinct taxa, which we regard as separate species: ~ the Indian spectacled cobra, Naja naia, from India and neighbouring areas; ~ the Central Asian cobra, Naja oxiana, ranging from the Caspian Sea to northern India; = the monocellate cobra, Naja kaouthia, which occurs from northern India to Malaysia, Vietnam and the ‘Andaman Islands; = the Chinese/Indochinese spitting cobra, Naja atra, which occurs from eastern China to Thailand. The affinities of the Indochinese populations here included in this species are still unclear, and under further inves- tigation, More than one species may be involved; = the equatorial spitting cobra, Naja swmatrana, from the Malayan Peninsula, equatorial Indonesia and Palawan; 26 Experientia 47 (1991), Birkhiuser Verlag, CH-4010 Basel/Switzeland| Research Articles Figure 1. Distbuton of the conventionally recognind subepecie of [Naja naia?“*. Note the geographically incoherent disebution of Nn. ‘spuai ‘Table. Lst of characters recorded from the preserved Naja material, Detailed descriptions of characters and recording methods have been published benbere 1 No. vail seals 2 No, eubeaudal sales 3 Sof sabeaudals undivided 4 No“euneates 5 No. posterior temporal 66 No. temporals and ucals in contact with pretals 4 No. sale roe at 10th ventral 1 No, sale ows at 20% VS length 9 No. scale rows at 40% VS length 10 No. sale rows at 60% VS length 1 No. scale roms at 80% VS length 12 No. scale rows at level of lst tnt 13 %CS tall segments with two vale roms 14 ‘KCS position of reduction fom 6 10 4 sale tows on al 15 °4CS postion of rection from 8 to 6 sae tows on al 16 “ACS postion of rection from 10 to 8 wae ows on tal 17 SGVS poston of anterior thyroid edse 18 94VS postion of posterior eat ip 19 *4VS postion of system junction 20 S4VS position of anterior Iver tp 21 %4VS position of posto Iver Up 22 %4VS position of anterior pancreas tip 23 4V portion of eye duet ntertine junction 24-960 length of estic duct. 25-4408 postion of saerior tip of right testis 26 46S position of posterior ip of right testis 21 94NS potion of tnteior tp of let tests 28 26S poultion of posterior ip of et ets 29 46S pouition of saterior tip of right Kidney 30 46V' postion of posterior tip of right kidney 31S postion of satenor tp of et kidey 32 46VS postion of posterior tp of et kidney 35 No. pieyeoid tet 436 No. detary teeth 37 %4VS positon of at ventral involved in formation of ligt throat area 38 No. lateral throat spots 39 No. sold teth on mala 40 S4VS length of longest cervical nib 441 SEDs width of hood mark 42 SVS poston of anterior edge of hood mark 4 SVS postion of posterior edge of ood mark ‘44 %¢DS encroachment of ight throat clout onto sides of neck 45 94VS position of anterior edge of largest pair of lateral toa spots 446 SLVS length of largest pair of lateral throat spots 447 %DS encroschment of largest pai of threat spots ont sides of neck 45 93 width of fist ventral baad 4 Length of fromal sale 50 Width of frontal sale 51 Distance betwee anterior edge of frontal sale and posterior ip of otra vale 52 Length of supraccuar sales 53 Lengh of suture between prefontl sales 54 Length of interparietal sure 55 Length of pail sealer 56 Length of fang 57 Length of fang discharge orice {58 Distance from tp of fang to proximal end ofdnchargeerfce $9 Seoutveat length © Tal eng {6 Maximam with of hed across supracalare {62 Distance from soout tip to posterior end of interparietal suture {3 Distance from snout tip to posterior end of lower jw bone (6 Head depth across middle of supraoculrs (65 Head depth fom lower edge of supralabias to top surface of supe ocala (6 Width of widest ventral ele Figure 2, Determination of Asiatic cobra taxa tough the us of CVA, For each CVA, the ordination ofthe speineas along the fist two canon teal variatesis how Pints corespond to OTU means, ines indicate the ‘eater of individual specimens, 4 CVA 1. Ordiaton of cobra popul ‘Sons from India, Central Asa and Indochina, Thee distinct xa a evident a Central Asan taxon, a taxon witha spctacle-shaped hood ‘bark, and a tan witha monocelste mark. B Distbution ofthe Con ‘eal Asin, spctaced and monoctliat aa revealed by CVA 1. Te thre taxa ae parully sympatic and therefore separate specie. CVA 2 (Ordination of obra populations from China and Indochina, Two distact ‘aia areevident:a thon witha monocellatehood mark and taxon With Variable, but soa-monocelte mark. D Distbution ofthe tro cobra taza ffom China and Indochina, revealed by CVA 2. They oceur sy AP ST Oo KR the Malayan Peinsula and Indonesia. Two s+ ‘an equatorial taxon, anda southern Indonesian ‘axon. F Disuibution of the two Malayan and Indonesian cobra taxa revealed by CVA 3. Because of consistent and considerable morpholop ‘i ferences, we consider tem o be separate species. CVA &: Ord ‘ation ofthe cobra populations af the Philippines and equatorial Indone- Sa. Thee distinet taxa ae evident the equatorial cobra, a soutbeastern ‘axon, anda pore taxon. Hf Ditton ofthe thee Philippine cobra tana revealed by CVA 4 The three forms are considered separate speci, since they constitute phenetialy cohesive and highly distinct ana (Ma ‘Marindugue; Mb: Masbate; BR: Bobo), 208 [Experientia $7 (199), Bikhiuser Verlag, CH-A010 Base/Switzeland Research Articles ai i a il Figure 3, Distribution of the eight Asai cobra specie. ~ the southern Indonesian spitting cobra, Naja sputatrix, from Java and the Lesser Sunda Islands; ~ the southeastern Philippine spitting cobra, Naja sa- ‘marensis, from Mindanao, Samar, Leyte, Bohol and Camiguin; = the northern Philippine cobra, Naja philippinensis, from Luzon, Mindoro, Masbate, and Marinduque. Their distribution is shown in figure 3. The extensive areas of sympatry between several pairs of species are clearly evident. Cobras are an important cause of snakebite morbidity and mortality in much of tropical Asia ®=!*. Cobra bites in humans result in a variety of symptoms, the most important being potentially fatal neurotoxic effects, and local tissue damage, which can be crippling. The relative incidence of these symptoms varies enormously between, and, toa lesser extent, within the newly recognized Asiat- ic cobra species 9-2! (table 2), ‘These enormous differences between these species indi- cate major differences in the composition of their ven- coms. Antigenic differences between these venoms are therefore likely, so that an antivenom against one species ‘ay be ineffective against the venom of another. Such problems have been demonstrated in some cases: Indian antivenom (manufactured on the basis of Naja raja venom) shows poor neutralizing ability against the venom of N. philippinensis*®, and antivenom against co- bras from southern Malaysia (. swmatrana) is ineffec- tive against the venom of N. kaouthia"®. Chinese N. atra ‘venom appears to be poorly neutralized by most com- aercially available cobra antivenoms?® These problems illustrate the need for a sound under- standing of the population systematics of venomous snakes. N. swmatrana and N. kaouthia, which are known to require different antivenoms, occur sympatrically in parts of the Malayan Peninsula® (fig. 3). N. kaouthia, and the Indochinese spitting cobras here referred to N. atra, occur sympatrically in much of Indochina. The te- ported differences in venom composition between . ‘kaouthia and the Chinese populations of N. atra?? sug- ‘Table 2. Geographic variation inthe elaive incidence ofthe symptoms of Aste cobra bites Tease Secass ‘Ratio swith netro- with eure (orc necrosis foie symptoms esos Tazoa (a= 39) mm 7% 1a (N.phiippnenss) C.Thaiand?® (a = 24 sim 30% sat (outa) N. Malaysia (a = 47) me am 137 WY. kaoutia and 1 smatrana) ‘Tie nomencatre faa been modified from the original referenes, and altered to the one proposed inthis paper. Many paints experienced father necrosis no bewotoxcty, symptoms being abset or Lied to local seeling andor pain. In the study trom northern Malaysia", wo patents were ten by Nal mamatrana the reminder probably al by ‘aja Rao Research Articles ‘gest that there may be problems with regard to antiven- om efficacy in Indochina. In both cases, no polyvalent antivenom against the venoms of both species involved is available, since it had not hitherto been realized that these forms are separate, sympatric species. Problems may also occur in countries with several non- sympatric species of cobra, since the antivenom manu- facturers generally produce antivenom only on the basis of the venom of snakes from their immediate vicinity. In the Philippines, only antivenom against N. philippinensis is produced, even though two other species (N. samaren- sis and N. swnatrana) occur in the country, and in In-

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