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Venoms are complex chemical arsenals that have evolved independently many times in the animal kingdom. Venoms have attracted the interest of researchers because they are an important innovation that has contributed greatly to the evolutionary success of many animals, and their medical relevance offers significant potential for drug discovery. During the last decade, venom research has been revolutionized by the application of systems biology, giving rise to a novel field known as venomics. More recently, biotechnology has also made an increasing impact in this field. Its methods provide the means to disentangle and study venom systems across all levels of biological organization and, given their tremendous impact on the life sciences, these pivotal tools greatly facilitate the coherent understanding of venom system organization, development, biochemistry, and therapeutic activity. Even so, we lack a comprehensive overview of major advances achieved by applying biotechnology to venom systems. This review therefore considers the methods, insights, and potential future developments of biotechnological applications in the field of venom research. We follow the levels of biological organization and structure, starting with the methods used to study the genomic blueprint and genetic machinery of venoms, followed gene products and their functional phenotypes. We argue that biotechnology can answer some of the most urgent questions in venom research, particularly when multiple approaches are combined together, and with other venomics technologies.

Additional file 10. zoomable CRISP tree.pdf.

Additional file 16. zoomable kunitz tree.pdf.

Additional file 25. zoomable SVMP propeptide tree.pdf.

Additional file 19. zoomable lectin tree.pdf.

Additional file 22. zoomalbe PIII SVMP tree.pdf.

Additional file 7. Zoomable CNP tree.pdf.

Additional file 4. Tables: Toxin types recovered for each species; Raw illumina data for each species; Statistics for each species for cleaned data; Molecular modelling templates.

Additional file 3. Zoomable 3ftx tree.pdf.

Snakes of the genera and (Viperidae: Viperinae) are known as the desert vipers due to their association with the arid environments of the Middle East. These species have received limited research attention and little is known about their venom or ecology. In this study, a comprehensive analysis of desert viper venoms was conducted by visualising the venom proteomes via gel electrophoresis and assessing the crude venoms for their cytotoxic, haemotoxic, and neurotoxic properties. Plasmas sourced from human, toad, and chicken were used as models to assess possible prey-linked venom activity. The venoms demonstrated substantial divergence in composition and bioactivity across all experiments. venom activated human coagulation factors X and prothrombin and demonstrated potent procoagulant activity in human, toad, and chicken plasmas, in stark contrast to the potent neurotoxic venom of . The venom of also induced coagulation, though this did not appear to be via the activation of factor X or prothrombin. The coagulant properties of and venoms varied among plasmas, demonstrating strong anticoagulant activity in the amphibian and human plasmas but no significant effect in that of bird. This is conjectured to reflect prey-specific toxin activity, though further ecological studies are required to confirm any dietary associations. This study reinforces the notion that phylogenetic relatedness of snakes cannot readily predict venom protein composition or function. The significant venom variation between these species raises serious concerns regarding antivenom paraspecificity. Future assessment of antivenom is crucial.

Vermicella parscauda sp. nov. Holotype. QM J95678, large male collected from boat ramp Weipa, Cape York, Queensland 12��31���53��� S 141��50���51������E in August 2014 by B.G. Fry and F.J. Vonk. Fixed in 10% formalin, stored in 70% ethanol at QM. Paratypes. QM J95679, small male squashed during mining company workplace activities, Evans Landing, Weipa, Qld (1240���21���S, 14152���09���E); AM R94414, unsexed individual, Mission River, Weipa, Qld (1240���59���S, 14152���59���E); CSIRO R02719, adult female, Rocky Point, Weipa, Qld (1237���00���S, 14152���00���E). Diagnosis. A small Vermicella to total 388 mm in length. Distinguished from all other Australian Vermicella by a combination of 55���92 black bands, white bands mottled posterior of scale, SVL 267���357 mm, tail length 21��� 31 mm, 213���230 ventral scales, head length 5���8 mm, head width 3���5 mm and present internasal scales (Figure 1). Ventrals dark black or mottled with distinct black rings on the posterior body and tail, each ring covering 2���3 divided caudal scales and 1���2 white divided caudal scales in between the black rings. Description of holotype (QM J95678). Measurements and counts: ToL 388mm; SVL 357; TaiL 31 mm (8.68% of SVL); HeadL 8mm; HeadW 4mm; HeadH 4mm; SnoutL 4mm; MouthL 6mm; ED 0.9mm; NeckH 4.5mm; NeckW 3mm; MBW 5mm, MBH 5.5mm, VS 230; VSW 4mm. Internasals present with undivided nasal scales, 6 supralabials, 6 infralabials, 0 anterior temporals, 3 posterior temporal scales. Fifteen dorsal scale rows present along the body, 27 total divided subcaudal scales and a ventral scale width of 4mm. Colouration: Neck scales: 6 black scales, 2 white scales; mid body scales: 5 black scales, 1 white scale; vent scales 5 black, 1 white scale. Ventral view mottled black for 2/3rds of body, ending with 17 black rings; 8 on the body and 9 on the tail. Coloration. Colour and pattern of dorsal and lateral surfaces of live specimen alternate black and white bands. White banding begins on the head including white markings classed as bands on the nasal scales and parietal scales. White bands are 1���2 white scales wide with mottling around each white scale resulting in white bands of smaller width that do not have a defined edge. Dark or mottled ventrals that terminate to black rings. Frozen and preserved specimens lose the white vibrancy, resulting in a yellowish tinge and brownish bands under light. Variation. SVL up to 357 mm; TailL 6���8% of SVL, mean 7% (N=4); 213���230 Ventral scales (N=4), Ventrals dark or mottled with 3���10 distinct posterior black rings terminating to 6���7 rings on the tail (N=4). The ring count on the underside of the body was not identified different due to sex of the individual as the single female count was within the range of the males. Distribution and habitat. V. parscauda may to restricted to the Western side of Cape York as current individuals were all found between the Weipa (museum specimens AM R94413, CSIRO R02719, QM J95678 and QM J95679) and Mapoon areas (photographed individuals). The surrounding vegetation is woodlands dominated by Eucalyptus tetrodonta and Eucalyptus miniata, with the soil classed as deep or gradational soils of red massive earth with concretions (Fox et al. 2001). Comparison with other species. Vermicella parscauda sp. nov. is considered geographically distant from other Vermicella species and show the following morphological differences: V. annulata (Grey, 1841) has the largest distribution through NT, QLD, NSW, VIC and SA, including Cape York), white bands can be larger (1���4 scales vs. 1���2) with a distinct edge and often a larger SVL (155��� 760mm vs. 267���357mm). V. intermedia (Keogh & Smith, 1996), from northern WA to northern NT, has fewer black bands (37���65 vs. 55���92) and no internasal scales. V. multifasciata (Longman, 1915), from north-eastern WA to northern NT, including Tiwi Islands, has no internasal scales and more ventral scales (231���350 vs. 213���230). V. snelli (Storr, 1968) from Pilbara region of WA, with a smaller tail length (9���23mm vs. 21���31mm) and normally higher ventral scale count (173���315 vs. 213���230). V. vermiformis (Keogh & Smith, 1996) from a localized population in southern Arnhem Land in NT and central Australia, lower number of black rings (38���51 vs. 55���92), with more ventral scales (253���303 vs. 213��� 230) and ring patterning on ventral scales. Etymology. The specific epithet is modified from the Latin words pars (part) and cauda (tail) in reference to the tail length and formed bands on the tail. Conservation Status. Only 6 individuals of Vermicella parscauda sp. nov. have been recorded and the presumed habitat on Cape York is the site of broad-scale mining. Activities that disturb the soil, such as commercial digging for mining purposes, may adversely affect V. parscauda, given the burrowing behavior of the Vermicella genus. A detailed assessment on abundance, distribution and potential threats to the species will clarify the need for listing by the International Union for Conservation of Nature (IUCN) as a species requiring protection under legislation., Published as part of Derez, Chantelle M., Arbuckle, Kevin, Ruan, Zhiqiang, Xie, Bing, Huang, Yu, Dibben, Lauren, Shi, Qiong, Vonk, Freek J. & Fry, Bryan G., 2018, A new species of bandy-bandy (Vermicella: Serpentes: Elapidae) from the Weipa region, Cape York, Australia, pp. 1-12 in Zootaxa 4446 (1) on pages 10-11, DOI: 10.11646/zootaxa.4446.1.1, http://zenodo.org/record/1437318, {"references":["Fox, I. D., Neldner, V. J., wilson, G. w. & Bannink, P. J. (2001) The Uegetation of the Australian Tropical Savannas. Environmental Protection Agency, Brisbane, pp. 12 - 209.","Keogh, J. S. & Smith, S. A. (1996) Taxonomy and natural history of the Australian bandy-bandy snakes (Elapidae: Uermicella) with a description of two new species. Journal of Zoology, London, 240, 677 - 701. https: // doi. org / 10.1111 / j. 1469 - 7998.1996. tb 05315. x"]}

Vermicella parscauda sp. nov. Holotype. QM J95678, large male collected from boat ramp Weipa, Cape York, Queensland 12°31’53” S 141°50’51’’E in August 2014 by B.G. Fry and F.J. Vonk. Fixed in 10% formalin, stored in 70% ethanol at QM. Paratypes. QM J95679, small male squashed during mining company workplace activities, Evans Landing, Weipa, Qld (1240’21”S, 14152’09”E); AM R94414, unsexed individual, Mission River, Weipa, Qld (1240’59”S, 14152’59”E); CSIRO R02719, adult female, Rocky Point, Weipa, Qld (1237’00”S, 14152’00”E). Diagnosis. A small Vermicella to total 388 mm in length. Distinguished from all other Australian Vermicella by a combination of 55–92 black bands, white bands mottled posterior of scale, SVL 267–357 mm, tail length 21– 31 mm, 213–230 ventral scales, head length 5–8 mm, head width 3–5 mm and present internasal scales (Figure 1). Ventrals dark black or mottled with distinct black rings on the posterior body and tail, each ring covering 2–3 divided caudal scales and 1–2 white divided caudal scales in between the black rings. Description of holotype (QM J95678). Measurements and counts: ToL 388mm; SVL 357; TaiL 31 mm (8.68% of SVL); HeadL 8mm; HeadW 4mm; HeadH 4mm; SnoutL 4mm; MouthL 6mm; ED 0.9mm; NeckH 4.5mm; NeckW 3mm; MBW 5mm, MBH 5.5mm, VS 230; VSW 4mm. Internasals present with undivided nasal scales, 6 supralabials, 6 infralabials, 0 anterior temporals, 3 posterior temporal scales. Fifteen dorsal scale rows present along the body, 27 total divided subcaudal scales and a ventral scale width of 4mm. Colouration: Neck scales: 6 black scales, 2 white scales; mid body scales: 5 black scales, 1 white scale; vent scales 5 black, 1 white scale. Ventral view mottled black for 2/3rds of body, ending with 17 black rings; 8 on the body and 9 on the tail. Coloration. Colour and pattern of dorsal and lateral surfaces of live specimen alternate black and white bands. White banding begins on the head including white markings classed as bands on the nasal scales and parietal scales. White bands are 1–2 white scales wide with mottling around each white scale resulting in white bands of smaller width that do not have a defined edge. Dark or mottled ventrals that terminate to black rings. Frozen and preserved specimens lose the white vibrancy, resulting in a yellowish tinge and brownish bands under light. Variation. SVL up to 357 mm; TailL 6–8% of SVL, mean 7% (N=4); 213–230 Ventral scales (N=4), Ventrals dark or mottled with 3–10 distinct posterior black rings terminating to 6–7 rings on the tail (N=4). The ring count on the underside of the body was not identified different due to sex of the individual as the single female count was within the range of the males. Distribution and habitat. V. parscauda may to restricted to the Western side of Cape York as current individuals were all found between the Weipa (museum specimens AM R94413, CSIRO R02719, QM J95678 and QM J95679) and Mapoon areas (photographed individuals). The surrounding vegetation is woodlands dominated by Eucalyptus tetrodonta and Eucalyptus miniata, with the soil classed as deep or gradational soils of red massive earth with concretions (Fox et al. 2001). Comparison with other species. Vermicella parscauda sp. nov. is considered geographically distant from other Vermicella species and show the following morphological differences: V. annulata (Grey, 1841) has the largest distribution through NT, QLD, NSW, VIC and SA, including Cape York), white bands can be larger (1–4 scales vs. 1–2) with a distinct edge and often a larger SVL (155– 760mm vs. 267–357mm). V. intermedia (Keogh & Smith, 1996), from northern WA to northern NT, has fewer black bands (37–65 vs. 55–92) and no internasal scales. V. multifasciata (Longman, 1915), from north-eastern WA to northern NT, including Tiwi Islands, has no internasal scales and more ventral scales (231–350 vs. 213–230). V. snelli (Storr, 1968) from Pilbara region of WA, with a smaller tail length (9–23mm vs. 21–31mm) and normally higher ventral scale count (173–315 vs. 213–230). V. vermiformis (Keogh & Smith, 1996) from a localized population in southern Arnhem Land in NT and central Australia, lower number of black rings (38–51 vs. 55–92), with more ventral scales (253–303 vs. 213– 230) and ring patterning on ventral scales. Etymology. The specific epithet is modified from the Latin words pars (part) and cauda (tail) in reference to the tail length and formed bands on the tail. Conservation Status. Only 6 individuals of Vermicella parscauda sp. nov. have been recorded and the presumed habitat on Cape York is the site of broad-scale mining. Activities that disturb the soil, such as commercial digging for mining purposes, may adversely affect V. parscauda, given the burrowing behavior of the Vermicella genus. A detailed assessment on abundance, distribution and potential threats to the species will clarify the need for listing by the International Union for Conservation of Nature (IUCN) as a species requiring protection under legislation.

Saw-scaled vipers (genus Echis) are one of the leading causes of snakebite morbidity and mortality in parts of Sub-Saharan Africa, the Middle East, and vast regions of Asia, constituting a public health burden exceeding that of almost any other snake genus globally. Venom-induced consumption coagulopathy, owing to the action of potent procoagulant toxins, is one of the most relevant clinical manifestations of envenomings by Echis spp. Clinical experience and prior studies examining a limited range of venoms and eected antivenoms have demonstrated for some antivenoms an extreme lack of antivenom cross-reactivity between different species of this genus, sometimes resulting in catastrophic treatment failure. This study undertook the most comprehensive testing of Echis venom effects upon the coagulation of human plasma, and also the broadest examination of antivenom potency and cross-reactivity, to-date. 10 Echis species/populations and four antivenoms (two African, two Asian) were studied. The results indicate that the venoms are, in general, potently procoagulant but that the relative dependence on calcium or phospholipid co-factors is highly variable. Additionally, three out of the four antivenoms tested demonstrated only a very narrow taxonomic range of effectiveness in preventing coagulopathy, with only the SAIMR antivenom displaying significant levels of cross-reactivity. These results were in conflict with previous studies using prolonged preincubation of antivenom with venom to suggest effective cross-reactivity levels for the ICP Echi-Tab antivenom. These findings both inform upon potential clinical effects of envenomation in humans and highlight the extreme limitations of available treatment. It is hoped that this will spur efforts into the development of antivenoms with more comprehensive coverage for bites not only from wild snakes but also from specimens widely kept in zoological collections.

Phospholipase A2 are esterolytic enzymes that catalyze the hydrolysis of 1,2-diacyl-3-sn-glycerophospholipids at the C2 position, resulting in the formation of lysophospholipids and free fatty acid. They constitute one of the largest families of lipid hydrolyzing enzymes. PLA2s are known to be one of the most potent components found in snake venoms, where they exhibit a diverse array of pharmacological activities. UCR::Vicerrectoría de Investigación::Unidades de Investigación::Ciencias de la Salud::Instituto Clodomiro Picado (ICP) UCR::Vicerrectoría de Docencia::Salud::Facultad de Microbiología

Sanders, Kate L., Rasmussen, Arne R., Elmberg, Johan, Guinea, Michael, Blias, Peter, Lee, Michael S. Y., Fry, Bryan G. (2012): Aipysurus mosaicus, a new species of egg-eating sea snake (Elapidae: Hydrophiinae), with a redescription of Aipysurus eydouxii (Gray, 1849). Zootaxa 3431: 1-18, DOI: 10.5281/zenodo.211492

Hydrophis donaldi sp. nov. (Figs. 1 and 2, Table 2) Holotype. QM J 90700 (Figs. 1 A���C and 2 A���B), an adult male from the Gulf of Carpentaria, Weipa, Queensland, Australia (12 �� 35 ' 10.88 " S, 141 �� 57 ' 47.21 " E) by B. G. Fry on 15 th October, 2000. Liver tissue stored in ethanol in the QM (Accession no: 007903). Paratypes (n = 7). SAMA R 65215 (juvenile), SAMA R 65216 adult male and SAMA R 66274 adult male, all collected from the Gulf of Carpentaria, Weipa, Queensland, Australia (12 �� 35 ' 10.88 " S, 141 �� 57 ' 47.21 " E) collected by B. G. Fry between the 15-20 October, 2000. Tail tissue stored in ethanol in the ABTC-SAM. QM J 63802, QM J 63803, QM J 63809 and QM J 63810 (all adults, sex undetermined), all collected from the Gulf of Carpentaria, Weipa, Queensland, Australia (12 �� 45 ' 4.03 " S, 141 �� 53 ' 58.10 " E) by B. G. Fry on 1 st March, 1997. Diagnosis. Hydrophis donaldi sp. nov. is distinguished from all other Hydrophis species except H. coggeri, H. sibauensis and H. torquatus diadema by the following combination of characters: ventrals not divided by a longitudinal furrow, 29���30 costal scale rows around neck, 33���35 costal scales around body, 6���7 maxillary teeth behind fang on each side, 246���288 ventrals (Rasmussen et al. 2001; Smith 1926). The new species differs from H. coggeri by having 47���56 (vs 30���42) bands on the body and tail, strongly spinous (vs feebly carinate) body scales, 246���288 (vs 280���360) ventrals, relatively larger and rounded (vs smaller, elongate) head, and anterior part of the maxilla not arched upwards and the tip of the fang projecting below the level of the maxillary teeth (Fig 1 D) (vs anterior part of the maxilla arched upwards and tip of fang not projecting below the level of the maxillary teeth) (Cogger 2000). The new species differs from H. sibauensis by a higher number of scale rows around the neck 29���30 (vs 25���26 in H. sibauensis) and strongly spinous (vs feebly carinate) body scales (Rasmussen et al. 2001). Hydrophis donaldi sp. nov. differs from H. torquatus diadema by a lower midbody scale count (33���35 vs 35���42 in H. torquatus diadema) and strongly spinous (vs feebly carinate) body scales (Smith 1926). Description of the Holotype. Six maxillary teeth are present behind the poison fang. One pre-ocular and postocular scale on each side. Seven supra-labials on each side. First and second supralabials touch the nasal scale and the 3 rd and 4 th touch the eye. Two anterior and posterior temporals on each side. Seven infralabials on each side. The 1 st, 2 nd and 3 rd infralabials are in contact with the anterior chin-shield and the 4 th contacts the posterior chin-shield. Costal scale rows around neck 29; around midbody 33. Dorsal scales strongly spinous. Ventral scales bear two strong spines. Ventrals 269; subcaudals 48; ventrals 1.5 times as wide as adjacent scales. SVL 756 mm; TAL 105 mm; TH 16.71 mm. The dorsal head scales have a rough and warty appearance. Variation (n = 7). Only differences to the holotype are noted. The number of maxillary teeth behind the poison fang is 6 or 7. Two anterior temporals are present in most of the paratypes, but some specimens have one anterior temporal (Table 2) and SAMA R 66274 has one anterior temporal on the left and two on the right. Costal scale rows around the neck 29���30 and around midbody range 33���35. Ventral scales bear two strong spines in all paratypes except in the juvenile SAMA R 65215. Ventrals range from 246���288; subcaudals + 42���51. SVL in mature individuals 649���777 mm; TL 92-101 mm; TH 13.02���16.69 mm (see Table 2). Gene/Locus Marker type Size Substitution Total polymor- Min. fixed Max. fixed (bp) Model phic sites nucleotide nucleotide differences differences Colour pattern. Before preservation in the holotype, the background colour was yellowish brown with 56 brownish bands from neck to tail. Bands are broader dorsally, narrow laterally and taper without connecting ventrally. They are wider than the paler interspaces. Bands in first third of body are darker in colour than rest of the body. Head is pale brown in colour. In preserved specimens bands are brownish grey with paler interspaces. The single juvenile specimen (SAMA R 65215) has a brighter colouration with a blackish head and bands with turquoise blue interspaces. Number of bands on body and tail in the type series range from 47 (SAMA R 65216) to 64 (holotype). See Fig 1 A and B. Etymology. The species is named to honour Dave Donald, the skipper of the boat who worked tirelessly with us and whose local knowledge facilitated the discovery of this species. We propose the common name ���rough scaled sea snake��� for this species due to its strongly keeled body scales. Distribution. Collection localities for H. donaldi sp. nov. are restricted to coastal regions of Weipa, Queensland, northern Australia. The specimens were collected from shallow (Remarks. Hydrophis donaldi sp. nov. can be assigned to the genus Hydrophis (Smith 1926; McDowell 1972; Cogger 2000) based on the following characters: fewer than 73 scale rows around body, single rostral shield, nasals not separated from internasals, more than four supralabials, ventrals small and not broader anteriorly than posteriorly, mental shield broader than long, shorter head without a bill like snout, shorter gape, ventrals entire, no spines on head shields, preocular scales present, maxillary bone not extending forward beyond the palatine, fang followed by a diastema (see Fig 1 D), ventrals distinct throughout the body and not enlarged compared to the dorsal scale rows, ventral scales not broader than twice the adjacent body scales and more than 24 scales around the thickest part of the body. Genetics. The final concatenated mitochondrial alignment consisted of 2704 bp for 25 individuals. Under AIC GTR+i+g is the best-fit model for each partition. There were 1237 invariable, 571 polymorphic and 362 parsimony informative sites and 599 sites with alignment gaps or missing data. The phased G 1888 nuclear DNA alignment contained 367 invariable, 12 polymorphic, six parsimony informative sites and 49 sites with alignment gaps/missing data. In the phased G 1894 nuclear DNA alignment, 385 sites were invariable, 14 polymorphic, 9 parsimony informative and 23 sites with alignment gaps/missing data. Maximum likelihood and Bayesian analysis of the mitochondrial alignment yielded similar branch lengths and topologies that differed only in the position of Astrotia stokesii (sister to a Hydrophis caerulescens + H. brooki clade under ML; sister to H. elegans under Bayesian analysis). In the results of both analyses, H. donaldi sp. nov. forms a well-supported monophyletic sister lineage to all the other sampled ingroup taxa except Hydrelaps darwiniensis and Parahydrophis mertonii, which formed successive sister lineages to the Hydrophis clade (Fig. 3). Hydrophis donaldi sp. nov. was closest to H. czeblukovi, with a mean pairwise corrected (Hasegawa-Kishino-Yano (HKY) correction method) mitochondrial distance of 4.46 % and 13 fixed substitutions separating these taxa. The greatest distance among ingroup taxa was between H. donaldi sp. nov. and P. mertoni (14.64 %), with 140 fixed nucleotide differences. Pairwise distances between H. donaldi sp. nov. and H. coggeri (the species it most closely resembles morphologically) ranged from 7.1 ���8.0% with 71 fixed substitutions. For each nuclear locus Hydrophis donaldi sp. nov. is represented by a single haplotype that is not shared with any other sampled taxon (parsimony trees are shown in Fig. 4 A and B). For G 1888 there was one fixed nucleotide difference between H. donaldi sp. nov. and both H. coggeri and Lapemis curtus, and 7 fixed nucleotide differences between H. donaldi sp. nov and Parahydrophis mertoni. For G 1894 there was one fixed difference between H. donaldi sp. nov. and Acalyptophis peronii, three between H. donaldi sp. nov. and H. coggeri, and 9 between H. donaldi sp. nov. and P. mertoni. The summary of genetic data from the analysis is shown in Table 3., Published as part of Ukuwela, Kanishka D. B., Sanders, Kate L. & Fry, Bryan G., 2012, Hydrophis donaldi (Elapidae, Hydrophiinae), a highly distinctive new species of sea snake from northern Australia, pp. 45-57 in Zootaxa 3201 on pages 48-52, DOI: 10.5281/zenodo.212606, {"references":["Smith, M. (1926) Monograph of the sea-snakes (Hydrophidae). Taylor and Francis, London, UK, 130 pp.","Cogger, H. G. (2000) Reptiles and Amphibians of Australia. Reed New Holland, Sydney, 808 pp.","McDowell, S. B. (1972) The genera of sea-snakes of the Hydrophis group (Serpentes: Elapidae). Transactions of the Zoological Society of London, 32, 189 - 247"]}

Aipysurus mosaicus sp. nov. Figs. 1���3, 4 a, 5 a���c, 6 a���c Aipysurus eydouxii (Gray, 1849) (part): Smith (1926) (part); MacPherson (1933); Cogger (1975); Limpus (1975); Minton (1975) (part); Reidfield et al. (1978); Cogger et al. (1983) (part); Cogger (1992); Golay et al. (1993) (part); Heatwole & Cogger (1994); Greer (1997); Porter et al. (1997); David & Ineich (1999) (part); Heatwole (1999); Ward (2000); Fry et al. (2001); Li et al. (2005 a, b); Lukoschek & Keogh (2006); Cogger (2007). Tomogaster eydouxii Gray, 1849 (specimens collected from Java and therefore not referable to A. mosaicus): Wells (2007). Holotype. SAMA R 65222. A small adult male collected in the Gulf of Carpentaria near Weipa, Australia, in October 2000 by B.F. Fry. Tail tissue stored in ethanol in the ABTC-SAM. Paratype. SAMA R 23493. An adult female collected in the Gulf of Carpentaria, Australia. Tissue stored frozen in the ABTC-SAM. Referred specimens. An additional 26 specimens (9 adult males, 17 adult females), see Appendix 2 for details. TABLE 2. Mean, range, standard deviation, and t-tests of morphological characters comparing Asian (Aipysurus eydouxii) and Australian (A. mosaicus sp. nov. ) specimens. Australian data and t-tests include counts from holotype (H) and paratype (P) when available. T-test probabilities are two-tailed, with significant contrasts given in bold type. See Methods for explanation of characters. Note that sample sizes for some characters differ from the gross sample size per origin and sex. Males Females Generic assignment. Aipysurus mosaicus has a combination of characters (see type description and variation below) that place it in the genus Aipysurus as defined by Smith (1926: 13): ���Maxillary bone as long as or longer than the ectopterygoid, extending forwards beyond the palatine; poison-fangs followed after an interval by from 5 to 11 teeth. Nostrils superior, nasal shields in contact; head-shields regular or broken up; scales imbricate in regular rows, 17���25; ventrals large, one-third to half the breadth of the body, a median keel usually present at least in the posterior part of the body.��� Diagnosis. Aipysurus mosaicus is distinct from seven of the eight species (except A. eydouxii) of Aipysurus using external characters. Aipysurus apraefrontalis and A. foliosquama have ventrals with a deep median notch on the posterior edge, A. mosaicus has ventrals lacking a median notch. Aipysurus tenuis has more than 170 ventrals, A. mosaicus has 140���154. Aipysurus duboisii, A. fuscus, A. laevis, and A. pooleorum have more than 17 scale rows around midbody and at least some of the head shields are usually broken up into smaller plates, A. mosaicus has 17 scale rows around body and head shields that are normally symmetrical and not broken up. Aipysurus mosaicus is clearly distinct from A. eydouxii in a number of external and internal characters. Aipysurus eydouxii has 29 or more bands on body, A. mosaicus has 22 body bands or fewer. In our sample, the following characters differ significantly between A. eydouxii and A. mosaicus when compared intrasexually: number of body bands, tail bands, ventrals, subcaudals, as well as heart and liver positions (absolute and relative) (Table 2). Compared to A. eydouxii, A. mosaicus has a larger premaxilla, a smaller maxilla, a cresent-shaped (rather than triangular) nasal, an anteriorly concave (rather than straight) frontoparietal suture, and dorsally-expanded caudal neural spines (Fig. 5). Description of holotype. Male. Snout-vent length 46.3 cm, tail 8.6 cm. One pre- and two postoculars on both sides. Six supralabials and six infralabials on both sides. First and second supralabials in contact with nasal. Second and third supralabials in contact with preocular on right side, second, third and fourth on left side. Fourth and fifth supralabial in contact with eye. Two anterior temporals on both sides. Sublinguals are well-developed and in contact with one another. First, second, third and fourth infralabials in contact with anterior pair of sublinguals. Fourth infralabial touching posterior pair of sublinguals. Seventeen scale rows on neck, 17 scale rows on body, 15 scale rows ten ventrals before vent. Ventrals 144, distinct throughout, about three to four times as broad as adjacent scales. Preanals enlarged and divided into paired scales. Subcaudals cannot be counted reliably due to a tail biopsy but probably number between 29 and 32. Head dark brown above and below. Body dark brown above, paler below. Twenty-one irregular brown bands on body and five on tail. Description of paratype (R 23493). Female. Snout-vent length 73.0 cm, tail 11.5 cm. One pre- and two postoculars on both sides. Six supralabials and six infralabials on both sides. First and second supralabials in contact with nasal. Second, third and fourth supralabials in contact with preocular on both sides. Fourth and fifth supralabial in contact with eye. Two anterior temporals on both sides. Sublinguals are well-developed and in contact with one another. First, second and third infralabial in contact with anterior pair of sublinguals. Third and fourth infralabial touching posterior pair of sublinguals. Seventeen scale rows on neck, 17 scale rows on body, 13 scale rows ten ventrals before vent. Ventrals 154, distinct throughout, about three to four times as broad as adjacent scales. Preanals enlarged and split up into two scales. Twenty-nine subcaudals. Head dark brown above and below. Body dark brown above, lighter below. Seventeen white/cream irregular brown bands on body and five on tail. Variation in Aipysurus mosaicus sp. nov. The following description refers to all 28 A. mosaicus specimens examined (10 adult males, 18 adult females), including the holotype and paratype: see Table 2 and Appendix 2. External morphological characters. One pre- and two postoculars on both sides in 25 specimens, 1-2 / 1 - 1 in two specimens, and 1-3 / 1-2 in one specimen. Six/six supralabials in 26 specimens, 7 / 6 in two specimens. First and second supralabials in contact with nasal in 26 specimens, first, second and third in two specimens. Second, third and fourth supralabial in contact with preocular in 22 specimens, second and third in six specimens. Fourth supralabial in contact with eye in 22 specimens, third and fourth in one specimen, fourth and fifth in four specimens, and third, fourth and fifth in one specimen. Two anterior temporals on both sides in 22 specimens, 1 / 1 in one specimen, 1 / 2 in one specimen, 2 / 3 in three specimens and 3 / 3 in one specimen. Six infralabials on both sides in 26 specimens, 7 / 7 in two specimens. Sublinguals well-developed and in contact with one another in all specimens. First, second and third infralabial in contact with anterior pair of sublinguals in 24 specimens, first through fourth in four specimens. Third and fourth infralabials touching posterior pair of sublinguals in 22 specimens, only the fourth in six specimens. Sixteen to seventeen scale rows on neck in males, 16���17 in females, 17 scale rows on body in males, 17 in females. Thirteen to seventeen scale rows ten ventrals before vent in males, 13���17 in females. Ventrals 140���147 in males, 140���154 in females, distinct throughout, about three to four times as broad as adjacent scales. Preanals enlarged and split up into two scales. Subcaudals 28���38 in males, 25���33 in females. Snout-vent length of largest male 68 cm, tail 14 cm; largest female 95 cm, tail 15 cm. Internal morphological characters. Tip of heart extending to ventral scale number 44���51 in males, 45���52 in females. %VS-heart 30.8���35.2% in males, 30.0���35.7% in females. Anterior end of liver situated at ventral scale number 47���53 in males, 47���54 in females. %VS-liver 32.9���36.6% in males, 32.0���36.7% in females. Heart and liver separated by 1���4 ventral scales in males, 1���5 in females. Colouration. Head dark brown above and below. Body usually cream, salmon or yellowish with brown to olive green irregular cross-bands. Bands are often incomplete with some forming staggered half-bands on either side of the body. Number of bands on body 17���22 in males and 15���21 in females (when distinct enough to be counted). Number of bands on tail 5���7 in males and 4���6 in females. Distribution. The new species is found along the coasts of northern Australia and southern New Guinea. We have examined specimens (Appendix 2) belonging to this species from the following localities in northern and north-eastern Australia: Coburg Peninsula (Arafura Sea), the Gulf of Carpentaria (southeast and northeast areas), Thursday Island (Torres Strait), Townsville (northeast Queensland), and Swain Reefs and Shoal Water Bay (southeast Queensland). A single specimen was examined from Kaap Valsch (= Tanjung Vals) on the south coast of West Papua. Limpus (1975) reports this species as breeding as far south as 26 o S in subtropical Queensland. Etymology. Named from the Latin mosaicus in reference to the mosaic-like pattern of dark and light scales arranged in irregularly staggered cross-bands. We suggest the common name mosaic sea snake for this species. Natural history. All collection records suggest that Aipysurus mosaicus (mentioned as A. eydouxii) inhabits offshore waters, estuaries and tidal rivers, where it is associated with soft sand and mud bottom substrates. Most records are from depths between 2m (Heatwole 1975) and 22m (Redfield et al. 1978) and there are no records from waters exceeding 50m. Limpus (1975) examined the guts of> 100 A. mosaicus specimens from southern Queensland; 38 % contained diet items, all of which were clumps of benthic fish eggs. A similarly specialist diet of fish eggs (and occasionally squid and cuttlefish) was reported for A. eydouxii in Malaysia (Voris 1966; Glodek & Voris 1982). In northern Australia, females of A. mosaicus are gravid between May and August but not in October (Fry et al. 2001)., Published as part of Sanders, Kate L., Rasmussen, Arne R., Elmberg, Johan, Guinea, Michael, Blias, Peter, Lee, Michael S. Y. & Fry, Bryan G., 2012, Aipysurus mosaicus, a new species of egg-eating sea snake (Elapidae: Hydrophiinae), with a redescription of Aipysurus eydouxii (Gray, 1849), pp. 1-18 in Zootaxa 3431 on pages 7-12, DOI: 10.5281/zenodo.211492, {"references":["Gray, J. E. (1849) Catalogue of the Specimens of Snakes in the Collection of the British Museum. Printed by order of the Trustees, 125 pp.","Smith, M. A. (1926) Monograph of the sea-snakes (Hydrophiidae). Printed by order of the Trustees of the British museum (Natural History) London, 130 pp.","MacPherson, J. (1933) Freshwater Snakes and Sea Snakes. The Australasian Nurses Journal, 31, 100 - 103.","Cogger, H. G. (1975) Sea Snakes of Australia and New Guinea. In Dunson, W. A. (Ed.) The Biology of Sea Snakes. University Park Press, Baltimore London & Tokyo, 59 - 139.","Limpus, C. J. (1975) In Dunson, W. A. (Ed), Coastal sea snakes of subtropical Queensland waters (230 to 280 south latitude). The Biology of Sea Snakes. University Park Press, London & Tokyo, 174 - 182.","Minton, S. A. (1975) Geographic distribution of sea snakes. In Dunson, W. A. (Ed), The Biology of Sea Snakes. London & Tokyo, Baltimore, 21 - 31.","Cogger, H. G., Cameron, E. E. & Cogger, H. M. (1983) Hydrophiidae. In Zoological catalogue of Australia. Amphibia and Reptilia. Government Publishing Service, Canberra: Australian, 241 - 256.","Cogger, H. G. (1992) Reptiles and Amphibians of Australia. Reed International Books, Chatswood New South Wales and Cornell University Press, New York., 775.","Golay, P., Smith, H. M., Broadley, D. G., Dixon, J. R., McCarthy, C., Rage, J. C., Schatti, B. & Toriba, M. (1993) Endoglyphs and other major venomous snakes of the world. A checklist. Herpetological Data Centre, Azemiops, Geneva, 478 pp.","Heatwole, H. & Cogger, H. G. (1994) Sea snakes of Australia. In: Gopalakrishnakone, P. (Ed), Sea snake toxinology. Singapore University Press, Singapore, 167 - 205.","Greer, A. E. (1997) The biology and evolution of Australian snakes. Surrey Beatty and Sons Sydney New South Wales, Australia, 358 pp.","Porter, R., Irwin, S., Irwin, T. & Rodrigues, K. (1997) Records of marine snake species from The Hey-embley and Mission Rivers far North Queensland. Herpetofauna (Sydney), 27, 2 - 7.","David, P. & Ineich, I. (1999) Les serpents venimeux du monde: systematique et repartition. Dumerilia (Paris), 3, 3 - 499.","Heatwole, H. (1999) Sea Snakes. University of New South Wales Press, Sydney, 148 pp.","Ward, T. M. (2000) Factors affecting the catch rates and relative abundance of sea snakes in the by-catch of trawlers targeting tiger and endeavour prawns on the northern Australian continental shelf. Marine and Freshwater Research, 51, 155 - 164.","Fry, G. C., Milton, D. A. & Wassenberg, T. J. (2001) The reproductive biology and diet of sea snake bycatch of prawn trawling in northern Australia: characteristics important for assessing the impacts on populations. Pacific conservation biology, 7, 55 - 73.","Li, M., Fry, B. G. & Kini, R. M. (2005 a) Eggs-Only Diet: Its Implications for the Toxin Profile Changes and Ecology of the Marbled Sea Snake (Aipysurus eydouxii). Journal of Molecular Evolution, 60, 81 - 89.","Li, M., Fry, B. G. & Kini, R. M. (2005 b) Putting the Brakes on Snake Venom Evolution: The Unique Molecular Evolutionary Patterns of Aipysurus eydouxii (Marbled Sea Snake) Phospholipase A 2 Toxins. Molecular Biology and Evolution, 22, 934 - 941.","Lukoschek, V. & Keogh, J. S. (2006) Molecular phylogeny of sea snakes reveals a rapidly diverged adaptive radiation. Biological Journal of the Linnean Society, 89, 523 - 539.","Cogger, H. (2007) Marine snakes. In Tzioumis, V. & Keable, S. (Eds), Description of key species groups in the east marine region. Australian Museum, 80 - 94.","Wells, R. W. (2007) Some taxonomic and nomenclatural considerations on the class Reptilia in Australia. The sea snakes of Australia. An introduction to the members of the families Hydrophiidae and Laticaudidae in Australia, with a new familial and generic arrangement. Australian Biodiversity Record, 8, 1 - 124.","Heatwole, H. (1975) Sea snakes of the Gulf of Carpentaria. In Dunson, W. A. (Ed), The Biology of Sea Snakes. London & Tokyo, Baltimore, 145 - 149.","Redfield, J. A., Holmes, J. C. & Holms, R. D. (1978) Sea snakes of the eastern Gulf of Carpentaria. Australian Journal of Marine and Freshwater Research, 29, 325 - 334.","Voris, H. K. (1966) Fish Eggs as the Apparent Sole Food Item for a Genus of Sea Snake, Emydocephalus (Krefft). Ecology, 47, 152 - 154.","Glodek, G. S. & Voris, H. K. (1982) Marine Snake Diets: Prey Composition, Diversity and Overlap. Copeia, 1982, 661 - 666."]}

Aipysurus eydouxii (Gray, 1849) Aipysurus eydouxii (Gray, 1849); Boulenger (1896); Flower (1899); Wall (1909); Smith (1926) (part); Smith (1943); Taylor (1965); Lim & Sawai (1975); Lemen & Voris (1981); Kharin (1981); Glodek & Voris (1982); McCarthy & Warrell (1991); Stuebing (1991); Golay et al. (1993) (part); Murphy et al. (1999); David & Ineich (1999) (part); Kharin & Hallermann (2009); Rasmussen et al. (2011) Tomogaster eydouxii Gray, 1849 Thalassophis anguillaeformis Schmidt, 1852 Thalassophis muraenaeformis Schmidt, 1852 Aipysurus anguillaeformis (Schmidt, 1852): Gunther (1864); Krefft (1869) Aipysurus margaritophorus Bleeker, 1858 Variation in Aipysurus eydouxii. This new characterisation was made following removal of A. mosaicus and is based on 37 specimens (19 adult males, 18 adult females) examined for this study including the type (see Table 2 and Appendix 2). External morphological characters. One pre- and two postoculars on both sides in 35 specimens, 1 - 1 / 2 - 2 in two specimens, 1 - 1 / 1-2 in one specimen, 1 - 1 / 1-3 in one specimen. Six/six supralabials in 34 specimens, 7 / 6 in two specimens, 7 / 7 in three specimens. First and second supralabials in contact with nasal in 38 specimens, first, second and third in one specimen. Second, third and fourth supralabial in contact with preocular in 22 specimens, second and third in 14 specimens, third and fourth in three specimens. Fourth supralabial in contact with eye in 32 specimens, third and fourth in six specimens, fourth and fifth in one specimen. Two anterior temporals on both sides in 28 specimens, 1 / 1 in two specimens, 1 / 2 in five specimens, 2 / 1 in three specimens and 3 / 3 in one specimen. Six infralabials on both sides in 30 specimens, 7 / 7 in two specimens, 6 / 7 in three specimens, 5 / 6 in one specimen, 7 / 6 in three specimens. Sublinguals well developed and in contact with one another. First, second and third infralabial in contact with anterior pair of sublinguals in 34 specimens, first through fourth in three specimens, first and second in two specimens. Third and fourth infralabials touching posterior pair of sublinguals in 36 specimens, only fourth in one specimen, second and third in two specimens. Scale rows 15���17 on neck in males, 15���17 in females. Scale rows on body 16���17 in males, 17 in females. Fifteen scale rows ten ventrals before vent in males, 13���15 in females. Ventrals 127���143 in males, 136���149 in females, distinct throughout, about three to four times as broad as adjacent scales. Preanals enlarged and split up into two scales. Subcaudals 27���34 in males, 21���27 in females. Snout-vent length in largest male 49 cm, tail length 9.5 cm; largest female 80 cm, tail 11.5 cm. Internal morphological characters: Tip of heart extending to ventral scale number 47���56 in males, 47���55 in females. %VS-heart 34.5��� 40 % in males, 33.1���39.6% in females. Anterior end of liver situated at ventral scale number 50���58 in males, 49���60 in females. %VS-liver 36.0���42.2% in males, 34.5���41.7% in females. Heart and liver separated by distance between 2���4 ventral scales in males, 1���4 in females. Compared to Aipysurus mosaicus, A. eydouxii has a smaller premaxilla, a larger maxilla, a triangular (rather than crescent-shaped) nasal, a straight (rather than anteriorly concave) frontoparietal suture, and caudal neural spines that lack dorsal expansions (Fig. 5). Colouration: Head brown or pale brown above, same or paler below. Body usually cream, salmon or yellowish with brown to olive green cross-bands. Bands are broadest dorsally, narrowing on the sides and ventral surface. Scales between bands usually with dark margins. Number of bands on body 30���42 on males, 29���45 in females. Number of bands on tail 7���11 in males, 4���10 in females. Distribution. Gulf of Thailand, Cambodia, Viet Nam, West Malaysia, Singapore and Java (Smith, 1926; 1943; David & Ineich, 1999). We also examined two specimens from New Guinea (Port Moresby: BMNH 1954.1.12.95 and ���N. Riv. Bvj Bivukeil���: RMNH 5784), but we did not find specimens from the area between the Sunda Shelf and New Guinea in the major museum collections (Berlin, Chicago, Leiden, London and Paris). Boulenger (1896) lists this species as also occurring in the Philippines but to the best of our knowledge no specimens have been collected to confirm this claim (Taylor, 1922; Herre, 1942; Dunson & Minton, 1978; Alcala, 1986)., Published as part of Sanders, Kate L., Rasmussen, Arne R., Elmberg, Johan, Guinea, Michael, Blias, Peter, Lee, Michael S. Y. & Fry, Bryan G., 2012, Aipysurus mosaicus, a new species of egg-eating sea snake (Elapidae: Hydrophiinae), with a redescription of Aipysurus eydouxii (Gray, 1849), pp. 1-18 in Zootaxa 3431 on pages 12-13, DOI: 10.5281/zenodo.211492, {"references":["Gray, J. E. (1849) Catalogue of the Specimens of Snakes in the Collection of the British Museum. Printed by order of the Trustees, 125 pp.","Boulenger, G. A. (1896) Catalogue of the snakes in the British Museum (Natural History). III Containing the Colubridae (Opistoglyphae and Proteroglyphae) Amblycephalidae and Viperidae. British Museum of Natural History, London, 727 pp.","Flower, S. S. (1899) Notes on a Second Collection of Reptiles made in the Malay Peninsula and Siam, from November 1896 to September 1898, with a List of the Species recorded from those Countries. Proceedings of the Zoological Society of London, 67, 600 - 697.","Wall, F. (1909) A monograph of the sea snakes. Memoirs of the Asiatic Society of Bengal, 2, 169 - 251.","Smith, M. A. (1926) Monograph of the sea-snakes (Hydrophiidae). Printed by order of the Trustees of the British museum (Natural History) London, 130 pp.","Smith, M. A. (1943) The fauna of British India, Ceylon and Burma. Reptilia and Amphibia Taylor & Francis, London, 1 - 583 pp.","Taylor, E. H. (1965) The serpentes of Thailand and adjacent waters. University of Kansas Science Bulletin, 9, 609 - 1096.","Limpus, C. J. (1975) In Dunson, W. A. (Ed), Coastal sea snakes of subtropical Queensland waters (230 to 280 south latitude). The Biology of Sea Snakes. University Park Press, London & Tokyo, 174 - 182.","Lemen, C. A. & Voris, H. K. (1981) A Comparison of Reproductive Strategies Among Marine Snakes. The Journal of Animal Ecology, 50, 89 - 101.","Kharin, V. E. (1981) A review of sea snakes of the genus Aipysurus (Serpentes: Hydrophiidae). Zoologicheskii Zhurnal, 60, 257 - 254.","Glodek, G. S. & Voris, H. K. (1982) Marine Snake Diets: Prey Composition, Diversity and Overlap. Copeia, 1982, 661 - 666.","McCarthy, C. J. & Warrell, D. (1991) A collection of sea snakes from Thailand with new records of Hydrophis belcheri (Gray). Bulletin of Natural History Museum (zoology), London, 57, 161 - 166.","Stuebing, R. B. (1991). A checklist of the snakes of Borneo. Raffles Bulletin of Zoology, 39, 323 - 362.","Golay, P., Smith, H. M., Broadley, D. G., Dixon, J. R., McCarthy, C., Rage, J. C., Schatti, B. & Toriba, M. (1993) Endoglyphs and other major venomous snakes of the world. A checklist. Herpetological Data Centre, Azemiops, Geneva, 478 pp.","Murphy, J. C., Cox, M. J. & Voris, H. K. (1999) A key to the sea snakes in the Gulf of Thailand. Natural History Bulletin of the Siam Society, 47, 95 - 108.","David, P. & Ineich, I. (1999) Les serpents venimeux du monde: systematique et repartition. Dumerilia (Paris), 3, 3 - 499.","Kharin, V. & Hallermann, J. (2009) Annotated catalogue of sea kraits (Laticaudidae) and sea snakes (Hydrophiidae) of the herpetological collection of the Zoological Museum, University of Hamburg. Mitteilungen aus dem Hamburgischen zoologische Museum und Institut , 106, 51 - 67.","Rasmussen, A. R., Murphy, J. C., Ompi, M., Gibbons, J. W. & Uetz, P. (2011) Marine Reptiles. PLos One, 6, 1 - 12.","Schmidt, P. (1852) Beitrage zur ferneren Kenntniss der Meerschlangen. Abhandlungen aus dem Gebiete der Naturwissenschaften, 2, 69 - 86.","Gunther, A. C. L. G. (1864) The reptiles of British India. Ray Society, London, 452 pp.","Krefft, G., Scott, H. & Forde, H. (1869) The Snakes of Australia: An Illustrated and Descriptive Catalogue of All the Known Species. T. Richards, Government Printer, 100 pp.","Bleeker, P. (1858) 17. No title. Natuurwetenschappelijk Tijdschrift Voor Nederlandsch - Indie, 16, 49 - 50.","Taylor, E. H. (1922) The snakes of the Philippine Islands. Bureau of Printing, Manila, 555 pp.","Herre, A. W. C. T. (1942) Notes on Philippine Sea-Snakes. Copeia, 1942, 7 - 9.","Dunson, W. A. & Minton, S. A. (1978) Diversity, Distribution, and Ecology of Philippine Marine Snakes (Reptilia, Serpentes). Journal of Herpetology, 12, 281 - 286.","Alcala, A. C. (1986) Amphibians and Reptiles. Guide to Philippine flora and fauna, 10, 1 - 195."]}

Additional file 1: Fig. S1. The effects of venom from an additional African cobra species (Naja nubiae) against the nAChR orthosteric site mimotopes from seven clades of primates. Fig. S2. The effects of venom from an additional African cobra species (Naja haje) against the nAChR orthosteric site mimotopes from seven clades of primates. Fig. S3. The effects of venom from an additional Asian cobra species (Naja kaouthia) against the nAChR orthosteric site mimotopes from seven clades of primates. Table S1. Orthosteric site sequences for each species and public database accessions codes.

The insights gained into the contentious evolutionary history of the venom system of toxicoferan reptiles (Serpentes [snakes] plus Anguimorpha [anguimorph lizards] and Iguania [iguanian lizards]) in recent years have generated considerable controversy regarding the accepted definition of "venom" and what constitutes a venomous animal. Although this may seem little more than a semantic debate, it is of crucial importance that toxinological researchers apply consistent terminology in order to avoid confusion and to facilitate the efficient formulation of research questions into the evolution, ecology, and medical application of venom and its constituent toxins.

Exendin peptides are not only the most enigmatic of all toxins, but they are also among the most medically useful. Their genetic ancestry remains to be conclusively resolved, but the evidence and the principle of parsimony point toward a single evolutionary origin after duplication of an ancestral multidomain gene from a gene encoding hormones of the VIP/glucagon superfamily.

The predatory ecology of Varanus komodoensis (Komodo Dragon) has been a subject of long-standing interest and considerable conjecture. Here, we investigate the roles and potential interplay between cranial mechanics, toxic bacteria, and venom. Our analyses point to the presence of a sophisticated combined-arsenal killing apparatus. We find that the lightweight skull is relatively poorly adapted to generate high bite forces but better adapted to resist high pulling loads. We reject the popular notion regarding toxic bacteria utilization. Instead, we demonstrate that the effects of deep wounds inflicted are potentiated through venom with toxic activities including anticoagulation and shock induction. Anatomical comparisons of V. komodoensis with V. ( Megalania ) priscus fossils suggest that the closely related extinct giant was the largest venomous animal to have ever lived.

Venoms are complex concoctions consisting of peptides, proteins, salts, and other compounds. As such, they present a unique set of challenges for the venom researcher. This chapter provides a broad overview of the techniques useful in venom research, ranging from gross characterization of the venom's protein content to purification and characterization of individual toxins, bioactivity testing, and evolutionary assessment of toxin-encoding genes. Special attention is paid to unique challenges posed by reptile venom or techniques that have proved to be particularly useful.