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Staff Profile of Dr Wolfgang Wüster

Dr Wolfgang Wüster
Senior Lecturer
+44(0)1248 382301
School of Biological Sciences, University of Wales, Bangor, Deiniol Road, Bangor, Gwynedd, LL57 2UW, UK


Venomous snakes are among the organisms provoking the strongest emotions in humans, ranging from intense fascination to visceral revulsion. Some represent serious public health problems, and yet their venoms offer the prospect of developing new drugs against common medical conditions. All are scientifically fascinating, providing opportunities to study the evolution of a chemical weapon system, its role in the biology of the animal and its interactions with other occupants of the snake’s habitat. At the same time, we know surprisingly little about the biodiversity of venomous snakes, and new species are still being discovered on a regular basis.

My research interests focus on the causes of variation in venom composition within species and between closely related species, the origin and evolution of venom and venom toxin families, the biodiversity of venomous snakes, the wider impact of venom on the interactions between snakes and other biota, and the biogeographical history of different groups.



Since 2012: Senior Lecturer in Zoology, School of Biological Sciences, Bangor University

1999-2012: Lecturer n Zoology, School of Biological Sciences, Bangor University

1995-1999: Wellcome Trust Research Fellow, School of Biological Sciences, Bangor University

1994-1995: Postdoc, School of Biological Sciences, Bangor University

1990-1993: NERC Postdoctoral Fellow, Department of Zoology, University of Aberdeen

1985-1986: Technician, MRC Radiobiology Unit, Harwell


PhD, 1990, University of Aberdeen: Population evolution of Asiatic cobras.

BA (Hons), II.1, 1985, Natural Sciences Tripos (Zoology), University of Cambridge.


Origin and evolution of snake venom and its consequences.

Snakes are unique among terrestrial vertebrates in possessing a chemical arsenal that, in at least some cases, allows them not only to increase their foraging success, but also to seriously injure or kill a potential predator. As such, this weapon system is likely to have had profound consequences for the evolution and diversification of snakes and their interactions with other organisms. Understanding the origin, continuing evolution and consequences of the biochemical arsenal of snakes is one of my main research goals, and follows three primary lines of investigation:

Origin of snake venom and venom toxins: using phylogenetic analysis of toxin amino acid sequences and related non-toxin genes allows the history of toxin gene families to be mapped on the phylogenetic tree of snakes. This has already lead to new insights into the origin of major toxin families (e.g., Fry & Wüster, Mol. Biol. Evol., 2004), the unexpectedly dynamic evolutionary model of toxin families (Casewell et al., Nature Comms., 2012) and the role of gene duplications and to co-opting of the pancreatic microRNA apparatus for the regulation of venom secretion (Vonk et al., PNAS, 2013).

Evolution of snake venom: venom composition varies at all taxonomic levels in snakes, and provides ample raw material to test hypotheses on the causes and consequences of snake venom variation. Compositional studies and experiments with natural prey have shown an association between snake venoms and diet composition (Daltry, Wüster & Thorpe, Nature, 1996; Barlow et al., Proc. R. Soc. London B, 2009; Richards et al., Toxicon, 2012). Advances in molecular genetics and proteomics now allow us to determine the mode of evolution of toxins and their impact on the activity of these toxins, and relate these to possible adaptive explanations. Current work also focusses on the role of gene flow and hybridisation as drivers of venom variation in rattlesnakes.

Impact of venom on snake evolution and diversification: this line of investigation on one hand use phylogenetics and comparative morphological studies to  test hypotheses on the use and adaptive evolution of venom and the venom apparatus under different selective regimes (e.g., dietary differences), and the interactions between venomous snakes and their predators. In particular, field studies have demonstrated that mimicry of venomous snakes is not restricted to brightly coloured models, but that the less conspicuous but nevertheless characteristic patterns of some venomous snakes also protect them from predatory attack, demonstrating that the protective umbrella of mimicry encompasses a higher proportion of snake diversity than previously known (Wüster et al., Proc. R. Soc. London B, 2004).


Phylogeographic patterns and biogeography of tropical snakes

Comparing phylogeographic patterns across multiple co-distributed species complexes can reveal shared patterns that can test for commonality of cause for shared patterns. Molecular data are particularly powerful in biogeographical analysis since they allow the absolute timing of lineage splits to be estimated. Current projects include investigations of zoogeographic relationships between Australia and New Guinea (Wüster et al., Mol. Phylogenet. Evol., 2005), testing biogeographical hypotheses in the Amazon Basin (Wüster et al., Mol. Ecol., 2005), and the Middle East (Pook et al., Mol Phylogenet. Evol., 2009) and phylogeographic patterns in widespread African venomous snakes (Wüster et al., Mol. Phylogenet. Evol., 2007).

Systematic revisions of venomous snakes

Understanding the systematics and phylogeny of venomous snakes provides the essential underpinnings to all other work on these animals, and constitutes a fundamental first step towards the rationalisation of antivenom manufacture and use. Using molecular (mtDNA and AFLP) and advanced multivariate morphological methods, I am revising the systematics of several groups of medically important venomous snakes, including the saw-scaled vipers (Echis), African and Asiatic cobras (Naja), and several Australasian elapids (Acanthophis and Pseudechis). This work is leading to a radical re-assessment of the systematic of several species groups, and has led to the description of five new species, including three cobras.


Recent and other Other Major Publications:

Ujvari, B., N.R. Casewell, K. Sunagar, K. Arbuckle, W. Wüster, N. Lo, D. O’Meally, C. Beckmann, G.F. King, E. Deplazes & T. Madsen (2015) Widespread convergence in toxin resistance by predictable molecular evolution. PNAS, 112(38): 11911-11916. PDF:

Maddock, S.T., R.J. Ellis, P. Doughty, L.A. Smith & W. Wüster (2015) A new species of death adder (Acanthophis: Serpentes: Elapidae) from north-western Australia. Zootaxa, 4007(3): 301-326. PDF:

Rhodin, A.G.J., H. Kaiser, P.P. van Dijk, W. Wüster, M. O’Shea, M. Archer, M. Auliya, L. Boitani, R. Bour, V. Clausnitzer, T. Contreras-MacBeath, B.I. Crother, J.M. Daza, C.A. Driscoll, O. Flores-Villela, J. Frazier, U. Fritz, A. Gardner, C. Gascon, A. Georges, F. Glaw, F.G. Grazziotin, C.P. Groves, G. Haszprunar, P. Havaš, J.M. Hero, M. Hoffmann, M.S. Hoogmoed, B.D. Horne, J.B. Iverson, M. Jäch, C.L. Jenkins, R.K.B. Jenkins, A.R. Kiester, J.S. Keogh, T.E. Lacher Jr., J.E. Lovich, L. Luiselli, D.L. Mahler, D. Mallon, R. Mast, R.W. Mcdiarmid, J. Measey, R.A. Mittermeier, S. Molur, V. Mossbrugger, R. Murphy, D. Naish, M. Niekisch, H. Ota, J.F. Parham, M.J. Parr, N.J. Pilcher, R.H. Pine, A.B. Rylands, J.G. Sanderson, J. Savage, W. Schleip, G.J. Scrocchi, H.B. Shaffer, E.N. Smith, R. Sprackland, S.N. Stuart, H. Vetter, L.J. Vitt, T. Waller, G. Webb, E.O. Wilson, H. Zaher, and S. Thomson (Corresponding Author). 2015. Comment on Spracklandus Hoser, 2009 (Reptilia, Serpentes, ELAPIDAE): request for confirmation of the availability of the generic name and for the nomenclatural validation of the journal in which it was published. (Case 3601; see BZN 70: 234–237; 71: 30–38, 133–135, 181–182, 252–253). Bulletin of Zoological Nomenclature 72(1): 65-78. PDF:

Casewell, N.C., S.C. Wagstaff, W. Wüster, D.A.N. Cool, F.M.S. Bolton, S.I. King, D. Pla, L. Sanz, J.J. Calvete & R.A. Harrison (2014) Medically important differences in snake venom composition are dictated by distinct post-genomic mechanisms. PNAS, 111: 9205-9210.

Vonk, F.J., N.R. Casewell, C.V. Henkel, A. Heimberg, R.J.R. McCleary, H.M.E. Kerkkamp, R. Vos, E. McGlinn, I. Guerreiro, J.J. Calvete, W. Wüster, J.M. Logan, A.E. Woods, R.A. Harrison, T.A. Castoe, A.P.J. de Koning, D.D. Pollock, C. Renjifo, R.B. Currier, D. Salgado, D. Pla, L. Sanz, H.P. Spaink, H.J. Jansen, A.S. Hyder, J.M.C. Ribeiro, P.W. Arntzen, G.E.E.J. van den Thillart, M. Boetzer, W. Pirovano, R.P. Dirks, D. Duboule, R.M. Kini & M.K. Richardson (2013) King cobra genome reveals dynamic gene evolution and adaptation in the snake venom system. PNAS 110 (51): 20651-20656. PDF:

Casewell, N.R., W. Wüster, F.J. Vonk, R.A. Harrison & B.G. Fry (2013) Complex cocktails: the evolutionary novelty of venoms. Trends in Ecology and Evolution. 28: 219-229. PDF:

Barlow, A., K. Baker, C.R. Hendry, L. Peppin, T. Phelps, K.A. Tolley, C.E. Wüster & W. Wüster (2013) Phylogeography of the widespread African puff adder (Bitis arietans) reveals multiple Pleistocene refugia in southern Africa. Molecular Ecology, 22: 1134-1157. PDF:

Kaiser, H., B.I. Crother, C.M.R. Kelly, L. Luiselli, M. O’Shea, H. Ota, P. Passos, W.D. Schleip & W. Wüster (2013) Best practices: in the 21st Century, taxonomic decisions in herpetology are acceptable only when supported by a body of evidence and published via peer-review. Herpetological Review 44: 8-23. PDF:

Casewell, N.R., G.A. Huttley & W. Wüster (2012) Dynamic evolution of venom proteins in squamate reptiles. Nature Communications, 3(1066): 1-10. PDF:

Barlow, A., W. Grail, M. de Bruyn & W. Wüster (2012) Anonymous nuclear markers for the African adders (Serpentes: Viperidae: Bitis). Conservation Genetics Resources, 4:967-969.  PDF:

Fry, B.G.,, N.R. Casewell, W. Wüster, N. Vidal, B. Young & T.N.W. Jackson (2012) The structural and functional diversification of the Toxicofera reptile venom system. Toxicon, 60: 434-448.  PDF:

Richards, D.P., A. Barlow & W. Wüster. (2012) Venom lethality and diet: Differential responses of natural prey and model organisms to the venom of the saw-scaled vipers (Echis), Toxicon, 59: 110-116.  PDF:

Casewell, N.R., S.C. Wagstaff, R.A. Harrison, C.Renjifo & W. Wüster (2011) Domain loss facilitates accelerated evolution and neofunctionalization of duplicate snake venom metalloproteinase toxin genes. Molecular Biology and Evolution 28: 2637-2649. PDF:

Williams, D.J., J.-M. Gutiérrez, J.J. Calvete, W. Wüster, K. Ratanabanangkoon, O. Paiva, N.I. Brown, N.R. Casewell, R.A. Harrison, P.D. Rowley, M. O’Shea, S.D. Jensen, K.D. Winkel & D.A. Warrell (2011) Ending the drought: new strategies for improving the flow of affordable, effective antivenoms in Asia and Africa. Journal of Proteomics, 74: 1735-1767.  PDF:

Casewell, N.R., S.C. Wagstaff, R.A. Harrison & W. Wüster (2011) Gene tree parsimony of multi-locus snake venom protein families reveals species tree conflict as a result of multiple parallel gene loss. Molecular Biology and Evolution, 28: 1157–1172.  PDF:

Wüster, W. & Bérnils, R.S. (2011) On the generic classification of the rattlesnakes, with special reference to the Neotropical Crotalus durissus complex (Squamata: Viperidae). Zoologia 28: 417–419. PDF:

Casewell, N.R., D.A.N. Cook, S.C. Wagstaff, A. Nasidi, N. Durfa, W. Wüster & R.A. Harrison (2010) Pre-clinical assays predict pan-African Echis viper efficacy for a species-specific antivenom. PLoS Neglected Tropical Diseases, 4: e851. DOI: 10.1371/journal.pntd.0000851

Antonelli, A., A. Quijada-Mascareñas, A.J. Crawford, J.M. Bates, P.M. Velazco & W. Wüster (2010) Molecular studies and phylogeography of Amazonian tetrapods and their relation to geological and climatic models. In Amazonia: Landscape and Species Evolution (C. Hoorn, F. Wesselingh & H. Vonhoff, eds.), p. 386-404. Blackwell Publishing, Oxford.  PDF:

Casewell, N.R., R.A. Harrison, W. Wüster & S.C. Wagstaff (2009) Comparative venom gland transcriptome surveys of the saw-scaled vipers (Viperidae: Echis) reveal substantial intra-family gene diversity and novel venom proteins. BMC Genomics 10: 564.  PDF:

Pook, C.E., U. Joger, N. Stümpel & W. Wüster (2009) When continents collide: phylogeny, historical biogeography and systematics of the medically important viper genus Echis (Squamata: Serpentes: Viperidae). Molecular Phylogenetics and Evolution 53: 792-807.  PDF:

Trape, J.-F., L. Chirio, D.G. Broadley & W. Wüster (2009) Phylogeography and systematic revision of the Egyptian cobra (Serpentes: Elapidae: Naja haje) species complex, with the description of a new species from West Africa. Zootaxa 2236: 1-25.  PDF:

Wallach, V., W. Wüster & D.G. Broadley (2009) In praise of subgenera: taxonomic status of cobras of the genus Naja Laurenti (Serpentes: Elapidae). Zootaxa 2236: 26-36.  PDF:

Barlow, A., C.E. Pook., R.A. Harrison & W. Wüster (2009) Co-evolution of diet and prey-specific venom activity supports the role of selection in snake venom evolution. Proceedings of the Royal Society B, 276: 2443-2449.  PDF:

Wüster, W., L. Peppin, C.E. Pook & D.E. Walker (2008) A nesting of vipers: phylogeny, historical biogeography and patterns of diversification of the Viperidae (Squamata: Serpentes). Molecular Phylogenetics and Evolution 49: 445-459.  PDF:

Williams, D.J., M. O'Shea, R.L. Daguerre, C.E. Pook, W. Wüster, C.J. Hayden, J.D. McVay, O. Paiva, T. Matainaho, K.D. Winkel & C.C. Austin (2008) Origin of the eastern brownsnake, Pseudonaja textilis (Duméril, Bibron and Duméril) (Serpentes: Elapidae: Hydrophiinae) in New Guinea: evidence of multiple dispersals from Australia, and comments on the status of Pseudonaja textilis pughi Hoser 2003. Zootaxa 1703: 47-61.  PDF:

Wüster, W., S. Crookes, I. Ineich, Y. Mane, C.E. Pook, J.-F. Trape & D.G.Broadley (2007) The phylogeny of cobras inferred from mitochondrial DNA sequences: evolution of venom spitting and the phylogeography of the African spitting cobras (Serpentes: Elapidae: Naja nigricollis complex). Molecular Phylogenetics and Evolution 45: 437-453.  PDF:

Wüster, W. & D.G. Broadley (2007) Get an eyeful of this: a new species of giant spitting cobra from eastern and north-eastern Africa (Squamata: Serpentes: Elapidae: Naja). Zootaxa 1532: 51-68.  PDF:

Quijada-Mascareñas, J.A., J.E. Ferguson, C.E. Pook, M.G. Salomão, R.S. Thorpe & W. Wüster (2007) Phylogeographic patterns of trans-Amazonian vicariants and Amazonian biogeography: the Neotropical rattlesnake (Crotalus durissus complex) as an example. Journal of Biogeography 34: 1296–1312.   PDF:

Williams, D., W. Wüster & B.G. Fry (2006) The good, the bad and the ugly: Australian snake taxonomists and a history of the taxonomy of Australia’s venomous snakes. Toxicon 48: 919-930. PDF:

Wüster, W., J.E. Ferguson, J.A. Quijada-Mascareñas, C.E. Pook, M.G. Salomão & R.S. Thorpe 2005) Tracing an invasion: landbridges, refugia and the phylogeography of the Neotropical rattlesnake (Serpentes: Viperidae: Crotalus durissus). Molecular Ecology 14: 1095-1108. PDF:

Wüster, W., M.R. Duarte & M.G. Salomão (2005) Morphological correlates of incipient arboreality and ornithophagy in island pitvipers, and the phylogenetic position of Bothrops insularis. Journal of Zoology, London 266: 1-10. PDF:

Wüster, W., A.J. Dumbrell, C. Hay, C.E. Pook, D.J. Williams & B.G. Fry (2005) Snakes across the Strait: Trans-Torresian phylogeographic relationships in three genera of Australasian snakes (Serpentes: Elapidae: Acanthophis, Oxyuranus and Pseudechis). Molecular Phylogenetics and Evolution 34: 1-14. PDF:

Wüster, W. C.S.E. Allum, I.B. Bjargardóttir, K.L. Bailey, K.J. Dawson, J. Guenioui, J. Lewis, J. Mcgurk, A.G. Moore, M. Niskanen & C.P. Pollard (2004) Do aposematism and Batesian mimicry require bright colours? A test, using European viper markings. Proceedings of the Royal Society of London, Biological Sciences 271: 2495 - 2499. PDF:

Fry, B.G. & W. Wüster (2004) Assembling an arsenal: origin and evolution of the snake venom proteome inferred from phylogenetic analysis of toxin sequences. Molecular Biology and Evolution 21: 870–883.  PDF:

Broadley, D.G. & W. Wüster (2004) A review of the southern African ‘non-spitting’ cobras (Serpentes: Elapidae: Naja). African Journal of Herpetology 53:101-122.   PDF:

Fry, B.G., N.G. Lumsden, W. Wüster, J.C. Wickramaratna, W.C. Hodgson & R.M. Kini (2003)  Isolation of a neurotoxin (alpha-colubritoxin) from a ‘non-venomous’ colubrid: evidence for early origin of venom in snakes. Journal of Molecular Evolution, 57: 446-452.   PDF:

Fry, B.G., W. Wüster, S.F.R. Ramjan, T. Jackson, P.  Martelli & R.M. Kini (2003) Analysis of Colubroidea snake venoms by liquid chromatography with mass spectrometry: evolutionary and toxinological implications. Rapid Communications in Mass Spectrometry 17: 2047-2062.  PDF:

Fry, B.G., W. Wüster, R.M. Kini, V. Brusic, A. Khan, D. Venkataraman & A.P. Rooney (2003). Molecular evolution and phylogeny of snake venom three finger toxins. Journal of Molecular Evolution 57: 110-129.   PDF:

Fry, B.G., K.D. Winkel, J.C. Wickramaratna, W.C. Hodgson & W. Wüster (2003) Effectiveness of snake antivenom: species and regional venom variation and its clinical impact. Journal of Toxicology -Toxin Reviews 22: 23-34.  PDF:

Wüster, W. & D.G. Broadley (2003) A new species of spitting cobra from northeastern Africa (Serpentes: Elapidae: Naja). Journal of Zoology, London 259: 345-359.   PDF:

Harrison, R.A., W. Wüster & R.D.G. Theakston (2003) The conserved structure of snake venom toxins confers extensive immunological cross-reactivity to toxin-specific antibody.Toxicon 41: 441-449.

Wüster, W., M.G. Salomão, J.A. Quijada-Mascareñas, R.S. Thorpe & B.B.B.S.P. (2002) Origin and evolution of the South American pitviper fauna: evidence from mitochondrial DNA sequence analysis. In Biology of the Vipers (G.W. Schuett, M. Höggren, M.E. Douglas & H.W. Greene, eds.), pp. 111-128. Eagle Mountain Publishing, Eagle Mountain, Utah.   PDF:

Wüster, W., R.S. Thorpe, M.G. Salomão, L. Thomas, G. Puorto, R.D.G. Theakston & D.A. Warrell (2002) Origin and phylogenetic position of the Lesser Antillean species of Bothrops (Serpentes: Viperidae): biogeographical and medical implications. Bulletin of the Natural History Museum London (Zoology), 68: 101-106.   PDF:

Daltry, J.C. & W. Wüster (2002) A new species of wolf snake (Serpentes: Colubridae: Lycodon) from the Cardamom Mountains, southwestern Cambodia. Herpetologica, 58: 498-504.   PDF:

Fry, B.G., J. Wickramaratna, W. Hodgson, P. Alewood, R.M. Kini, H. Ho & W. Wüster (2002) Electrospray liquid chromatography/mass spectrometry fingerprinting of Acanthophis (death adder) venoms: taxonomic and toxinological implications. Rapid Communications in Mass Spectrometry, 16: 600-608.   PDF:

Puorto, G., M.G. Salomão, R.D.G. Theakston, R.S. Thorpe, D.A. Warrell & W. Wüster (2001) Combining mitochondrial DNA sequences and morphological data to infer species boundaries: phylogeography of lanceheaded pitvipers in the Brazilian Atlantic forest, and the status of Bothrops pradoi (Squamata: Serpentes: Viperidae). Journal of Evolutionary Biology, 14: 527-538.   PDF:

Wüster, W., J.L. Yrausquin & A. Mijares-Urrutia (2001) A new species of indigo snake from northwestern Venezuela (Serpentes: Colubridae: Drymarchon). Herpetological Journal, 11: 157-165.   PDF:

Wüster, W., B. Bush, J.S. Keogh, M. O'Shea & R. Shine (2001) Taxonomic contributions in the "amateur" literature: comments on recent descriptions of new genera and species by Raymond Hoser. Litteratura Serpentium, 21: 67-79, 86-91.  PDF:

Pook, C.E., W. Wüster & R.S. Thorpe (2000) Historical biogeography of the western rattlesnake (Serpentes: Viperidae: Crotalus viridis), inferred from mitochondrial DNA sequence information. Molecular Phylogenetics and Evolution, 15: 269-282.   PDF:

Slowinski, J.B. & W. Wüster (2000) A new cobra (Elapidae: Naja) from Myanmar (Burma). Herpetologica, 56: 257-270.   PDF:

Wüster, W., M.G. Salomão, G.J. Duckett, R.S. Thorpe & BBBSP (1999) Mitochondrial DNA evolution of the Bothrops atrox species complex (Squamata: Serpentes: Viperidae). Kaupia, 8: 135-144.   PDF:

Salomão, M.G., W. Wüster, R.S. Thorpe & BBBSP (1999) MtDNA evolution in Neotropical pitvipers of the genus Bothrops (Squamata: Serpentes: Viperidae). Kaupia, 8: 127-134.   PDF:
Lenk, P. & W. Wüster (1999) A multivariate approach to the systematics of Italian rat snakes of the Elaphe longissima complex (Reptilia, Colubridae): revalidation of Camerano's Callopeltis longissimus var. lineata. Herpetological Journal 9: 153-162.   PDF:

Wüster, W., P. Golay & D.A. Warrell (1999) Synopsis of recent developments in venomous snake systematics, No. 3. Toxicon 37: 1123-1129.   PDF:

Wüster, W., P. Golay & D.A. Warrell (1998) Synopsis of recent developments in venomous snake systematics, No. 2. Toxicon, 36: 299-307.    PDF:

Daltry, J.C., T. Ross, R.S. Thorpe & W. Wüster (1998) Evidence that humidity influences snake activity patterns: a field study of the Malayan pit viper Calloselasma rhodostoma. Ecography, 21: 25-34.    PDF:

Daltry, J.C., W. Wüster & R.S. Thorpe (1998) Intraspecific variation in the feeding ecology of the crotaline snake Calloselasma rhodostoma in Southeast Asia. Journal of Herpetology, 32: 198-205.    PDF:

Wüster, W., D.A. Warrell, M.J. Cox, P. Jintakune & J. Nabhitabhata (1997) Redescription of Naja siamensis Laurenti, 1768 (Serpentes: Elapidae), a widely overlooked spitting cobra from Southeast Asia: geographic variation, medical importance and designation of a neotype. Journal of Zoology, 243: 771-788.   PDF:

Wüster, W., P. Golay & D.A. Warrell (1997) Synopsis of recent developments in venomous snake systematics. Toxicon, 35: 319-340.   PDF:

Wüster, W., M.G. Salomão, R.S. Thorpe, G. Puorto, M.F.D. Furtado, S.A. Hoge, R.D.G. Theakston, & D.A. Warrell (1997) Systematics of the Bothrops atrox species complex: insights from multivariate analysis and mitochondrial DNA sequence information. In R.S. Thorpe, W. Wüster & A. Malhotra (Eds.), Venomous Snakes: Ecology, Evolution and Snakebite. Symposia of the Zoological Society of London, No. 70, Pp. 99-113. Clarendon Press, Oxford.    PDF:

Salomão, M.G., W. Wüster, R.S. Thorpe & BBBSP (1997) DNA evolution of South American pitvipers of the genus Bothrops. In R.S. Thorpe, W. Wüster & A. Malhotra (Eds.), Venomous Snakes: Ecology, Evolution and Snakebite. Symposia of the Zoological Society of London, No. 70, Pp. 89-98. Clarendon Press, Oxford.    PDF:

Daltry, J.C., W. Wüster & R.S. Thorpe (1997) The role of ecology in determining venom variation in the Malayan pit viper. In R.S. Thorpe, W. Wüster & A. Malhotra (Eds.), Venomous Snakes: Ecology, Evolution and Snakebite. Symposia of the Zoological Society of London, No. 70, Pp. 155-171. Clarendon Press, Oxford.    PDF:

Belt, P., D.A. Warrell, A. Malhotra, W. Wüster & R.S. Thorpe (1997) Russell's viper in Indonesia: snakebite and systematics. In R.S. Thorpe, W. Wüster & A. Malhotra (Eds.), Venomous Snakes: Ecology, Evolution and Snakebite. Symposia of the Zoological Society of London, No. 70, Pp. 219-234. Clarendon Press, Oxford.   PDF:

Daltry, J.C., W. Wüster & R.S. Thorpe (1996) Diet and snake venom evolution. Nature, 379: 537-540. PDF:

Wüster, W. (1996) Taxonomic changes and toxinology: systematic revisions of the Asiatic cobras (Naja naja species complex). Toxicon, 34: 399-406.   PDF:

Wüster, W., R.S. Thorpe, G. Puorto & BBBSP (1996) Systematics of the Bothrops atrox complex (Reptilia: Serpentes: Viperidae) in Brasil: a multivariate analysis. Herpetologica, 52: 263-271.    PDF:

Daltry, J.C., G. Ponnudurai, C.K. Shin, N.-H. Tan, R.S. Thorpe & W. Wüster (1996) Electrophoretic profiles and biological activities: intraspecific variation in the venom of the Malayan pit viper (Calloselasma rhodostoma) Toxicon, 34: 67-79.    PDF:

Wüster, W. & C.J. McCarthy (1996) Venomous snake systematics: implications for snakebite treatment and toxinology. In C. Bon & M. Goyffon (Eds.), Envenomings and their Treatments: 13-23. Fondation Mérieux, Lyon.   PDF:

Wüster, W., R.S. Thorpe, M.J. Cox, P. Jintakune & J. Nabhitabhata (1995) Population systematics of the snake genus Naja (Reptilia: Serpentes: Elapidae) in Indochina: multivariate morphometrics and comparative mitochondrial DNA sequencing (cytochrome oxidase I). Journal of Evolutionary Biology, 8: 493-510.   PDF:

Thorpe, R.S., A. Malhotra, H. Black, J.C. Daltry & W. Wüster (1995) Relating geographic pattern to phylogenetic process. Philosophical Transactions of the Royal Society of London, B., 349: 61-68.    PDF:

Theakston, R.D.G., G.D. Laing, C.M. Fielding, A. Freire Lascano, J.-M. Touzet, F. Vallejo, R.H. Guderian, S.J. Nelson, W. Wüster, A.M. Richards, J. Rumbea Guzmán & D.A. Warrell. (1995) Treatment of snake bites by Bothrops species and Lachesis muta in Ecuador: laboratory screening of candidate antivenoms. Transactions of the Royal Society of Tropical Medicine and Hygiene, 89: 550-554.   PDF:

Wüster, W. & R.S. Thorpe (1994) Naja siamensis, a cryptic species of venomous snake revealed by mtDNA sequencing. Experientia, 50: 75-79.    PDF:

Thorpe, R.S., R.P. Brown, M. Day, A. Malhotra, D.P. McGregor & W. Wüster (1994) Testing ecological and phylogenetic hypotheses in microevolutionary studies: an overview. In P. Eggleton and R. Vane-Wright (Eds.): Phylogenetics and Ecology: 189-206. Academic Press, London.    PDF:

Wüster, W. & R.S. Thorpe (1992) Dentitional phenomena in cobras revisited: fang structure and spitting in the Asiatic species of Naja (Serpentes: Elapidae). Herpetologica, 48: 424-434.    PDF:

Wüster, W. & R.S. Thorpe (1992) Asiatic cobras: population systematics of the Naja naja species complex (Serpentes: Elapidae) in India and Central Asia. Herpetologica, 48: 69‑85.    PDF:

Wüster, W. & M.J. Cox (1992) Defensive hemipenis display in the Kukri snake Oligodon cyclurus. Journal of Herpetology, 26: 238-241.    PDF:

Wüster, W., S. Otsuka, A. Malhotra & R.S. Thorpe (1992) Population systematics of Russell's viper: a multivariate study. Biological Journal of the Linnean Society, 47: 97-113.    PDF:

Wüster, W. & R.S. Thorpe (1991) Asiatic cobras: systematics and snakebite. Experientia, 47: 205‑209.    PDF:

Thorpe, R.S., R.P. Brown, A. Malhotra & W. Wüster (1991) Geographic variation and population systematics: distinguishing between ecogenetics and phylogenetics. Bolletino di Zoologia, 58:329-335.    PDF:

Wüster, W. & R.S. Thorpe (1990) Systematics and biogeography of the Asiatic cobra (Naja naja) species complex in the Philippine Islands. In G. Peters & R. Hutterer (Eds.), Vertebrates in the Tropics: 333‑344. Museum A. Koenig, Bonn.   PDF:

Wüster, W. & R.S. Thorpe (1989) Population affinities of the Asiatic cobra (Naja naja) species complex in south‑east Asia: reliability and random resampling. Biological Journal of the Linnean Society, 36: 391‑409.    PDF:

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