Whip it into shape: Revision of the Demansia psammophis (Schlegel, 1837) complex (Squamata: Elapidae), with a description of a new species from central Australia.

The genus Demansia Günther is the most diverse genus of Australian terrestrial elapids. A phylogenetic framework for the familiar but problematic taxa D. psammophis and D. reticulata (Gray) has been long overdue to ascertain interspecific relationships and resolve unclear taxonomic issues. Here we present an integrated molecular phylogenetic and morphological analyses to review species delineation, resulting in confirmation that both D. psammophis and D. reticulata are full species and that some populations referred to D. r. cupreiceps Storr are not distinguishable from more typical D. reticulata. We also find the widespread central Australian population (treated by most authors as part of cupreiceps) to be specifically distinct. We redescribe D. psammophis and D. reticulata to clarify morphological and geographical boundaries and describe D. cyanochasma sp. nov. based on a combination of molecular genetic markers, details of colour and pattern, adult total length and a few morphometric attributes. We also designate a lectotype for D. psammophis from the original syntype series and comment on the necessity for further taxonomic refinement of this distinctive group.

Morphology, Diet, and Reproduction of Coastal Hydrophis Sea Snakes (Elapidae: Hydrophiinae) at Their Northern Distribution Limit.

The Ryukyu Archipelago represents the northern distribution limit for hydrophiine sea snakes, the largest group of marine reptiles. Ryukyuan sea snakes may have developed distinct local adaptations in morphology and ecology, but they have been poorly studied. We examined preserved specimens of 111 Hydrophismelanocephalusand 61 Hydrophis ornatusfrom the Ryukyu Archipelago to obtain data on morphology, diet, and reproduction. Sexual size dimorphism was detected in H. melanocephalus (mean ± standard deviation of adult snout-vent length: SVL, females 1062 ± 141 mm vs. males 959 ± 96 mm) but not in H. ornatus. Female H. melanocephalus had larger head widths and shorter tail lengths relative to SVL compared to males. Relative girth was low in neonates of both species (1.0-1.3), but increased in adults to about 1.7-2.6 in H. melanocephalus and 1.3-1.8 in female H. ornatus. Stomach contents of H. melanocephalus consisted of ophichthid and congrid eels, a sand diver, and gobies, whereas in H. ornatus, gobies and a goat fish were found. Litter size of three reproductive H. melanocephalus ranged from five to seven, and parturition seems to occur from August to October. Litter size of six H. ornatus ranged from two to seven, and was correlated with maternal SVL. Parturition in H. ornatus probably occurs around November. Different selective forces related to locomotion, feeding and predation risk, which influence the pregnant mother and neonates, may have resulted in having few, long but slender offspring that show positive allometric growth in hind-body girth.

Systematics of Calliophis intestinalis with the Resurrection of Calliophis nigrotaeniatus (Elapidae, Serpentes).

The red-bellied form of Calliophis intestinalis (Laurenti, 1768) sensu lato was originally reported from Pahang, west Malaysia. To determine the taxonomic status of this form, we examined the type specimens of Elaps sumatranus Lidth De Jeude, 1890, Calliophis intestinalis everetti (Boulenger, 1896), and Callophis furcatus var. nigrotaeniatus Peters, 1863. The results indicated that the red-bellied form of C. intestinalis should be named as Calliophis nigrotaeniatus comb. nov., whose valid species status was based on morphological and molecular analyses. We designate a lectotype and redescribe the species, which is genetically close to Calliophis bilineatus (Peters, 1881) from the Philippines, and is clearly distinguishable from other congeners by possessing a pair of gray or dark blue lateral stripes and by being bright red on the ventrum. Elaps sumatranus and C. i. everetti are relegated to subjective junior synonyms of C. nigrotaeniatus.

A new species of the genus Sinomicrurus (Serpentes: Elapidae) from China and Vietnam.

A new species of Sinomicrurus Slowinski, Boundy, and Lawson, 2001 is described herein based on a series of specimens. The new species, Sinomicrurus peinani sp. nov., occurs in southern China and northern Vietnam. Sinomicrurus peinani sp. nov. is distinguished from its congeners by the following combination of characters: (1) 30-35 black cross-bands on body and tail; (2) 13 dorsal scale rows throughout, all smooth; (3) white belly with black cross-bands or irregular spots; (4) broad white transverse bar on top of head with inverted V-shaped anterior margin, white bar wider than anterior black bar; and (5) frontal V-like, 1.3 times as long as wide. In addition, new occurrences of S. houi in Guangxi, China, and Vietnam are discussed.

Burrowing with a kinetic snout in a snake (Elapidae: Aspidelaps scutatus).

Of the few elongate, fossorial vertebrates that have been examined for their burrowing mechanics, all were found to use an akinetic, reinforced skull to push into the soil, powered mostly by trunk muscles. Reinforced skulls were considered essential for head-first burrowing. In contrast, I found that the skull of the fossorial shield-nosed cobra (Aspidelaps scutatus) is not reinforced and retains the kinetic potential typical of many non-fossorial snakes. Aspidelaps scutatus burrows using a greatly enlarged rostral scale that is attached to a kinetic snout that is independently mobile with respect to the rest of the skull. Two mechanisms of burrowing are used: (1) anteriorly directed head thrusts from a loosely bent body that is anchored against the walls of the tunnel by friction, and (2) side-to-side shovelling using the head and rostral scale. The premaxilla, to which the rostral scale is attached, lacks any direct muscle attachments. Rostral scale movements are powered by, first, retractions of the palato-pterygoid bar, mediated by a ligament that connects the anterior end of the palatine to the transverse process of the premaxilla and, second, by contraction of a previously undescribed muscle slip of the m. retractor pterygoidei that inserts on the skin at the edge of the rostral scale. In derived snakes, palatomaxillary movements are highly conserved and power prey capture and transport behaviors. Aspidelaps scutatus has co-opted those mechanisms for the unrelated function of burrowing without compromising the original feeding functions, showing the potential for evolution of functional innovations in highly conserved systems.

Morphology of the snake spectacle reflects its evolutionary adaptation and development.

BACKGROUND: Covering the eye of all snakes is a transparent integumental structure known as the spectacle. In order to determine variations in spectacle thickness among species, the spectacles of 217 alcohol-preserved museum specimens of 44 species belonging to 14 different families underwent optical coherence tomography (OCT) to measure spectacular thickness. Multivariable analyses were made to determine whether family, activity period (diurnal/nocturnal) and habitat (arboreal/terrestrial/fossorial/aquatic) influenced spectacle thickness. RESULTS: The thinnest spectacles in absolute terms were found in the Usambara bush viper (Viperidae) with a thickness of 74 ± 9 µm and the absolute thickest spectacle was found in the red-tailed pipe snake (Cylindrophiidae) which had a spectacle thickness of 244 ± 57 µm. Fossorial and aquatic snakes had significantly thicker spectacles than arboreal and terrestrial snakes. When spectacle thickness was correlated to eye size (horizontal spectacle diameter), Gray's earth snake (Uropeltidae) had the lowest ratio (1:7) and the cottonmouth (Viperidae) had the highest ratio (1:65). Multivariable and phylogenetic analyses showed that spectacular thickness could be predicted by taxonomic family and habitat, but not activity period. CONCLUSION: This phylogenetically broad systematic study of the thickness of the snake spectacle showed that spectacular thickness varies greatly across snake species and may reflect evolutionary adaptation and development.

Two snakes from eastern Australia (Serpentes: Elapidae); a revised concept of Antaioserpens warro (De Vis) and a redescription of A. albiceps (Boulenger).

Antaioserpens warro sensu lato is known from two populations, one in north-eastern Queensland (Qld), the other from south central Qld. Morphological and genetic assessments demonstrate that these widely separated populations represent two species. A re-examination of museum specimens and the type descriptions show that the name A. warro (De Vis) has been erroneously applied to the north-eastern Qld species. The type specimen of A. warro, from the Gladstone district in south-east Qld, is badly faded but the colour pattern as described by De Vis (1884a) is consistent with that of recently collected specimens from south central Qld and it is this species to which the name applies. The earliest available name for the species from north-eastern Qld is A. albiceps (Boulenger, 1898). Both A. warro and A. albiceps are redescribed herein.

A new species of monadal coral snake of the genus Micrurus (Serpentes, Elapidae) from western Amazon.

We described a new species of monadal coral snake of the genus Micrurus from the region of Tabatinga and Leticia, along the boundaries of Brazil, Colombia, and Peru. The new species can be distinguished from the other congeners by the combination of the following characters: absence of a pale nuchal collar; black cephalic-cap extending from rostral to firstdorsal scale and enclosing white tipped prefrontal scales; upper half of first to four supralabials and postoculars black; tricolor body coloration, with 27-31 black rings bordered by narrower white rings and 27-31 red rings; tail coloration similar to body, with alternating black rings bordered by irregular narrow white rings, red rings of the same width as the black rings; ventral scales 205-225; subcaudal scales 39-47.

The development of the skull of the Egyptian Cobra Naja h. haje (Squamata: Serpentes: Elapidae).

BACKGROUND: The study of craniofacial development is important in understanding the ontogenetic processes behind morphological diversity. A complete morphological description of the embryonic skull development of the Egyptian cobra, Naja h. haje, is lacking and there has been little comparative discussion of skull development either among elapid snakes or between them and other snakes. METHODOLOGY/PRINCIPAL FINDINGS: We present a description of skull development through a full sequence of developmental stages of the Egyptian cobra, and compare it to other snakes. Associated soft tissues of the head are noted where relevant. The first visible ossification centres are in the supratemporal, prearticular and surangular, with slight ossification visible in parts of the maxilla, prefrontal, and dentary. Epiotic centres of ossification are present in the supraoccipital, and the body of the supraoccipital forms from the tectum posterior not the tectum synoticum. The venom glands are visible as distinct bodies as early at stage 5 and enlarge later to extend from the otic capsule to the maxilla level with the anterior margin of the eye. The gland becomes more prominent shortly before hatching, concomitant with the development of the fangs. The tongue shows incipient forking at stage 5, and becomes fully bifid at stage 6. CONCLUSIONS/SIGNIFICANCE: We present the first detailed staging series of cranial development for the Egyptian cobra, Naja h. haje. This is one of the first studies since the classical works of G. de Beer and W. Parker that provides a detailed description of cranial development in an advanced snake species. It allows us to correct errors and misinterpretations in previous accounts which were based on a small sample of specimens of uncertain age. Our results highlight potentially significant variation in supraoccipital formation among squamates and the need for further research in this area.

Molecules and morphology reveal overlooked populations of two presumed extinct Australian sea snakes (Aipysurus: Hydrophiinae).

The critically endangered leaf-scaled (Aipysurus foliosquamaI) and short-nosed (A. apraefrontalis) sea snakes are currently recognised only from Ashmore and Hibernia reefs ~600km off the northwest Australian coast. Steep population declines in both species were documented over 15 years and neither has been sighted on dedicated surveys of Ashmore and Hibernia since 2001. We examine specimens of these species that were collected from coastal northwest Australian habitats up until 2010 (A.foliosquama) and 2012 (A. apraefrontalis) and were either overlooked or treated as vagrants in conservation assessments. Morphological variation and mitochondrial sequence data confirm the assignment of these coastal specimens to A. foliosquama (Barrow Island, and offshore from Port Hedland) and A.apraefrontalis (Exmouth Gulf, and offshore from Roebourne and Broome). Collection dates, and molecular and morphological variation between coastal and offshore specimens, suggest that the coastal specimens are not vagrants as previously suspected, but instead represent separate breeding populations. The newly recognised populations present another chance for leaf-scaled and short-nosed sea snakes, but coastal habitats in northwest Australia are widely threatened by infrastructure developments and sea snakes are presently omitted from environmental impact assessments for industry. Further studies are urgently needed to assess these species' remaining distributions, population structure, and extent of occurrence in protected areas.

Sea snakes in Australian waters (Serpentes: subfamilies Hydrophiinae and Laticaudinae)--a review with an updated identification key.

Sea snakes (Elapidae, subfamilies Hydrophiinae and Laticaudinae) reach high species richness in the South China Sea and in the Australian region; however, most countries in the two regions still lack up-to-date checklists and identification tools for these snakes. We present an updated reviewed checklist and a new complete identification key to sea snakes in Australian waters. The identification key includes 29 species documented and 4 possibly occurring taxa and is based mostly on easy-to-use external characters. We find no evidence for breeding populations of Laticauda in Australian waters, but include the genus on the list of possibly occurring taxa. 

Rapid evolution of mimicry following local model extinction.

Batesian mimicry evolves when individuals of a palatable species gain the selective advantage of reduced predation because they resemble a toxic species that predators avoid. Here, we evaluated whether-and in which direction-Batesian mimicry has evolved in a natural population of mimics following extirpation of their model. We specifically asked whether the precision of coral snake mimicry has evolved among kingsnakes from a region where coral snakes recently (1960) went locally extinct. We found that these kingsnakes have evolved more precise mimicry; by contrast, no such change occurred in a sympatric non-mimetic species or in conspecifics from a region where coral snakes remain abundant. Presumably, more precise mimicry has continued to evolve after model extirpation, because relatively few predator generations have passed, and the fitness costs incurred by predators that mistook a deadly coral snake for a kingsnake were historically much greater than those incurred by predators that mistook a kingsnake for a coral snake. Indeed, these results are consistent with prior theoretical and empirical studies, which revealed that only the most precise mimics are favoured as their model becomes increasingly rare. Thus, highly noxious models can generate an 'evolutionary momentum' that drives the further evolution of more precise mimicry-even after models go extinct.

A new species of triadal coral snake of the genus Micrurus Wagler, 1824 (Serpentes: Elapidae) from northeastern Brazil.

The genus Micrurus comprises 123 currently recognized taxa (species and subspecies) that are traditionally arranged in four species groups diagnosable mainly by color pattern characteristics. Here, we describe a new species of triadal coral snake from northeastern Brazil. The new species is distinguished from other sympatric triadal congeners (M. lemniscatus carvalhoi, M. ibiboboca and M. brasiliensis) mainly by the entirely black parietals and by a suite of external characters and hemipenial morphology. The new species appears to be restricted to tropical ombrophilous lowland coastal forests of northeastern Brazil and all recently collected specimens are known to occur in small forest patches surrounded by periurban environment, which calls for an urgent evaluation on its conservation status.

Trophic divergence despite morphological convergence in a continental radiation of snakes.

Ecological and phenotypic convergence is a potential outcome of adaptive radiation in response to ecological opportunity. However, a number of factors may limit convergence during evolutionary radiations, including interregional differences in biogeographic history and clade-specific constraints on form and function. Here, we demonstrate that a single clade of terrestrial snakes from Australia--the oxyuranine elapids--exhibits widespread morphological convergence with a phylogenetically diverse and distantly related assemblage of snakes from North America. Australian elapids have evolved nearly the full spectrum of phenotypic modalities that occurs among North American snakes. Much of the convergence appears to involve the recurrent evolution of stereotyped morphologies associated with foraging mode, locomotion and habitat use. By contrast, analysis of snake diets indicates striking divergence in feeding ecology between these faunas, partially reflecting regional differences in ecological allometry between Australia and North America. Widespread phenotypic convergence with the North American snake fauna coupled with divergence in feeding ecology are clear examples of how independent continental radiations may converge along some ecological axes yet differ profoundly along others.

Morphology, reproduction and diet in Australian and Papuan death adders (Acanthophis, Elapidae).

Death adders (genus Acanthophis) differ from most other elapid snakes, and resemble many viperid snakes, in their thickset morphology and ambush foraging mode. Although these snakes are widely distributed through Australia and Papua New Guinea, their basic biology remains poorly known. We report morphological and ecological data based upon dissection of >750 museum specimens drawn from most of the range of the genus. Female death adders grow larger than conspecific males, to about the same extent in all taxa (20% in mean adult snout-vent length,  =  SVL). Most museum specimens were adult rather than juvenile animals, and adult males outnumbered females in all taxa except A. pyrrhus. Females have shorter tails (relative to SVL) than males, and longer narrower heads (relative to head length) in some but not all species. The southern A. antarcticus is wider-bodied (relative to SVL) than the other Australian species. Fecundity of these viviparous snakes was similar among taxa (mean litter sizes 8 to 14). Death adders encompass a broad range of ecological attributes, taking a wide variety of vertebrate prey, mostly lizards (55%), frogs and mammals (each 21%; based on 217 records). Dietary composition differed among species (e.g. frogs were more common in tropical than temperate-zone species), and shifted with snake body size (endotherms were taken by larger snakes) and sex (male death adders took more lizards than did females). Overall, death adders take a broader array of prey types, including active fast-moving taxa such as endotherms and large diurnal skinks, than do most other Australian elapids of similar body sizes. Ambush foraging is the key to capturing such elusive prey.

Coastal nurseries and their importance for conservation of sea kraits.

Destruction and pollution of coral reefs threaten these marine biodiversity hot stops which shelter more than two thirds of sea snake species. Notably, in many coral reef ecosystems of the Western Pacific Ocean, large populations of sea kraits (amphibious sea snakes) have drastically declined during the past three decades. Protecting remaining healthy populations is thus essential. In New Caledonia, coral reefs shelter numerous sea krait colonies spread throughout an immense lagoon (24,000 km2). Sea kraits feed on coral fish but lay their eggs on land. However, ecological information on reproduction and juveniles is extremely fragmentary, precluding protection of key habitats for reproduction. Our 10 years mark recapture study on Yellow sea kraits (L. saintgironsi >8,700 individuals marked) revealed that most neonates aggregate in highly localized coastal sites, where they feed and grow during several months before dispersal. Hundreds of females emigrate seasonally from remote populations (>50 km away) to lay their eggs in these coastal nurseries, and then return home. Protecting these nurseries is a priority to maintain recruitment rate, and to retain sea krait populations in the future.

3D flow in the venom channel of a spitting cobra: do the ridges in the fangs act as fluid guide vanes?

The spitting cobra Naja pallida can eject its venom towards an offender from a distance of up to two meters. The aim of this study was to understand the mechanisms responsible for the relatively large distance covered by the venom jet although the venom channel is only of micro-scale. Therefore, we analysed factors that influence secondary flow and pressure drop in the venom channel, which include the physical-chemical properties of venom liquid and the morphology of the venom channel. The cobra venom showed shear-reducing properties and the venom channel had paired ridges that span from the last third of the channel to its distal end, terminating laterally and in close proximity to the discharge orifice. To analyze the functional significance of these ridges we generated a numerical and an experimental model of the venom channel. Computational fluid dynamics (CFD) and Particle-Image Velocimetry (PIV) revealed that the paired interior ridges shape the flow structure upstream of the sharp 90° bend at the distal end. The occurrence of secondary flow structures resembling Dean-type vortical structures in the venom channel can be observed, which induce additional pressure loss. Comparing a venom channel featuring ridges with an identical channel featuring no ridges, one can observe a reduction of pressure loss of about 30%. Therefore it is concluded that the function of the ridges is similar to guide vanes used by engineers to reduce pressure loss in curved flow channels.

Recent rapid speciation and ecomorph divergence in Indo-Australian sea snakes.

The viviparous sea snakes (Hydrophiinae) are a young radiation of at least 62 species that display spectacular morphological diversity and high levels of local sympatry. To shed light on the mechanisms underlying sea snake diversification, we investigated recent speciation and eco-morphological differentiation in a clade of four nominal species with overlapping ranges in Southeast Asia and Australia. Analyses of morphology and stomach contents identified the presence of two distinct ecomorphs: a 'macrocephalic' ecomorph that reaches >2 m in length, has a large head and feeds on crevice-dwelling eels and gobies; and a 'microcephalic' ecomorph that rarely exceeds 1 m in length, has a small head and narrow fore-body and hunts snake eels in burrows. Mitochondrial sequences show a lack of reciprocal monophyly between ecomorphs and among putative species. However, individual assignment based on newly developed microsatellites separated co-distributed specimens into four significantly differentiated clusters corresponding to morphological species designations, indicating limited recent gene flow and progress towards speciation. A coalescent species tree (based on mitochondrial and nuclear sequences) and isolation-migration model (mitochondrial and microsatellite markers) suggest between one and three transitions between ecomorphs within the last approximately 1.2 million to approximately 840,000 years. In particular, the macrocephalic 'eastern' population of Hydrophis cyanocinctus and microcephalic H. melanocephalus appear to have diverged very recently and rapidly, resulting in major phenotypic differences and restriction of gene flow in sympatry. These results highlight the viviparous sea snakes as a promising system for speciation studies in the marine environment.

Molecular evidence that the deadliest sea snake Enhydrina schistosa (Elapidae: Hydrophiinae) consists of two convergent species.

We present a striking case of phenotypic convergence within the speciose and taxonomically unstable Hydrophis group of viviparous sea snakes. Enhydrina schistosa, the 'beaked sea snake', is abundant in coastal and inshore habitats throughout the Asian and Australian regions, where it is responsible for the large majority of recorded deaths and injuries from sea snake bites. Analyses of five independent mitochondrial and nuclear loci for populations spanning Australia, Indonesia and Sri Lanka indicate that this 'species' actually consists of two distinct lineages in Asia and Australia that are not closest relatives. As a result, Australian "E. schistosa" are elevated to species status and provisionally referred to Enhydrinazweifeli. Convergence in the characteristic 'beaked' morphology of these species is probably associated with the wide gape required to accommodate their spiny prey. Our findings have important implications for snake bite management in light of the medical importance of beaked sea snakes and the fact that the only sea snake anti-venom available is raised against Malaysian E. schistosa.