AUSTRALIAN BIODIVERSITY RECORD - Calodema

AUSTRALIAN BIODIVERSITY RECORD______________________________________________________________2007 (No 8) ISSN 1325-2992 May, 2007______________________________________________________________Some Taxonomic and Nomenclatural Considerations on the Class Reptilia in Australia.The Sea Snakes of Australia. An Introduction to the Members of the FamiliesHydrophiidae and Laticaudidae in Australia, with a New Familial and GenericArrangement.byRichard W. WellsP.O. Box 826, Lismore, New South WalesAustralia, 2480IntroductionIn this revision of the Australian sea snakes it will be readily apparent that I have departedconsiderably from the classifications adopted in the most recent comprehensive regionaltreatments of this group (see Cogger, Cameron and Cogger (1983), Cogger (2000), andWilson and Swan (2003). The major areas where change has occurred relates to the genericand familial assignment of the 31-32 species of the Hydrophiidae and 2 species of theLaticaudidae so far known from Australia. No new species of sea snakes are describedherein, but I have continued to treat Aipysurus pooleorum as a full species as was done byWells and Wellington (1984, 1985), Storr, Smith and Johnstone (1986, 2002), Ehmann (1992)and most recently by Wilson and Swan (2003). I have also substantially reclassified thetraditional genera Hydrophis and Aipysurus, a number of genera have been resurrected fromsynonymy, and a new genus is erected within the Aipysurus group. The familial classificationof the Hydrophiidae is revised.I have continued to accept the widely supported view that the genera Acalyptophis andLapemis are valid. I also accept the generic distinctiveness of Astrotia and Disteira, althoughsome workers continue to believe that both should be submerged into Hydrophis or at aminimum, Astrotia synonymized with Disteira.In regards the genera Aipysurus and Hydrophis I consider that both (in traditional treatments)represent highly artificial assemblages. The pioneering work of Kharin has already begun thedismemberment of Hydrophis with his resurrections of Chitulia, Leioselasma andPolydontognathus - which I agree with, and continue to follow here, but at a higher level. Inmy division of the genus Chitulia I have included ‘Chitulia ornata’ as part of the Australianfauna because of its traditional association, but at the conclusion of that species description Itake the formal step of referring the Australian population of ornata to Chitulia ocellata (Gray,1849). Further, I treat the genus Microcephalophis as valid for what has been previouslyreferred to as Hydrophis gracilis. One result of the above changes is the restriction of thegenus Hydrophis in Australia to a much smaller group of species - Hydrophis atriceps,Hydrophis elegans, Hydrophis mcdowelli, Hydrophis melanosoma, and Hydrophis vorisi.The genus Aipysurus is similarly in need of revision. It has been recently treated as having ahigh degree of similarity between its included species, however I consider that this genus ishighly divergent and herein offer a new arrangement. The genus Aipysurus has until nowincluded a number of species assemblages that have broadened the morphological andecological basis for that genus almost to the absurdity of the situation in Hydrophis (sensulato). Indeed, it has even been suggested that Emydocephalus should be synonymized withAipysurus due to presumptions of morphological similarity between the two genera. In my

Australian Biodiversity Record, 2007 (8): 1-124view some support could be had for believing that Emydocephalus is indeed distantly relatedto the traditional assemblages represented by Aipysurus yet Emydocephalus does clearlyrepresent a distinct genus as most authorities continue to maintain. This has phylogeneticconsequences for the interpretation of the genus Aipysurus, for if one accepts that themorphological divergence of Emydocephalus is distinctive enough to warrant separategeneric recognition while recognizing its lineage to Aipysurus, then those more significantlydiverged assemblages within Aipysurus must require similar recognition. Kharin (1981) hasobviously recognized this with his description of Smithohydrophis for apraefrontalis - which Ialso accept, but at the generic level. Therefore, I have concluded that Aipysurus should besplit into six separate genera within the Australian region: viz Aipysurus for laevis andpooleorum; Pelagophis for duboisii; Stephanohydra for fusca; Tomogaster for eydouxii, andSmithohydrophis for apraefrontalis, and foliosquama and a new genus for tenuis.Despite the above generic rearrangements, the higher level classification of the sea snakeshas until recent years remained fairly stable with all the sea snakes being either placed in asingle Family (Hydrophiidae), or with perhaps broader appeal, into two Families(Hydrophiidae and Laticaudidae). However, a flurry of recent research, initially pioneered bySamuel McDowell, has focussed on the evolutionary relationships of sea snakes to theterrestrial proteroglyphs. A widening view is emerging that the sea snakes are not distinct atthe family level, and so should be placed within the ‘Elapids’, but this is a view that I cannotsupport.I have rejected the recent merging of both these families within the Family Elapidae -particularly in light of the polyphyletic nature of that Family. The Elapidae sensu stricto should,in my opinion, be restricted to only those proteroglyphic snakes of the Americas (Micruridaeand related assemblages), and the rest of the so-called Elapidae (including those ofAustralasia) consequently relegated to entirely separate Families. In the case of Australasia -the snakes currently assigned to the Elapidae should be placed within the FamilyOxyuranidae.While the sea snakes may have had part of their origins in the distant evolution of theAustralasian Oxyuranid radiation, the sea snakes can be separated from all otherproteroglyphs by a range of morphological characters, including but not limited to, thepossession of valvular nostrils and lingual fossa, the absence of choanal processes on thepalatine bones, and a unique physiology that has evolved a range of critical adaptations to atotally aquatic life-style in a high-pressure, saline environment (see numerous includedreferences). For instance, their circulatory and central nervous systems are so delicatelyadapted to the higher pressures experienced in water that exposure to normal air pressure foreven a brief period can prove fatal to most species as they have very limited control of arterialblood flow when out of water. The vertically compressed body forms of most species,including their unique vertically compressed paddle-like tail, and greatly reduced ventral scalestructure are obvious adaptations for aquatic locomotion which have no parallels in theterrestrial proteroglyphs. The integument is structurally and functionally also very distinct fromthe terrestrial proteroglyphs due not only to the requirements of aquatic locomotion, but alsoto the extreme physiological consequences of swimming in what chemically amounts to a seaof sodium. The highly vascularized skin of sea snakes also allows for cutaneous respiration -obtaining oxygen while underwater - with the excess salt being removed by a speciallyadapted salt gland under the tongue. No other proteroglyphic snake is capable of cutaneousrespiration.So, it is clear to me that the sea snakes, while obviously distantly related genetically, arephylogenetically distinct from the rest of the proteroglypha. The modern view that sea snakesshould be placed within the Elapidae largely on the basis of their origins is at the price ofignoring the significance of some of the most spectacular evolutionary advances made byreptiles in their history. The adaptations of terrestrial snakes to the marine environmentjustifies them being treated as separate Families, and quite bluntly, as a genus isrepresentative of a group of species that are each others closest relatives, then a Family mustbe treated as representative of a group of genera that are each others closest relatives. Ineffect this simply means that the sea snakes actually represent MORE than the two traditionalFamilies. At its most readily appreciated focus, a Family of sea snakes could represent an2

Australian Biodiversity Record, 2007 (8): 1-124implicit statement of linkage to marine adaptation from its terrestrial ancestors. Therecognition of the Family Laticaudidae - a group of seven species in two genera with no closerelatives in any other group of sea snakes - could be viewed this way and probably indicatesthat terrestrial proteroglyphs have experimented with colonizing the marine environment onmore than one phylogenetic front.One of the consequences of accepting the Laticaudidae as a phylogenetically distinct lineagefrom the Hydrophiidae, is that the Hydrophiidae is itself polyphyletic for its traditionallyincluded species in some cases depart significantly in morphology to a similar degree to theLaticaudidae. Therefore, using just the criteria that justify the recognition of Laticaudidae, theAipysurus and Hydrophis lineages should be placed within their own families in my opinion.Further, the Australian mangrove snakes of the genera Ephalophis and Parahydrophis, andthe genus Hydrelaps are so divergent morphologically that they have even been placed byothers in separate subfamilies (the Ephalophinae and the Hydrelapinae). The pan-tropicalPelamis - including Lapemis and related genera - also represent a separate lineage worthy ofhigher recognition. So, rather than follow the trend by degrading the higher classification ofthe sea snakes, I have opted for the opposite, and expanded it with what I believe is a morephylogenetically appropriate arrangement.Thus in this paper I have removed a number of genera from the Hydrophiidae, and placedthem within their own separate Families - viz: the Aipysurus group of genera (Aipysurus,Emydocephalus, Oceanius gen. nov., Pelagophis, Smithohydrophis, Stephanohydra, andTomogaster) are now placed in the Family Aipysuridae. The genus Hydrelaps is now in theFamily Hydrelapidae, and the genera Ephalophis and Parahydrophis are now in the FamilyEphalophiidae. Pelamis, and Lapemis (along with the extralimital Kolpophis and Praescutata)are now referred to the Family Pelamiidae. Finally, my present concept of the FamilyHydrophiidae sensu stricto is that it should only include the following genera in Australia -Acalyptophis, Astrotia, Chitulia, Disteira, Enhydrina, Hydrophis, Leioselasma,Microcephalophis and Polydontognathus. The extralimital genera Mediohydrophis (hereinregarded as generically distinct), Kerilia and Thallasophis are included in the Hydrophiidaebut further remarks on these genera will follow in a subsequent paper on the Asian seasnakes.AcknowledgementsDuring the mid 1960s I obtained many preserved sea snakes from trawler operators in northQueensland, and it was apparent in those days that vast numbers were being accidentallykilled in netting operations. My original intention of identifying all the sea snake species inAustralia was soon abandoned when faced with the multitude of different specimens thatarrived in alcohol-dripping parcels from Cairns. Following the enactment of protectivelegislation for reptiles in Queensland, my collecting had to be abandoned, but the slaughter ofsea snakes of course continued - as it still does to this day. When I later lived in Darwin in the1970s and the 1980s one of my main intentions was to study the snakes of the mangrovesaround the city, and I have many fishermen to thank for the information that they gave me onsea snakes. The late Graeme Gow of the Northern Territory Museum in Darwin, and I alsoexamined many sea snakes from trawling operators in northern Australia during the late1970s and some of best memories of our friendship relate to our attempts at trying to identifyand photograph specimens amid the ruins of the old Museum in Smith Street. Today there isa huge literature on sea snakes thanks largely to the impetus provided by the pioneering workof Harold Cogger and William Dunson in the 1970s (and of course their collaborators).Although a picture has begun to emerge of the diversity and complexity of Australia’s seasnake fauna, there is still much to be done, and so few resources, and so little time to do it.But more importantly, it requires the enthusiasm to actually get out and have a go atsomething worth doing. Recently, a good friend and colleague in herpetology - Alex Dudley -sent me a photo of our days in Darwin looking for sea snakes - virtually up to our knees inblack mud wading through mangrove swamps for sea snakes, now some twenty years ago. Ilooked a lot younger and fitter in the photo - but what the photo didn’t show was that on theday of the photo, we were unemployed and down to our last dollar and had bought achocolate bar between us so that we could have the energy to check out the mangroves andlook for sea snakes. Now that’s enthusiasm. So for those interested - go down the shop, buya Mars Bar and go looking for sea snakes…3

Australian Biodiversity Record, 2007 (8): 1-124The Sea Snakes of AustraliaFamily AipysuridaeGenus Aipysurus Lacepede, 1804Aipysurus Lacépède, B.G.E. (1804). Ann. Mus. Natl Hist. Nat. Paris 4: 184-211 [p. 210]. Typespecies: Aipysurus laevis Lacépède, 1804 by monotypy.Aspisurus Gray, J.E. (1841): In: Grey, G. Journals of Two Expeditions of Discovery in Northwestand Western Australia… Vol. 2 [p. 433] [non Aspisurus Lacépède, 1802; lapsus proAipysurus Lacépède, 1804].Aepyurus Agassiz, L. (1846). Nomenclatoris Zoologici [p. 9] [emend. pro Aipysurus Lacépède,1804].Hypotropis Gray, J.E. (1846): Ann. Mag. Nat. Hist. 18: 284 [284]. Type species: Hypotropisjukesii Gray, 1846 by monotypy.Diagnosis: A genus of very large and robust-bodied marine snakes of the Family Aipysuridae,readily separated from all other genera by the following combination of characters: Headshields large and symmetrical with only minor partial fragmentation posteriorly; prefrontalspresent, and occasionally divided; frontal usually divided; supraoculars divided; parietalsdivided; preoculars present (usually 1-2 (rarely 3); nasal not contacting preocular, separatedby the presence of a loreal; nasals in contact; 2-3 (rarely 1) postoculars; temporals small;supralabials 8-10; posterior chin-shields small and separated by one or more small scales;portion of rostral scale bearing median, valve-like fold sometimes separated from remainderof scale by suture; valvular nostrils and lingual fossa; body scales smooth in females andimbricate and in about 20-25 rows at mid-body; ventrals about 140-160; ventrals large andextending about half way across belly, and each about three times as wide as adjacent bodyscales; ventrals with a median keel and a posterior notch (but smooth in females); analdivided; subcaudals around 20-40 entire; tail strongly compressed vertically, paddle-like, witha unique photoreceptor just anterior to the tip; venom apparatus advanced and highly toxic;ovoviviparous. Etymology: The name Aipysurus means ‘high tail’ (from the Greek, ‘aipys’ -high, and ‘ura’ - tail, and refers to the typical sea snake laterally flattened tail shape. Content:Aipysurus laevis Lacepede, 1804; Aipysurus pooleorum Smith, 1974.Aipysurus laevis Lacepede, 1804Aipsurus laevis Lacépède, B.G.E. (1804). Ann. Mus. Natl Hist. Nat. Paris 4: 184-211 [p. 210pl. 56 fig 3]. Type data: Neotype WAM R22384. Subsequent designation: Smith, L.A. (1974):Rec. West. Aust. Mus. 3: 93-110 [p. 99]. Type locality: near Locker Is., off Onslow, WA[21º44'S 114º46'E].Hypotropis jukesii Gray, J.E. (1846): Ann. Mag. Nat. Hist. 18: 284 [p. 284]. Type data:holotype BMNH 1946.1.9.53. Type locality: nr Darnley Island, Torres Strait (as - nr DarnleyIls, Port Essington), QLD.Aipsurus fuliginosus Duméril, A.M.C., Bibron, G. and Duméril, A. (1854): Erpét. Gén. Hist.Nat. Compl. Reptiles. Roret, Paris [p. 1327 pl. 77 fig. 1]. Type data: holotype MNHP 639.Type locality: New Caledonia.Description: This is one the largest and bulkiest of the sea snakes at its full size, but evenwhen only average-sized, specimens are robust-bodied, with a short and deep head that isindistinct from the neck. Its colouration is highly variable with the base colour ranging frompale yellowish-brown through olive green to dark purple on the upper body. There is bothsexual and ontogenetic colour and pattern variation in this species. At birth, juveniles aremarked with a series of pale and dark bands, but as they mature, the pattern fades to a moreuniform colouration. In some a pattern of scattered darker brown or black body scales amonglarger creamish scales may form a weak lateral blotching or partial reticulum effect along the4

Australian Biodiversity Record, 2007 (8): 1-124body which contrasts strongly with the usually darker upper body, but this also fades with age.Completely patternless specimens of various shades of uniform olive brown on the upperbody occur with increasing age. The venter is usually pale creamish-white. The is markedsexual dimorphism in the topotypic population of this species, with males being a uniformbrown, and females dark grey with a brownish head. Some significant features of this species'morphology are: Head shields large and symmetrical with only minor partial fragmentationposteriorly; prefrontals present, and occasionally divided; frontal usually divided; supraocularsdivided; parietals divided; preoculars present (usually 1-2 (rarely 3); nasal not contactingpreocular, separated by the presence of a loreal; nasals in contact; 2-3 (rarely 1) postoculars;temporals small; supralabials 8-10; posterior chin-shields small and separated by one or moresmall scales; portion of rostral scale bearing median, valve-like fold sometimes separatedfrom remainder of scale by suture; body scales smooth in females and imbricate and in about21-25 rows at mid-body (males and females do not appear to have significantly different midbodyscale counts); ventrals about 140-155 (females do not appear to have a higher ventralcount than that of males); ventrals large and extending about half way across belly, and eachabout three times as wide as adjacent body scales; ventrals with a median keel and aposterior notch (but smooth in females); anal divided; subcaudals around 20-40 entire (maleshave a significantly higher number of subcaudals); venom apparatus advanced. Attains amaximum total length of around 1.7 m., and a snout-vent length of about 1.6 m., andmaximum body girth of around 25 cm. Specimens are sexually mature at around 700 mm(males) and 800 mm (females). Females are larger than males. It is possible that eastAustralian specimens are taxonomically distinct.Distribution: Largely confined to tropical Australian seas, being found off the coasts ofQueensland, Northern Territory and Western Australia (to about as far south as ExmouthGulf). It is occasionally observed in New South Wales waters to as far south as the Sydneyregion. Also known from the waters of New Guinea and the Coral Sea, as far out in the SouthWest Pacific as New Caledonia.Habitat: It mainly inhabits relatively shallow water coral reefs, the slopes of outer reefs andthe margins of rocky islands, but the exceptionally shallow fringing reef flats are less suitable.Preferred water depths are around 10-20 metres, although they have been commonly foundanywhere from 1m. to about 30 metres depths. Also known from sandy bottom coastal watersbut these areas are only traversed through the necessity of movements between its corehabitats of rocky or coral outcrops. Specimens are also occasionally detected in estuarineareas in relatively shallow continental waters.Biology/Ecology: This is essentially an abundant species, with males tending to be morecommonly encountered in the middle of the year and females more so in summer due to thespecies’ breeding behaviour/activity patterns. It may be either diurnal or nocturnal althoughthey are apparently more active later in the day than earlier, and specimens may be regularlysighted in the evenings apparently foraging for resting fishes. Occasionally specimens arefound encrusted with barnacles and algae on their skin, and this can make them less obviousamid the coral and rocks. The diet is fairly mixed, although clearly dominated by small fishes.It feeds on a variety of species of small fishes and on some occasions fish eggs, and evenmarine invertebrates, such as crabs and prawns. Captive specimens readily feed on fishes.Foraging occurs amongst coral or rock outcrops, usually for around 15 minutes durationbefore they need to return to the surface for air - although activity periods between breathsmay be as low as only a few minutes, up to around 30 minutes. Interestingly, it has also beenreported by fishermen to feed on discarded fish guts thrown overboard as well as fish heads,so it may forage for dead fish (carrion) as well. Hunting is mainly confined to the edges ofreefs, by slowly searching coral or rock crevices for sheltering fishes near sandy bottoms.When a fish is seized it is often constricted, and quickly swallowed - usually head-first - andthe snake immediately returns to the surface to breath. Although fishes slightly larger than thesnake’s head can be eaten, they usually prefer smaller size classes, rather than larger prey.Generally they do hunt in open water or chase free-swimming fish. It can occur in hugenumbers in suitable habitat, and has long been considered one of Australia’s most abundantspecies of sea snake. Despite large population sizes no reports of territorial or fightingbehaviour has been reported in this species. Although some sort of limited individual homerange activity occurs, home ranges of several individuals can overlap extensively without5

Australian Biodiversity Record, 2007 (8): 1-124apparent conflict. Shelter sites are for short term use only (hours or days) and are regularlyabandoned for more suitable sites elsewhere on the reef - although on occasions a site maybe continuously occupied for 2 or 3 weeks by a large individual. When under cover, they donot seem to tongue flick or otherwise seek prey - they appear to be resting until suitableconditions for open active foraging occurs. One of the more interesting physiological traits ofAipysurus species (both laevis and pooleorum) concerns use of its tail - when sheltering, asnake will often leave its tail exposed - and this is done both during the day and night,although it appears to be more common at night. Physiologists have discovered that the tailhas a unique photoreceptor just anterior to the end, but its precise function is at presentunclear. It may be some sort of metabolic regulating mechanism to allow extended periods ofsubmergence during periods of darkness, or it is possibly used as a predator/prey detectiondevice. When in repose under cover, a snake will leave its tail exposed at the edge of theshelter, and when the light intensity changes suddenly the tail is moved instantly, and thesnake appears to investigate the source of the lighting change. Reproduction is highlyseasonal in this ovoviviparous species. Mating occurs between May and August and thegestation period is around 9 months. Males actively court receptive females with vigour duringthe mating season, and at this time this species can be particularly inclined to respondaggressively to divers who may disturb their courting or mating activities. By Spring(September to October females are carrying yolky follicles, and oviducal young are carriedfrom mid Spring through Summer to Autumn when they are born. Litter size is variable and iscorrelated with size - larger females tend to produce higher numbers of young. From 1 to 11young may be produced in a litter, although the average is only around 6-7 large young in abrood. At birth they juveniles are quite large, at around 300-350 mm in length. Growth is rapidfor the first few years, but slows upon attaining sexual maturity. Males attain sexual maturityat around 3 years of age, while females are not sexually mature until around 4 or 5 years old.It would appear that most females only reproduce every 2nd year, and a life-span of around15 years in attained. Major predators are sharks and sea eagles; parasitic nematodes havealso been found in their stomachs but their effects if any are unknown.Toxicity: Although often credited as being one of the more ‘aggressive’ species of sea snake,this is actually a very curious species that has been known to leave protective shelter or coveron becoming aware of a diver and swim towards them while rapidly flicking its tongue. Othersmay be very shy and rapidly flee to the protection of a crevice when approached by a diver,until they can assess the extent of the threat. During apparently exploratory or curious activity,an Olive Sea Snake may also just casually swim right up to human divers and proceed totouch arms, legs and body with its tongue or head. Naturally, this can be somewhatdisconcerting to the more inexperienced divers - particularly when some snakes vigorouslycoil around a diver's arm or leg during their ‘investigations’. Divers may be tempted to overreactto the snake by attempting to fend it off with a violent panic response of arm-waving oreven by directly hitting the snake by kicking and punching it. This often has the effect ofsufficiently harassing the snake to the point where it may actually attempt to bite, so cautionshould be exerted, because its venom is highly toxic to humans. On the other hand, duringthe breeding season in Autumn - Spring, males can become very nervous to any perceivedthreat, and may on rare occasions seek to bite a diver swimming too close. Indeed, suchlarge males can be quite belligerent and on occasions have forced divers from the water withtheir repeated lunges and determined attempts at biting. So how does one discriminatebetween a ‘curious’ sea snake and one that is intent on biting? An indication of the snake’sintentions is about the best one can hope for, so it’s best to give all sea snakes a fairly wideberth if possible. As a general rule - but this is not always the case - the swimming pattern ofan attacking sea snake usually leaves no doubt as to its intentions, as it is much faster andmore direct than that of a curious specimen. When a genuine ‘attack’ occurs, it may biterepeatedly and with determination if sufficiently aroused, and with its relatively large teeth, awet-suit should not be considered as adequate protection from penetration by the fangs of anadult specimen. Similarly, specimens landed on trawlers vigorously bite anything within range,including other snakes, fishes and fishing gear, and those handled by researchers readily tryto bite if restrained or handled roughly, so urgent medical attention should always be sought ifone is unfortunate enough to receive a bite from this species. Although there are no knownhuman fatalities associated with its bite, tests have shown that the venom is extremely toxic.6

Australian Biodiversity Record, 2007 (8): 1-124Survival Status: Protected under the New South Wales National Parks and Wildlife Act (1974)but not listed in that State as a Threatened Species in any of the Schedules of the NSWThreatened Species Conservation Act (1995). Also protected under the Qld NatureConservation Act (1992), the Territory Parks and Wildlife Conservation Act (1998) and theWA Wildlife Conservation Act 1950 (as amended). Listed as a Marine Protected Speciesunder the Australian Environmental Protection and Biodiversity Conservation Act (1999).Although it is generally regarded as one of the commonest of all sea snake species andtherefore probably not under threat, its patchy distribution in Australian waters couldpotentially make it vulnerable in some parts of its range.Etymology: Aipysurus is derived from the Greek aipys meaning high, and ura for tail, andrefers to the laterally compressed tail that is higher than the depth of the body. The name'laevis' means 'smooth', and refers to the skin texture of the species.Aipysurus pooleorum Smith, 1974Aipsusus laevis pooleorum Smith, L.A. (1974): Rec. West. Aust. Mus. 3: 93-110 [p. 97]. Typedata: holotype WAM R21366, paratype(s) WAM 6 specimens. Type locality: Shark Bay, WA.Description: This is another relatively large and bulky species with a short deep head that isbarely distinct from the neck, although it doesn’t quite reach the large size of Aipysurus laevis.Generally, darker in colouration than its congenor Aipysurus laevis, the males and females ofA pooleorum differ significantly from each other in colouration. Males are uniform dark browndorsally and lighter brown along the lower lateral of the body. Females are more purplishdorsally, with pale barring along the flanks. Some significant features of this species'morphology are: Head shields large and symmetrical with only minor partial fragmentationposteriorly; prefrontals present, and occasionally divided; frontal occasionally divided;supraoculars divided; parietals divided; preoculars present (1-2); nasal not contactingpreocular, separated by the presence of a loreal; nasals in contact; 2-3 postoculars; 3 primarytemporals; supralabials 8-10; posterior chin-shields small and separated by one or more smallscales; portion of rostral scale bearing median, valve-like fold sometimes separated fromremainder of scale by suture; body scales smooth in females and imbricate and in about 20-23 rows at mid-body; body scales tuberculate in males; ventrals about 145-160; ventrals largeand extending about half way across belly, and each about three times as wide as adjacentbody scales; ventrals with a weak median keel and a posterior notch (but smooth in females);anal divided; subcaudals around 25-33 entire; venom apparatus advanced. Attains amaximum total length of only around 1.1 m., and a snout-vent length of about 95 cm, althoughthe largest would be about 1.2m. Despite its smaller maximum length, A pooleorum, like A.laevis can be quite bulky, with a mid-body girth of about 20 cm. in large specimens.Distribution: Known principally from around Shark Bay, on the mid-west coast of WesternAustralia, but sometimes found further south to about Perth.Habitat: It mainly inhabits coral reefs in relatively shallow continental waters.Biology/Ecology: A locally common, diurnal species that feeds on a variety of small fishes.Ovoviviparous, producing up to 2-4 living young in a brood.Toxicity: This species will readily attempt to bite if harassed, so caution should be exerted,because its venom is highly toxic to humans. Urgent medical attention should always besought in the event of a bite from this species.Survival Status: Protected under the WA Wildlife Conservation Act 1950 (as amended). Listedas a Marine Protected Species under the Australian Environmental Protection andBiodiversity Conservation Act (1999). Probably not under threat, but its highly restricteddistribution could potentially make it vulnerable.Etymology: The name 'pooleorum' honours the Australian fishermen W. Poole and W. Pooleof Fremantle who collected some of the Types of this species.7

Australian Biodiversity Record, 2007 (8): 1-124Further Notes: It was originally believed that this sea snake was merely a subspecies ofAipysurus laevis, but Wells and Wellington (1984) elevated it to specific status, a move soonafter followed by Storr, Smith and Johnstone (1986, 2002) and Ehmann (1992). Widespreadrecognition of its distinctiveness however, has been slow coming, but it is now accepted byrecent authorities as being a distinct species in its own right (see Wilson and Swan, 2003).Genus Emydocephalus Krefft, 1869Emydocephalus Krefft, G. (1869): Snakes of Australia [p. 92]. Type species: Emydocephalusannulatus Krefft, 1869 by subsequent designation [see Cogger, H.G. (1983): Zool. Cat. Aust.1. Amphibia and Reptilia [p. 246].Diagnosis: A genus of two very similar species of blunt-headed sea snakes (of which oneoccurs in Australia) in the family Aipysuridae, and readily identified by the followingcombination of characters: Head shield large and symmetrical; nasals in contact; preocularpresent; 3 supralabials (2nd contacting orbit); prominent conical projection on angulate rostralshield, and in males this takes the form of a conspicuous spine; posterior chin shields small,separated by one or more smaller scales; valvular nostrils and lingual fossa; body scalessmooth, imbricate and in 15-17 rows at mid-body; ventrals 125-145, large, at least 3 times thewidth of adjacent body scales; ventrals with small tubercles and a low median keel; analusually entire, but sometimes divided; subcaudals entire, about 20-40; tail stronglycompressed vertically, paddle-like; ovoviviparous; venom apparatus vestigial. Etymology: Thename Emydocephalus is derived from the Greek ‘emys’ for turtle, and ‘cephale’ for ‘head’, andrefers to the unique head-shape (in profile) of its included species. Content: Emydocephalusannulatus Krefft, 1869; Emydocephalus ijimae Stegneger, 1898. Content in Australia:Emydocephalus annulatus Krefft, 1869.Emydocephalus annulatus Krefft, 1869Emydocephalus annulatus Krefft, G. (1869): Snakes of Australia [p. 92]. Type data: syntypesAM R454, AM R6633. Type locality: probably … Australian seas.Emydocephalus tuberculatus Krefft, G. (1869): Snakes of Australia [p. 93]. Type data:holotype AM R455. Type locality: probably … Australian seas.Aipysurus chelonicephalus Bavay, A. (1869). Mém. Soc. Linn. Normandie 15: 1-37 [p. 34].Type data: Holotype - Marine Museum Brest. Type locality: Lifu (as Lufou), Loyalty Ils, NewCaledonia.Description: This is a relatively small, highly specialized sea snake of shallow water coralreefs. Its body-form is more robust than most species, but its head is short and thick-set.When viewed from the side, the head is reminiscent of a sea turtle and this is due to theenlarged plate-like supralabials and angulate rostral area – hence the common name of‘turtle-headed seasnake’. There are both banded and unbanded forms. However, mostspecimens are marked with creamish-yellow bands on a dark brown or purplish-blackbackground over the entire body and tail, creating a black and yellow ringed effect. Somespecimens however, are unbanded and may be either a uniform dark grey, or various shadesof brown to even black in overall colour. In the case of banded specimens, the dark zones cancontain pale cream or white blotching, and the paler interspaces can contain brownishblotching. In banded specimens the head is usually pale creamish to yellow dorsally with darkbrown speckles. There is a dark brown temporal stripe which sometimes extends from just infront of the eye, over the eye to the nape where it meets the first dark transverse band. Somesignificant features of this species’ morphology are: Head shield large and symmetrical;nasals in contact; preocular present; ; 3 supralabials (2nd the largest, and contacting orbit);prominent conical projection on angulate rostral shield, and in males this takes the form of aconspicuous spine, whereas in females the snout is more rounded; posterior chin shieldssmall, separated by one or more smaller scales; body scales smooth with longitudinal alignedseries of tiny tubercles, imbricate and in 15-17 rows at mid-body; ventrals 125-145, large, atleast 3 times the width of adjacent body scales; ventrals with small tubercles and a lowmedian keel; anal usually entire, but sometimes divided; subcaudals entire, about 20-408

Australian Biodiversity Record, 2007 (8): 1-124(males have significantly higher subcaudal counts than females). Attains a maximum totallength of only around 75 cm., although an exceptional specimen around 90 cm has beenrecorded.Distribution: Largely confined to tropical Australian seas, particularly between the Timor Seaand Arafura Sea, and extending into the Coral Sea to about New Caledonia. It occurs patchilyoff the coasts of Queensland, Northern Territory and Western Australia (to about as far southas Shark Bay), and has been occasionally detected in New South Waters as well.Habitat: Mainly inhabits tropical coral reefs, usually in relatively shallow continental waterswith sandy bottoms and less than 30 metres in depth. Commonly observed active in depthsranging from 1m to around 15 m. on the Great Barrier Reef.Biology/Ecology: This is essentially a diurnal species that readily burrows into the sandybottoms of the sea for shelter, as well as for its sole food - the eggs of fishes such as Blenniesand Gobies. Foraging is concentrated at the sandy margins of coral reefs, as well as in moreexposed open areas of sandy bottom, but feeding is reduced or even ceases during thebreeding season. Concealed nests of eggs are detected by head probing and tongue-flickinginto burrows or other suitable fish-egg-laying sites, with food being located by smell ratherthan sight. Large numbers of eggs (in the thousands) can be consumed at a feeding episodeby mature individuals, but interestingly, different size classes of snake appear to consumedifferent size classes of eggs. This could indicate that different species of fish eggs areutilized as the snake grows, or there may be some geographic/seasonal variation in diet overthe species range. The eggs are also removed from coral and rocks by using its specializedsupralabial plates as a scraping device and then eaten as they float away, or more usuallydirectly taken from holes (nests) in the sandy substrates. Sometimes moderately large groupsare found in loose aggregations following currents that run through sandy-bottomed gullies orchannels in coral reefs, and on contacting one another there seems to be some sort ofrecognition behaviour practiced by each touching snouts, before returning to their respectiveforaging patterns. Submergence times can be up to an hour before they need to quickly returnto surface to breathe for a few seconds before submerging again, although most foragingepisodes would be under 30 minutes in duration. This species is essentially a benthic speciesthat spends as much time as possible active on the bottom - only racing to the surface tobreathe before returning to the safety of the sand. It rests amid pieces of coral or on/underthe sand. They are active year-round - although there does appear to be seasonal changes inpreferred water depths, with shallower waters being utilised during the Winter months anddeeper waters in the hotter months. Courtship and mating can occur in June-July, or as lateas September (Dry Season), and when attempting to mate, males gently prod the head, neckand forebody of a receptive female with its pointed snout as a stimulus. Gravid females havebeen found in December (Wet Season) and small litters of live young (2-4) are apparentlyproduced in the beginning of the Dry Season. On occasions specimens may become carriersof barnacles and algal growth on the skin.Toxicity: This species is totally inoffensive in temperament and so rarely if ever attempts tobite. In any case, this species is technically regarded as non-venomous, as its specialized dietof fish-eggs has led to the reduction of fangs and venom glands to the stage where theiroriginal function is now apparently lost. They do however still possess tiny venom glands.Survival Status: Protected under the Qld Nature Conservation Act (1992), the Territory Parksand Wildlife Conservation Act (1998), the WA Wildlife Conservation Act 1950 (as amended).Also protected under the New South Wales National Parks and Wildlife Act (1974) but notlisted in that State as a Threatened Species in any of the Schedules of the NSW ThreatenedSpecies Conservation Act (1995). Listed as a Marine Protected Species under the AustralianEnvironmental Protection and Biodiversity Conservation Act (1999). Probably not underthreat, but its restricted distribution in Australian waters, and its specialised ecology couldpotentially make it vulnerable.Etymology: The name 'annulatus' (from the Latin ‘anus’ = ring) is in reference to the bodypattern of dark annular rings.9

Australian Biodiversity Record, 2007 (8): 1-124Oceanius gen. nov.Type Species: Aipsurus tenuis Lönnberg, E. and Andersson, L.G. (1913). K. Sven. Vetensk.-Akad. Handl. 52(3): 1-173 [p. 13].Diagnosis: A monotypic genus of small Aipysurid sea snake readily identified by the followingcombination of characters: head shields enlarged and symmetrical, with each head shieldlarger than nape scales (not partly fragmented as in Smithohydrophis); snout rounded (vspointed in Smithohydrophis); valvular nostrils and lingual fossa; frontal divided; parietalsdivided; supraoculars divided; nasals in contact; loreal present (vs absent inSmithohydrophis); 1 preocular; 2 postoculars; temporals small; prefrontals present; posteriorchin shields small and separated by smaller scales; body scales smooth and imbricate in 19rows at mid-body, with the lowest rows of males tuberculate; ventral scales about three timesas wide as the rest of the body scales; ventrals with a slight median notch and keel,tuberculate along the hind edge; subcaudals entire; tail strongly compressed vertically,paddle-like; venom apparatus advanced and highly toxic; ovoviviparous. Etymology: Oceanusin classical mythology was the Son of Heaven and Earth (Uranus and Ge) and Lord of theWater that encircled the World. Content: Oceanius tenuis comb. nov. (Lonnberg andAndersson, 1913).Oceanius tenuis comb. nov. (Lonnberg and Andersson, 1913)Aipsurus tenuis Lönnberg, E. and Andersson, L.G. (1913). K. Sven. Vetensk.-Akad. Handl.52(3): 1-173 [p. 13]. Type data: syntypes NHRM 2400 3 specimens. Type locality: CapeJaubert, nr Broome, WA.Description: This is usually a moderate-sized species of sea snake, in which the body-form isrelatively elongate, the head small with a somewhat rounded snout, and the head is notdistinct from the neck. Overall the base body colour is usually purplish, with some being moredark olive-brown. There is a series of narrow or broad, light olive-brown transverse bands,which tend to be less distinct dorsally, and most prominent laterally. In some specimens twonarrow light bands may be very close together with a narrow darker interspace purplishbrown,followed by a broader interspace of dark purplish-brown, then another two narrowlighter bands and so on. In others, these narrow light bands may partially or completelycoalesce to form a series of broad pale bands, or in some a mixture of thin and broad bands.Occasional specimens may have scattered whitish or creamish lateral or ventrolateral scales,or in others, the body scales may have darker centres at the scale-tip aligned longitudinally toform a series of faint body-length striations or lines along the dorsum. The ventral colour isgreyish-brown, with only scattered splotches of creamish-white on the venter, but with the(hidden) anterior edge of each ventral pale creamish; the throat has a speckled appearance,with the scales whitish anteriorly and brownish posteriorly. Some significant features of thisspecies' morphology are: head shields enlarged and symmetrical, with each head shieldlarger than nape scales; frontal divided; parietals divided; supraoculars divided; nasals incontact; loreal present; 1 preocular; 2 postoculars; temporals small; prefrontals present;posterior chin shields small and separated by smaller scales; body scales smooth andimbricate in 19 rows at mid-body, with the lowest rows of males tuberculate; ventral scalesabout three times as wide as the rest of the body scales; ventrals with a slight median notchand keel, tuberculate along the hind edge, and numbering 185-194; anal divided; subcaudalsentire, 36-37. Attains a maximum total length of around 1m.Distribution: Largely confined to tropical Australian seas, between the Arafura Sea, NorthernTerritory and north-west Western Australia, south almost to the Dampier Archipelago.Habitat: Mainly inhabits reefs as well as the vicinity of muddy deltas in relatively shallowcontinental waters.Biology/Ecology: Forages diurnally for a variety of small fishes in relatively shallow inshoreand near shore reefs and outflows from deltas. Ovoviviparous, producing up to 4 living youngin a brood.10

Australian Biodiversity Record, 2007 (8): 1-124Toxicity: This species would probably attempt to bite if annoyed, so caution should beexerted, because its venom is likely highly toxic to humans. Urgent medical attention shouldalways be sought in the event of a bite from this species.Survival Status: Protected under the Territory Parks and Wildlife Conservation Act (1998) andthe WA Wildlife Conservation Act 1950 (as amended). Listed as a Marine Protected Speciesunder the Australian Environmental Protection and Biodiversity Conservation Act (1999).Probably not under threat, but its restricted distribution in Australian waters could potentiallymake it vulnerable in some parts of its range.Etymology: The name 'tenuis' means 'slender', and refers to the body-form of the species.Genus Pelagophis Peters and Doria, 1878Pelagophis Peters, W. and Doria, G. (1878): Ann. Mus. Civ. Stor. Nat. Genova 13: 323-450[p. 413 pl. 5]. Type species: Pelagophis lubricus Peters and Doria, 1878 by monotypy.Diagnosis: A genus of marine snakes of the Family Aipysuridae readily separated from allother genera by the following combination of characters: Head shields mostly fragmented intosmaller scales (except rostral and nasals), with each fragment about the same size as napescales; nasals in contact; supralabials 8-9; posterior chin shields small and separated bysmaller scales; valvular nostrils and lingual fossa; body scales mostly smooth (but some bodyscales may be slightly keeled or with a centrally-aligned row of tubercles) and imbricate in 19rows at mid-body; ventral scales about three times as wide as the rest of the body scales;ventrals 150-180, only weakly notched, and with a shallow median keel; anal divided;subcaudals entire; tail strongly compressed vertically, paddle-like; venom apparatusadvanced and highly toxic; ovoviviparous. Etymology: The name Pelagophis is from theGreek, pelag - ‘sea’, and ophis - ‘snake’. Content: Pelagophis duboisii (Bavay, 1869).Pelagophis duboisii (Bavay, 1869)Aipysurus duboisii Bavay, A. (1869) : Mém. Soc. Linn. Normandie 15: 1-37 [p. 33]. Type data:holotype Marine Museum Brest [see Smith, M.A. (1926). Monograph on the Sea Snakes.(Hydrophiidae). London : British Museum xvii 130 pp.]. Type locality: Lifu (as Lifou), LoyaltyIls, New Caledonia.Aipysurus australis Sauvage, H.E. (1877): Bull. Soc. Philomath. Paris (7)1: 107-115 [p. 114].Type data: holotype MNHP (not found). Type locality: Australia.Pelagophis lubricus Peters, W. and Doria, G. (1878): Ann. Mus. Civ. Stor. Nat. Genova 13:323-450 [p. 414, pl. 5] [erroneously referred to Aipysurus laevis by Capocaccia, L. (1961):Ann. Mus. Civ. Stor. Nat. 'Giacomo Doria' 72: 86-111]. Type data: holotype MCG C.E.30765.Type locality: Yule Is., Papua New Guinea.Description: This is medium-sized species of sea snake, in which the body-form is moderatelyelongate, the head short with a rounded snout, and the head distinct from the neck. Overallthe base body colour is usually pale brown or brownish-white, with the head darker brown.Pattern may be present or absent, and when present is somewhat variable. In somespecimens the base colour appears pale creamish-white or even salmon pink, with bodyscales having a brownish spot and so forming a series of obscure darker bands over thebody. In most individuals however, each body scale is creamish-white on its concealedanterior part, the pale colour being just visible and so forming a vague light reticulum over abrownish or purplish body. Some of the scales along the lower lateral of the body arecreamish-white, and these may be clustered together in triangular or wedge-shapes, forminga point at the upper lateral. The overall effect of this pattern is that of broad dark transversebanding with paler interspaces, such that there is the appearance of a series of narrow orbroad, purple or brownish transverse bands, which tend to coalesce dorsally, and be moreprominent laterally where they narrow to a point; the creamish triangular lower lateral patternforms the paler interspaces. Ventrally creamish or brownish under the body, but white underthe throat, with each scale brownish-tipped. Some significant features of this species'11

Australian Biodiversity Record, 2007 (8): 1-124morphology are: Head shields mostly fragmented into smaller scales (except rostral andnasals), with each fragment about the same size as, or smaller than, the nape scales; nasalsin contact; supralabials 8-9; posterior chin shields small and separated by smaller scales;body scales mostly smooth (but some body scales may be slightly keeled or with a centrallyalignedrow of tubercles) and imbricate in 19 rows at mid-body; ventral scales about threetimes as wide as the rest of the body scales; ventrals only weakly notched and with a shallowmedian keel; ventrals 150-180; anal divided; subcaudals entire, 23-35. Attains a maximumtotal length of only about 1.1 m. and a snout-vent length of around 1 m. (although individualsaround 70-80 cm overall length are sexually mature).Distribution: Known from a large area of tropical Australian seas, being found off the coasts ofQueensland, Northern Territory and Western Australia and rarely northern New South Wales.Also occurs in New Guinea waters as well as the Coral Sea and around islands of the South-West Pacific Ocean to about New Caledonia.Habitat: Mainly inhabits reef flats and coral reefs in relatively shallow continental shelf waters,reef lagoons and fringing platforms, from a depth of about 1 metre to 20 metres, butsometimes reaching depths of 50m. At low tides they can also be found temporarily trapped invery shallow pools of only a few centimetres depth and have been observed attempting tocrawl over exposed corals to deeper water.Biology/Ecology: This is essentially a diurnal or crepuscular species that feeds on a variety ofsmall fishes, including eels, and even fish eggs. Fishes are usually hunted on the sea bottomaround the edges of coral during daylight and early evening. Its sedentary nature allows quitea range of marine organisms (algae, bryozoans, polychaetes and tube worms) to becomeattached to its body, making it sometimes hard to detect amongst the coral. It isovoviviparous, with gravid females being found in the late Wet Season (February-March), andlitter sizes ranging from 2-6 - average of about 4 or 5 - relatively large living young in a brood.Toxicity: There are no known human fatalities from the bite of this species. It is usuallyinoffensive, but caution should nevertheless be exerted, because it has been known to exhibitaggressive behaviour towards divers without any apparent provocation. It will definitely bitewhen carelessly handled and as its venom is highly toxic to humans, urgent medical attentionshould always be sought in the event of a bite.Survival Status: Protected under the Qld Nature Conservation Act (1992), the Territory Parksand Wildlife Conservation Act (1998) and the WA Wildlife Conservation Act 1950 (asamended). Also protected under the New South Wales National Parks and Wildlife Act (1974)but not listed in that State as a Threatened Species in any of the Schedules of the NSWThreatened Species Conservation Act (1995). Listed as a Marine Protected Species underthe Australian Environmental Protection and Biodiversity Conservation Act (1999). It isprobably not under direct threat, but its restricted distribution in Australian waters couldpotentially make it vulnerable in some parts of its range to localized impacts such as canoccur with some intensive fishing operations.Etymology: The name 'duboisii' honours the 19th century French naturalist Charles FredericDubois.Genus Smithohydrophis Kharin, 1981Diagnosis: This is a genus of small elongate species of marine snakes of the familyAipysuridae, believed to be restricted to coral reefs of north-western Australia. All have smallheads with pointed snouts, head shields more or less symmetrical and always larger thanneck scales, imbricate dorsal scales with pointed or bifid posterior edges, and ventrals at least3 times wider than adjacent body scales. This new genus can be readily separated from allother genera by the following combination of characters: Body-form moderately elongate;head small with a pointed snout; head not distinct from neck; head shields large and mostlysymmetrical, but partly fragmented; each shield nevertheless larger than nape scales; nasalsin contact; valvular nostrils and lingual fossa; prefrontals either absent (fused to nasals) or ifpresent, fragmented; loreal absent; temporals small; preocular scale present, wider than high;12

Australian Biodiversity Record, 2007 (8): 1-1242 postoculars, the lower being the largest; median, valve-like fold present on part of rostral,and sometimes separated from remainder of rostral by suture; supralabials 7-8; posterior chinshields small and separated by smaller scales; body scales smooth and imbricate in 17-21rows at mid-body, with the hind edge of each scale pointed or bifid, and with a short mediankeel posteriorly; ventral scales about three times as wide as the adjacent body scales;ventrals deeply notched, and ragged-edged on the posterior margins and numbering 140-194;anal divided; subcaudals entire, 18-37; tail strongly compressed vertically, paddle-like;ovoviviparous. Etymology: The name 'Smithohydrophis' recognizes the pioneering work ofMalcomb A. Smith with all sea snakes. Content: Smithohydrophis apraefrontalis comb. nov.(Smith, 1926); and Smithohydrophis foliosquama comb. nov. (Smith, 1926).Smithohydrophis apraefrontalis comb. nov. (Smith, 1926)Aipysurus apraefrontalis Smith, M.A. (1926). Monogr. Sea Snakes. (Hydrophiidae). [p. 24, fig.13]. Type data: holotype BMNH 1946.1.1.94. Type locality: Ashmore Reefs, Australia.Description: This is quite a small species of sea snake, in which the body-form is moderatelyelongate, the head small with a pointed snout, and the head is not distinct from the neck.Overall the base body colour is usually purplish, with some being more dark olive-brown.There is a series of about 24-27 narrow or broad, light olive-brown transverse bands, whichtend to be less distinct dorsally, and most prominent laterally, where they taper to a point. Insome specimens two narrow light bands may be very close together with a narrow darkerinterspace purplish-brown, followed by a broader interspace of dark purplish-brown, thenanother two narrow lighter bands and so on. In others, these narrow light bands may partiallyor completely coalesce to form a series of broad pale bands, or in some a mixture of thin andbroad bands. Occasional specimens may have scattered whitish or creamish lateral orventrolateral scales. The ventral colour is greyish-brown, with only scattered splotches ofcreamish-white on the venter, but with the (hidden) anterior edge of each ventral palecreamish; the throat has a speckled appearance, with the scales whitish anteriorly andbrownish posteriorly. Some significant features of this species' morphology are: head shieldslarge and mostly symmetrical, but with parietals partly fragmented, but each shieldnevertheless larger than nape scales; nasals in contact; no prefrontals (fused to nasals);nasals contacting frontal; temporals small; preocular scale present, wider than high; lorealabsent; preocular contacts nasal; 2 postoculars, the lower being the largest; supralabials 7;posterior chin shields small and separated by smaller scales; body scales smooth andstrongly imbricate in 17 rows at mid-body (males and females have the same mid-body scalecount), with the hind edge of each scale pointed or bifid, and with a short keel posteriorly;ventral scales about three times as wide as the rest of the body scales; ventrals deeplynotched, and ragged-edged on the posterior margins and numbering 140-160; anal divided;subcaudals entire, 18-25. Attains a maximum total length of around 1.1 m., and a snout-ventlength of only about 98 cm, although smaller mature specimens around 50-60 cm in totallength are more often observed.Distribution: Known only from continental shelf reefs off the north-west of Western Australia,in particular Ashmore Reef, south to about Exmouth Gulf. Reputedly occurs around theeastern parts of Indonesia.Habitat: Mainly inhabits the edges or flat areas of coral reefs in relatively shallow waters fromabout 1 m to less than 10 m. in depth. More often associated with sparse coral outcroppingsamid extensive sandy bottoms.Biology/Ecology: This is essentially a nocturnal species but it can also occasionally beobserved active during the day. Specimens have been observed hiding amongst coral, and assome may have growths of algae on their body, they can be difficult to detect. Specimensrest/hide in cavities or coral growths and overhangs or beneath broken coral in very shallowwater during low tides. It feeds on a variety of small fishes - mainly comprising eels.Ovoviviparous, producing up to 4 living young in a brood.13

Australian Biodiversity Record, 2007 (8): 1-124Toxicity: This species rarely attempts to bite, but caution should nevertheless be exerted,because its venom is likely highly toxic to humans. Urgent medical attention should always besought in the event of a bite from this species.Survival Status: Protected under the WA Wildlife Conservation Act 1950 (as amended). Listedas a Marine Protected Species under the Australian Environmental Protection andBiodiversity Conservation Act (1999). Probably not under threat, but its restricted distributionin Australian waters could potentially make it vulnerable in some parts of its range.Etymology: The name 'apraefrontalis' means 'without prefrontal', in reference to the headshields of this species.Smithohydrophis foliosquama comb. nov. (Smith, 1926)Aipysurus foliosquama Smith, M.A. (1926). Monogr. Sea Snakes. (Hydrophiidae). [p. 22, figs11-12 pl. 2 fig. 3]. Type data: holotype BMNH 1946.1.1.96. Type locality: Ashmore Reefs,Australia.Description: This is quite a small species of sea snake, in which the body-form is moderatelyelongate, the head small with a pointed snout, and the head is barely distinct from the neck.Overall the base body colour is usually purplish, with some being more dark olive-brown.There is a series of narrow or broad, light olive-brown transverse bands, which tend to be lessdistinct dorsally, and most prominent laterally. In some specimens two narrow light bandsmay be very close together with a narrow darker interspace of purplish-brown, followed by abroader interspace of darker purplish-brown, then another two narrow lighter bands and soon. In others, these narrow light bands may partially or completely coalesce to form a seriesof broad pale bands, or in some a mixture of thin and broad bands. Generally, the bands tendto be most intense on the lower flanks. Occasional specimens may have scattered whitish orcreamish lateral or ventrolateral scales between the dark crossbands. The ventral colour isgreyish-brown, with only scattered splotches of creamish-white on the venter, but with the(hidden) anterior edge of each ventral pale creamish; the throat has a speckled appearance,with the scales whitish anteriorly and brownish posteriorly. Some significant features of thisspecies' morphology are: head shields large and partly symmetrical, but with parietals,prefrontals and frontal partly fragmented, but each shield nevertheless larger than napescales; nasals in contact; prefrontals present, but usually fragmented, with pairs between thefrontal and the nasals; preocular scale present, loreal absent; not in contact with nasal;postoculars 2; supralabials 7-8; posterior chin shields small and separated by smaller scales;body scales smooth and strongly imbricate in 19-21 rows at mid-body mid-body (males andfemales do not appear to have significantly different mid-body scale counts), with the hindedge of each scale pointed or bifid, and with a short median keel; ventral scales about threetimes as wide as the rest of the body scales; ventrals deeply notched, and ragged-edged onthe posterior margins and numbering around 135-155 (females do not appear to have ahigher ventral count than that of males); anal divided; subcaudals entire, 20-29 (males havesignificantly more subcaudals than females). Attains a maximum total length of only around55 cm., and a snout-vent length of about 47 cm.Distribution: Apparently restricted to large continental shelf reefs, (Ashmore Reef, HiberniaReef and Scott Reef), off the coast of north-west Western Australia.Habitat: Mainly inhabits the edges continental shelf reefs or reef flats in relatively shallowwaters less than 10 metres in depth. It is usually found in association with dense coral growth.Biology/Ecology: Essentially a diurnal species, it feeds on a variety of small fishes which itactively hunts during the day. Specimens are occasionally found with a growth of algae overthe skin, and no doubt this assists in camouflaging the animal as it forages amongst the coral.Ovoviviparous, producing up to 4 large living young in a brood.Toxicity: This is a small inoffensive species that rarely attempts to bite, but caution shouldnevertheless be exerted, because its venom is likely toxic to humans. No known fatalities14

Australian Biodiversity Record, 2007 (8): 1-124have been recorded from this species, but urgent medical attention should always be soughtshould envenomation occur.Survival Status: Protected under the WA Wildlife Conservation Act 1950 (as amended). Listedas a Marine Protected Species under the Australian Environmental Protection andBiodiversity Conservation Act (1999). Probably not under threat, but its restricted distributionin Australian waters could potentially make it vulnerable in some parts of its range.Etymology: The name 'foliosquama' means 'leaf-scale', in reference to the stronglyoverlapping body scales in this species.Genus Stephanohydra Tschudi, 1837Stephanohydra Tschudi, J.J. von (1837): Arch. Naturgesch. 3: 331-335 [p. 335]. Typespecies: Stephanohydra fusca Tschudi, 1837 by monotypy.Diagnosis: A genus of relatively short but very large (robust-bodied) marine snakes of theFamily Aipysuridae, readily separated from all other genera by the following combination ofcharacters: Head broad and deep, barely distinct from the neck with shields large andsymmetrical with only minor (partial) fragmentation posteriorly; frontal usually divided;supraoculars divided; parietals divided; prefrontals present, and occasionally divided; nasalsin contact; portion of rostral scale bearing median, valve-like fold sometimes separated fromremainder of scale by suture; valvular nostrils and lingual fossa; usually 1 preocular; lorealpresent; 2 postoculars; temporals small; supralabials 6-9; posterior chin-shields small andseparated by one or more small scales; body scales smooth in females and imbricate (but thelower lateral rows may be weakly keeled) and usually in 19 rows at mid-body (vs usually 21-25 rows in Aipysurus); ventrals about 155-175 (vs always less than 155 in Aipysurus);ventrals large, about three times as wide as adjacent body scales, and each with a weakmedian keel and a shallow posterior notch (but smooth in females); anal divided; subcaudalsaround 25-35 entire; tail strongly compressed vertically, paddle-like; venom apparatusadvanced and highly toxic; ovoviviparous. Content: Stephanohydra fusca Tschudi, 1837.Stephanohydra fusca Tschudi, 1837Stephanohydra fusca Tschudi, J.J. von (1837): Arch. Naturgesch. 3: 331-335 [p. 335 pl. 8].Type data: holotype ZMB 2824. Type locality: Asia (Sulawesi), [see Smith, M.A. (1926).Monogr. on the Sea Snakes. (Hydrophiidae); also erroneously referred to an Ashmore Reefspecimen as the type].Description: This is a thick-set or stout-bodied sea snake with a broad, deep head that isbarely distinct from the neck. The snout is rounded and blunt in profile. Mature specimens areusually uniform brownish on the dorsal of the body, with a series of faint transverse bandspresent on the lower flanks. The base colouration can sometimes have a purplish sheen, oreven be more of a darker almost blackish brown. On the lateral of the body, each scale canhave a darker centre, and this can create a pattern of very faint body-length striations. Theventer is brown. Some significant features of this species' morphology are: head shields moreor less regular and symmetrical, with the partial fragmentation of the parietals the mostsignificant; prefrontals present, and occasionally divided; frontal usually divided; supraocularsdivided; parietals divided; usually 1 preocular; loreal present; 2 postoculars; temporals small;supralabials 6-9; body scales smooth and imbricate (but the lower lateral rows may be weaklykeeled) and usually in 19 rows at mid-body (but rare specimens with 21 rows are known)(however, males and females do not appear to have significantly different mid-body scalecounts); ventrals about 155-175 (females do not have a significantly higher ventral count thanmales); ventrals about three times as wide as adjacent body scales, and each with a weekmedian keel and a shallow posterior notch (but smooth in females); anal divided; subcaudals20-40 entire (males have significantly higher subcaudal counts than females). A maturespecimen is around 60 cm in total length, although it can attain a maximum total length ofaround 95 cm. (snout-vent length of about 83 cm.).15

Australian Biodiversity Record, 2007 (8): 1-124Distribution: Mainly restricted to the larger reefs, (Ashmore Reef and Scott Reef), off the coastof north-west Western Australia. Also known from eastern Indonesia.Habitat: Inhabits the edges of coral and rocky reefs, coral reef lagoons and reef flats inrelatively shallow continental shelf waters around 10 metres or less in depth. Prefers areas offairly rich, dense coral growth.Biology/Ecology: This is an essentially diurnal species that feeds on a variety of small fishesand their eggs, principally of the families Gobiidae and Labridae. It is active throughout theyear and is ovoviviparous, producing up to 4 large living young in a brood. Sometimesspecimens are found with a growth of algae over parts of its skin.Toxicity: This is another small and inoffensive species that is usually reluctant to bite unlessroughly handled, but caution should nevertheless be exerted, because its venom is likely toxicto humans. Urgent medical attention should always be sought in the event of a bite from thisspecies.Survival Status: Protected under the WA Wildlife Conservation Act 1950 (as amended). Listedas a Marine Protected Species under the Australian Environmental Protection andBiodiversity Conservation Act (1999). Probably not under threat, but its restricted distributionin Australian waters could potentially make it vulnerable in some parts of its range.Etymology: The name 'fusca' means 'dark', and refers to the colouration of the species.Genus Tomogaster Gray, 1849Tomogaster Gray, J.E. (1849): Cat. Spec. Snakes Coll. Brit. Mus. [p. 59]. Type species:Tomogaster eydouxii Gray, 1849 by monotypy.Diagnosis: A genus of highly specialized marine snakes of the Family Aipysuridae, andreadily separated from all other genera by the following combination of characters: head shortand deep, snout rounded, with dentition reduced for its specialized diet; head not distinct fromthe neck; body-form moderately robust; head shields (including parietals) enlarged andsymmetrical (not fragmented); nasals contacting; prefrontals present; 1 preocular; 2postoculars; temporals small; primary temporals 1-3; supralabials 6; valvular nostrils andlingual fossa; body scales smooth, imbricate and in 17 rows at mid-body; ventrals around125-165; ventrals about three times the width of adjacent body scales, and each with ashallow keel in the centre; the ventrals usually lack notching on the posterior edge, but ifpresent it is shallow; anal divided; tail strongly compressed vertically, paddle-like; subcaudals23-34; venom apparatus present but rudimentary; ovoviviparous. Etymology: The name‘Tomogaster’ literally means ‘on the stomach’, and probably refers to the large ventral scalesof the only species in the genus. Content: Tomogaster eydouxii Gray, 1849.Tomogaster eydouxii Gray, 1849Tomogaster eydouxii Gray, J.E. (1849): Cat. Spec. Snakes Coll. Brit. Mus. [p. 59]. Type data:holotype BMNH 1946.1.6.86. Type locality: Indian Ocean.Thalassophis anguillaeformis Schmidt, P. (1852): Abh. Geb. Naturw. Hamburg 2: 69-86 [p.76, pl. 1]. Type data: holotype ZMH 386 [see Smith, M.A. (1926). Monogr. Sea Snakes.(Hydrophiidae)] Type locality: Java, Indonesia.Thalassophis muraeneformis Schmidt, P. (1852): Abh. Geb. Naturw. Hamburg 2: 69-86 [p.77]. Type data: holotype ZMH 385 (see Smith, M.A. (1926). Monogr. Sea Snakes.(Hydrophiidae)]. Type locality: Samarang, Java, Indonesia.Aipysurus margaritophorus Bleeker, P. (1858). Natuurk. Tijdschr. Ned.-Indie. (4)16: 49-50 [p.49]. Type data: holotype BMNH 1946.1.7.1. Type locality: Java, Indonesia.16

Australian Biodiversity Record, 2007 (8): 1-124Description: This is a very small species of sea snake, with a short, deep head that is notdistinct from the neck, and a moderately robust body-form. The head is usually brownish, withor without darker brown flecking. The base body colour is variable from one region to another,and may be cream to pinkish-brown, with prominent darker banding along the entire body.The series of broad transverse bands are black or dark slaty grey, and these bands may taperto a point at the ventrolateral margin, or extend partly onto the ventrals. In the paleinterspaces between the dark bands, the scales may be either faintly edged with black, ormore solidly marked with short black streaks, or on occasions blackish blotches; thesesmaller markings are usually more intense along the lower flanks. The venter is usually darkgrey to black, with the median area being darkest; on occasions specimens may have a darkcreamish venter. Some significant features of this species' morphology are: head shieldsenlarged and symmetrical; nasals contacting; prefrontals present; 1 preocular; 2 postoculars;temporals small; primary temporals 1-3; supralabials 6; body scales smooth, imbricate and in17 rows at mid-body (males and females have the same mid-body scale count); ventrals 125-165 (females have a higher ventral count than males); ventrals about three times the width ofadjacent body scales, and each with a shallow keel in the centre; the ventrals usually lacknotching on the posterior edge, but if present it is shallow; anal divided; subcaudals 23-34(males have slightly more subcaudals than females). Attains a maximum total length of onlyaround 90 cm., and a snout-vent length of about 80 cm., but the average adult total lengththough only rarely exceeds 50 cm. Females are sexually mature at 47 cm SVL. Variation inmorphology suggest that this species is composite.Distribution: Largely confined to tropical Australian seas, being found off the coasts ofQueensland, Northern Territory and Western Australia. Also widely known in South-EastAsian waters from about Vietnam to New Guinea.Habitat: Often associated with inshore areas of strong current or muddy conditions incontinental shelf waters. It can be particularly common around major river deltas, muddybottomed bays and estuaries of fairly shallow water depths ranging from about 3m. to 22 m. Itis also known from deeper waters of around 30 to 50 metres depth as well, and has beenobserved even living in the upper reaches of major tidal rivers - up to 10 or more kilometresfrom the coast. It has mainly been observed in association with muddy bottoms, and hasseldom been observed with other substrates.Biology/Ecology: This is mainly a diurnal species that feeds only on the eggs of fishes,although there is a record of an eel being taken. It is ovoviviparous, with gravid specimensbeing found in Australian waters during the middle of the year (May-August, Dry Season) incontrast to most other sea snakes that tend to be gravid during the Wet Season. In variousparts of Asia there is variation in the timing of reproduction, with gravid females being found inJanuary to April (late winter Monsoon Season) in the Straits of Malacca, but with none gravidin April in the South China Sea. In Australia, litter sizes range from 1 to 9, with the averagebeing about 4 relatively large living young being produced at a time, and larger females tendto produce larger litters. Litter sizes in Asia however, appear to be slightly higher (up to 12,but with a similar average of only about 4 or 5), which could indicate that Asian populationsmay attain a slightly larger maximum size. In Australia, there is some evidence to suggest thatfemales significantly outnumber males.Toxicity: Although this species is most reluctant to bite, it does actually possess smallfunctional venom glands, so caution should be exerted when handling this species. Despitethe fact that there are no known human fatalities due to the bite of this species - possiblybecause of the rudimentary nature of its venom apparatus - medical assistance should besought in the event of a bite as a precaution.Survival Status: Protected under the Qld Nature Conservation Act (1992), the Territory Parksand Wildlife Conservation Act (1998) and the WA Wildlife Conservation Act 1950 (asamended). Listed as a Marine Protected Species under the Australian EnvironmentalProtection and Biodiversity Conservation Act (1999). Probably not under threat, but itsrestricted distribution in Australian waters could potentially make it vulnerable in some parts ofits range.17

Australian Biodiversity Record, 2007 (8): 1-124Etymology: The name 'eydouxii' honours the French naturalist Joseph Fortune TheodoreEydoux (1803-1841).Family EphalophiidaeThis Family contains only the genera Ephalophis Smith, 1931, and Parahydrophis Burger andNatsuno, 1974 [see also Kharin (2006)].Genus Ephalophis Smith, 1931Ephalophis Smith, M.A. (1931): Proc. Zool. Soc. Lond. 1931: 397-398 [397]. Type species:Ephalophis greyi Smith, 1931 by monotypy.Diagnosis: A monotypic genus of small snakes of the family Ephalophiidae, readily identifiedby the following combination of characters: Head shields large and symmetrical; preocularpresent; valvular nostrils and lingual fossa; 7-8 maxillary teeth following fang; rostral with amedian sharp fold located within the mouth, behind the median and undivided lingual fossa,this fold formed from the posterior edge of the portion of the keratinized scale extending withinthe lining of the lip; preocular large; nasal does not contact preocular; 1 postocular; temporals1+2; supralabials 6; 3rd and 4th supralabials contacting orbit; posterior chin shields large andbordering mental groove separated by small scale; ventrals 151-184; ventrals large, almost asbroad as body, each about 3-5 times the width of adjacent body scales; body scales imbricatewith spine-like to blunt keels posteriorly, but smooth laterally and becoming juxtaposed onventrolateral margins; mid-body scales in 19 or 21 rows; anal divided; subcaudals 24-33entire; tail strongly compressed vertically, but not as high as body; ovoviviparous. Etymology:The name Ephalophis means ‘snake by the sea’, and is derived from the Greek ‘ephalos’ - bythe sea, and ‘ophis’ - snake. Content: Ephalophis greyae Smith, 1931.Ephalophis greyae Smith, 1931Ephalophis greyi Smith, M.A. (1931). Proc. Zool. Soc. Lond. 1931: 397-398 [397, pl. 1]. Typedata: holotype BMNH 1946.1.6.92. Type locality: Cape Boileau, WA.Description: This is a rather diminutive species of seasnake, with a small head that is barelydistinct from the neck, a gracile but solid body, with a tail that is much reduced in its lowerlateral compression height. The body varies from pale brownish to creamish with a series ofdark grey blotch-like bands (about 25 to 40) which extend from the head along the body to thetail; there are around 7-8 rings of dark grey on the tail. The markings are broadest along thedorsum, and often these dark botches may coalesce to form an irregular, wavy zig-zagpattern along the vertebral line. Immature specimens tend to be paler with stronger blackishbanding. Some significant morphological characters of this species are: Head shields largeand symmetrical; rostral with a median sharp fold located within the mouth, behind themedian and undivided lingual fossa, this fold formed from the posterior edge of the portion ofthe keratinized scale extending within the lining of the lip; preocular present; nasal does notcontact preocular; postocular single; temporals 1+2; supralabials 6; 3rd and 4th supralabialscontacting orbit; posterior chin shields large and bordering mental groove separated by smallscale; 7-8 maxillary teeth follow fang; body scales imbricate with spine-like to blunt keelsposteriorly, but smooth and juxtaposed ventro-laterally; mid-body scales in 19 or 21 rows;ventrals about 151-184; ventrals large, almost as broad as body, each about 3-5 times thewidth of adjacent body scales; anal divided; subcaudals 24-33 entire; unlike most other seasnakes, the vertically compressed tail is noticeably down-curved and not as high as the body.Attains a maximum snout-vent length of only around 50-65 cm.Distribution: Known only from the mid-west, and north-west coast of Western Australia, fromKing George Sound on the Kimberley coast, south to about Shark Bay.Habitat: Largely restricted to mangrove flats, river deltas and estuaries in relatively shallowcontinental waters.Biology/Ecology: This is mainly a nocturnal species, although it has been observed activeduring daylight foraging on low-tide mud-flats. The diet is fishes - mainly comprising mud-18

Australian Biodiversity Record, 2007 (8): 1-124skipper fish (Gobiidae) - which are hunted in their burrows on the mud-flats at low tide. Thisspecies is essentially an inhabitant of the littoral and sub-littoral zones and it is very adept atmoving over exposed mud-flats at low tide in marked contrast to most other sea snakes.Toxicity: Although this is a small, largely inoffensive species that rarely attempts to bite undernormal circumstances, caution should nevertheless be exerted, because its venom is likelytoxic to humans. Urgent medical attention should always be sought in the event of a bite fromthis species.Survival Status: Protected under the WA Wildlife Conservation Act 1950 (as amended). Listedas a Marine Protected Species under the Australian Environmental Protection andBiodiversity Conservation Act (1999). Probably not under threat, but its restricted distributionin Australian waters could potentially make it vulnerable in some parts of its range.Etymology: The name 'greyae' honours Australian naturalist Beatrice Grey of Broome whocollected the Type Specimen.Genus Parahydrophis Burger and Natsuno, 1974Parahydrophis Burger, W.L. & Natsuno, T. (1974): Snake 6: 61-75 [p. 65]. Type species:Distira mertoni Roux, 1910 by original designation.Diagnosis: A monotypic genus of small sea snakes of the family Ephalophiidae and readilyidentified by the following combination of characters: head shield enlarged and symmetrical;5-8 maxillary teeth following fang; valvular nostrils and lingual fossa; a small median sharpfold on rostral (within the mouth), which is formed from the posterior edge of the rostral(where it extends within the lining of the lip); large posterior chin shields bordering mentalgroove; preocular present; body scales smooth and imbricate, in 36-39 rows at mid-body;large ventral scales, each being about 3 times the width of the body scales; ventrals 158-160;anal divided; subcaudals 29-35, entire except for first 1 or 2 that are divided; tail stronglycompressed vertically, paddle-like; ovoviviparous. Etymology: The name Parahydrophis (Gk.para - ‘near’, means ‘near Hydrophis’, a belief at the time of its original description‘ [Hydrophis- ‘water snake’, derived from the Greek ‘hydros’ - water, and ‘ophis’ - snake. Content:Parahydrophis mertoni (Roux, 1910).Parahydrophis mertoni (Roux, 1910)Distira mertoni Roux, J. (1910): Abh. Senckenb. Naturforsch. Ges. 33: 210-247 [see p. 222,pl. 13 fig. 4]. Type data: holotype NHMB. Type locality: Sungei Waskei, Wodam, Aru Ils,Indonesia.Description: This is a rather small species of sea snake, with an evenly rounded, robust bodyformthat tends to become more vertically compressed with age. The base body colour variesfrom pale bluish-grey or greenish-grey, to very light brown, or even pale yellow with a patternof distinctive transverse bands or rings across the body. the edges of the pale scales may bedark-edged giving the body a finely dotted appearance. The bands are blackish and numberabout 40 to 50 over the body (with a few on the tail as well), are broadest on the dorsum andgradually taper to the ventro-lateral boundary. Additionally, the banding is irregular in shapeand does not completely encircle the body, and some bands may coalesce at the mid-line(vertebral) to form a darker dorsum than the lateral area of the body; some bands may beoffset at the midline on the anterior of the body to create a sort of limited checked patterndorsally. The width of the dark bands may be either slightly narrower or slightly broader thanthe paler interspaces as well, and some bands may have pale centres. The ventral surface islight brown anteriorly gradually fading to a pale greyish posteriorly. The dorsum of the head isbrownish with fine black variegations, and the snout and side of head is uniform black. Somesignificant morphological characters are: Head shield enlarged and symmetrical; 5-8 maxillaryteeth following fang; a small median sharp fold on rostral (within the mouth), which is formedfrom the posterior edge of the rostral (where it extends within the lining of the lip); largeposterior chin shields bordering mental groove; preocular present; body scales smooth andimbricate, in 36-39 rows at mid-body; large ventral scales, each being about 3 times the width19

Australian Biodiversity Record, 2007 (8): 1-124of the body scales; ventrals 158-160; anal divided; subcaudals 29-35, entire except for first 1or 2 scales that are divided. Content: Parahydrophis mertoni (Roux, 1910). This is a verysmall species of sea snake, attaining a maximum total length of only around 50 cm.Distribution: Confined to the Arafura Basin, ranging from about the Aru Islands in easternIndonesia to north Australian seas, being found locally along the coasts of Queensland (Gulfof Carpentaria) and the Northern Territory, as well as the waters along southern New Guinea.Habitat: Known to live in the turbid waters of river deltas, bays, mud flats, estuaries and gulfson continental shelf areas.Biology/Ecology: This is essentially a nocturnal species that is rarely encountered. It is knownto be ovoviviparous, producing up to 3 living young in a brood. Its diet comprises a variety ofsmall fishes.Toxicity: Although this is a small, inoffensive species that rarely attempts to bite under normalcircumstances, caution should nevertheless be exerted, because its venom is likely toxic tohumans. Urgent medical attention should always be sought in the event of a bite from thisspecies.Survival Status: Protected under the Qld Nature Conservation Act (1992) and the TerritoryParks and Wildlife Conservation Act (1998). Listed as a Marine Protected Species under theAustralian Environmental Protection and Biodiversity Conservation Act (1999). Probably notunder threat, but its restricted distribution in Australian waters could potentially make itvulnerable in some parts of its range.Etymology: The name ‘mertoni’ honours the 19th century European biologist Dr H. Merton.Family HydrelapidaeThis Family only contains the genus Hydrelaps Boulenger, 1896.Genus Hydrelaps Boulenger, 1896Hydrelaps Boulenger, G.A. (1896): Cat. Snakes Brit. Mus. [p. 270]. Type species: Hydrelapsdarwiniensis Boulenger, 1896 by monotypy.Diagnosis: A monotypic genus of sea snakes, readily diagnosed by the following combinationof characters: Head shields large and symmetrical; 3-6 maxillary teeth following fang; valvularnostrils; lingual fossa - part of rostral possessing a median valve-like fold continuous withremainder of scale (median lobe of rostral at its anterior-most end fitting into a median notchin the mental and completely dividing the lingual fossa of the rostral into separate left andright grooves); nasals in broad contact; preoculars absent; postocular single; prefrontalextends laterally to front of eye and so effectively fusing with preocular; 6 supralabials;temporals 1+1, or 1+2, with the primary temporal the largest; posterior chin shields notreduced, separated by a mental groove, and similar in size to the anterior chin shields; bodyscales, smooth and imbricate but tending to be juxtaposed ventrolaterally; mid-body scales in25-29 rows; ventrals about 160-180, narrow and about 2-3 times as wide as adjacent bodyscales; anal divided; subcaudals about 20-36 usually first few scales divided, but mostlyentire; tail paddle-like but less compressed vertically, being of a similar height to the body;ovoviviparous. The fusion of the prefrontals with the preoculars in Hydrelaps is unique to thisgenus. Etymology: The name Hydrelaps means ‘water snake’, and is derived from the Greek,‘hydros’ - ‘water’, and ‘Elaps’, a genus of snakes. Content: Hydrelaps darwiniensis Boulenger,1896.20

Australian Biodiversity Record, 2007 (8): 1-124Hydrelaps darwiniensis Boulenger, 1896Hydrelaps darwiniensis Boulenger, G.A. (1896): Cat. Snakes Brit. Mus. [p. 270, pl. 12 fig. 1].Type data: syntypes BMNH 1946.1.1.91-92. Type locality: Port Darwin, NT.Description: This is another diminutive species of sea snake of the sub-littoral tidal zone. Ithas a slender, and less compressed body than other seasnake species, a small head that isbarely distinct from the neck, and a tail that is similar in profile height to the body. The head ismostly black with scattered cream to yellow variegations. The base body colour is palecreamish to yellow with a series of 30-45 broad dark rings that encircle the body, and a further5-8 dark rings on the tail. These transverse body bands are each about twice as wide as thepale interspaces, and in some specimens, some of the bands may be broken at the vertebral.Ventrally, same as dorsum as the most rings tend to encircle the body, although some breakat the venter also; the subcaudals are usually black. Some significant features of this species’morphology are: Head shields large and symmetrical; nasals in broad contact; 3-6 maxillaryteeth following fang; 6 supralabials; 3rd and 4th supralabials contacting orbit; preocularsabsent; postocular single; prefrontal extends laterally to front of eye and so effectively fusingwith preocular; rostral with median lobe at its anterior-most end, this lobe fitting into a mediannotch in the mental and completely dividing the lingual fossa of the rostral into separate leftand right grooves; temporals 1+1, or 1+2, with the primary temporal the largest; posterior chinshields not reduced, separated by a mental groove, and similar in size to the anterior chinshields; body scales, smooth and imbricate but tending to be juxtaposed ventrolaterally; midbodyscales in 25-29 rows; ventrals about 160-180, narrow and about 2-3 times as wide asadjacent body scales; anal divided; subcaudals about 20-36 mostly entire; tail lesscompressed vertically, than other sea snakes being of a similar height to the body. Attains amaximum total length of only around 50-60 cm.Distribution: Known mainly from tropical Australian waters west of the Torres Strait. It occurspatchily along the coasts of north-west Queensland (mainly in the Gulf of Carpentaria),Northern Territory and Western Australia, to about as far south as Dampier. Also known fromsouthern areas of coastal Papua New Guinea and southern Irian Jaya.Habitat: Largely restricted to mangrove flats and freshwater river deltas and estuaries inrelatively shallow continental waters.Biology/Ecology: Essentially a nocturnal species that is believed to feed on a variety of smallfishes, such as Mudskippers. It may also be found active on mudflats during the day onincoming tides, and has been observed crawling over exposed mudflats, and activelysearching crab burrows - presumably for fish. Produces living young, but reproductionotherwise unknown. Sharks are known predators of this species.Toxicity: Although this is a small, largely inoffensive species that rarely attempts to bite undernormal circumstances, caution should nevertheless be exerted, because its venom is likelytoxic to humans. Urgent medical attention should always be sought in the event of a bite fromthis species.Survival Status: Protected under the Qld Nature Conservation Act (1992), the Territory Parksand Wildlife Conservation Act (1998), and the WA Wildlife Conservation Act 1950 (asamended). Listed as a Marine Protected Species under the Australian EnvironmentalProtection and Biodiversity Conservation Act (1999). Probably not under threat, but itsrestricted distribution in Australian waters could potentially make it vulnerable in some parts ofits range.Etymology: The name 'darwiniensis' refers to the Type Locality for the species, Darwin, NT.21

Australian Biodiversity Record, 2007 (8): 1-124Family HydrophiidaeGenus Acalyptophis Boulenger, 1896Acalyptus Duméril, A.M.C. (1853): Mém. Acad. Sci. Inst. Fr., 23: 399-536 [p. 522; nonAcalyptus Schonheer, 1836]. Type species: Acalyptus peronii Duméril, 1853 by monotypy.Acalyptophis Boulenger, G.A. (1896): Cat. Snakes Brit. Mus., 3 [p. 269] [nom. substit. proAcalyptus Duméril, 1853].Pseudodistira Kinghorn, J.R. (1926): Proc. Zool. Soc. Lond. 1926: 71-74 [p. 71]. Typespecies: Pseudodistira horrida Kinghorn, 1926 by monotypy.Diagnosis: A monotypic genus of marine snakes of the family Hydrophiidae and readilyseparated from all other genera by the following combination of characters: Form small butrobust-bodied; head small and indistinct from the neck; head shields fragmented and irregularor asymmetrical in shape, with most scales on the head of mature specimens having a raisedprojection or spine on the posterior edge (particularly prominent on the supraoculars andpost-oculars); parietals and frontal fragmented; no prefrontal scales; nasal contacts preocular;body scales with a single keel, imbricate anteriorly, but nearly juxtaposed on the posterior partof the body and in 21-31 rows at mid-body; ventrals entire, very small 140-210 (each about aswide as the adjacent body scales); anal divided; valvular nostrils and lingual fossa (tonguenotch); tail strongly compressed vertically, paddle-like; ovoviviparous. Etymology: The nameAcalyptophis is derived from the Greek acalyptos which means uncovered, and ophis forsnake - and is possibly in reference to the almost uniform ventral scalation - instead of thebroad ventral scales of other species. Content: Acalyptophis peronii (Dumeril, 1853).Acalyptophis peronii (Dumeril, 1853)Acalyptus peronii Duméril, A.M.C. (1853): Mém. Acad. Sci. Inst. Fr. 23: 399-536 [p. 522].Type data: holotype MNHP 7711. Type locality: ?New Holland.Acalyptus superciliosus velperoni Duméril, A.M.C., Bibron, G. and Duméril, A. (1854): Erpét.Gén. Hist. Nat. Comp. Reptiles [p. 1340 - as vel peroni]. Type data: holotype MNHP 7711.Type locality: Australia.Pseudodistira horrida Kinghorn, J.R. (1926): Proc. Zool. Soc. Lond. 1926: 71-74 [p. 71]. Typedata: holotype AM R8640. Type locality: McCulloch Reef, Great Barrier Reef, QLD [17º20'S146º30'E].Description: A medium-sized species of sea snake in which the head is relatively small butdeep and indistinct from the neck; the anterior of the body is quite slender, and the posteriorthick-set, with the lateral compression of the tail extending slightly higher than the body depth.The base body colour varies from light brown to creamish or greyish, with the head noticeablypaler than the body (at any age), and the lower lateral intermittently brighter creamish tocreamish-yellow. The body pattern comprises a series of broad transverse very dark brown(almost black) bands that are widest on the dorsum and each is about as wide as the palerinterspaces; there may also be a series of much narrower and somewhat fainter brownishbands within the pale interspaces as well - although this is not always the case. The principalbroad bands number about 25-30, and slightly taper towards the lower lateral area, wherethey may form a point ventro-laterally. These bands are usually more distinct on the anteriorbody and rather less prominent on the tail, and the bands are usually quite prominent inmature individuals, but rather more obscure on older individuals. The ventral colour is pale -almost white, without any markings. The head shields are fragmented and irregular in shape,with most scales on the head possessing either prominent tubercles (juveniles) or a raisedprojection or spine (in adults) on the posterior edges of the scales. These spines areparticularly large on the supraoculars and post-oculars giving the head a distinctive hornedappearance. Some other significant features of this species’ morphology are: body scaleskeeled, imbricate anteriorly, but nearly juxtaposed on the posterior part of the body and in 21-31 rows at mid-body; ventrals 140-210 entire and each very small (only about as wide as the22

Australian Biodiversity Record, 2007 (8): 1-124adjacent body scales); anal divided; nasal contacts the single preocular; parietals and frontalfragmented; no prefrontal scales. There are 5-8 solid maxillary teeth behind the fang. Attainsa maximum total length of around 1.2 m., and a snout-vent length of about 1 m.Distribution: Although this species is virtually endemic to the Australian region, includingsouthern New Guinea, it also occurs in Indonesia and the south-west Pacific to about NewCaledonia. Within Australian territory, it is largely confined to tropical seas off the coasts ofQueensland, Northern Territory and the north-west of Western Australia (to about as far southas Barrow Island). Occasionally specimens may be found along the subtropical far north eastcoast of New South Wales as well, but no permanent population occurs in this State (notealso that this species was not mentioned as part of the NSW fauna in the most recent guide tothat region (Swan, Shea and Sadlier, 2004).Habitat: It mainly inhabits sand beds of coral reefs in relatively shallow (about 4 to 20 metresdeep) continental shelf waters. It appears to be more common around the outer edges ofreefs.Biology/Ecology: This is essentially a nektonic, nocturnal predator of several species of smallfishes, particularly Gobies, which it either actively hunts around coral, or by searching burrowsand crevices. It can also be observed active in shallow reef waters during daylight as well -and at any time of the year. It is ovoviviparous, producing up to 10 living young in a brood,although the average is only around 4, with larger females tending to have larger litters. In theTorres Strait, a gravid female has been recorded in November (Summer/early Wet Season),while one in Asia (South China Sea) was gravid in April - weather conditions (early MonsoonSeason). On occasions specimens of this species are accidentally captured during fishingoperations in northern Australia.Toxicity: In general this species is not overtly aggressive when disturbed, and indeed willusually refrain from biting unless provoked by very rough treatment or handling duringcapture. Caution should nevertheless be exerted, because its venom is known to be highlytoxic. Although there are no recorded human fatalities from this species, urgent medicalattention should always be sought in the event of a bite.Survival Status: Protected under the Qld Nature Conservation Act (1992), the Territory Parksand Wildlife Conservation Act (1998) and the WA Wildlife Conservation Act 1950 (asamended). Protected under the New South Wales National Parks and Wildlife Act (1974) butnot listed in that State as a Threatened Species in any of the Schedules of the NSWThreatened Species Conservation Act (1995). Listed as a Marine Protected Species underthe Australian Environmental Protection and Biodiversity Conservation Act (1999). It isprobably not under threat, but its restricted distribution in Australian waters could potentiallymake it vulnerable in some parts of its range.Etymology: The name ‘peronii’ honours the French naturalist Francois Peron (1775-1810).Genus Astrotia Fischer, 1856Astrotia Fischer, J.G. (1856): Abh. Geb. Naturw. Hamburg 3: 1-78 [p. 38]. Type species:Hydrophis schizopholis Schmidt, 1846 by monotypy.Diagnosis: As presently defined a monotypic genus of extremely large marine snakes of thefamily Hydrophiidae, readily separated from all other genera by the following combination ofcharacters: Head long, deep, and squarish (or weakly rounded) in shape when viewed fromabove, (but with the snout slightly or steeply angular in profile), and barely distinct from neckor not at all; head shields large and symmetrical, with each shield much larger than adjacentnape scales; valvular nostrils and lingual fossa; nasals in contact; prefrontals present; 1preocular; 2 postoculars; 2 primary temporals; supralabials 10-11 (occasionally divided);mental shield triangular in shape, wider than long; distinct but shallow mental groove; mentalshield not partially hidden in the mental groove; body scales relatively smooth, stronglyimbricate, each with either a central week keel or small line of tubercles, and in 46-63 rows atmid-body; ventrals longitudinally divided (i.e. divided into pairs of foliform scales), strongly23

Australian Biodiversity Record, 2007 (8): 1-124imbricate, and numbering 226-286, each being only slightly wider than the adjacent bodyscales; ventrals in mature specimens (except on throat) form a distinct mid-line keel at thepoint of overlap of each ventral scale; tail strongly compressed vertically, paddle-like; preanalscales enlarged; ovoviviparous. Etymology: The name Astrotia is derived from ‘astrotos’(Greek) meaning ‘not covered’, and is believed to refer to the lack of broad ventral scales.Content: Astrotia stokesii (Gray, 1846).Astrotia stokesii (Gray, 1846)Hydrus stokesii Gray, J.E. (1846): Disc. Aust. Voy. Beagle…[p. 502, pl. 3]. Type data:holotype BMNH 1946.1.17.12. Type locality: Australian Seas.Hydrophis schizopholis Schmidt, P. (1846): Abh. Geb. Naturw. Hamburg 1: 164-172 [166, pl.15 figs 1-7]. Type data: holotype ZMH 387. Type locality: China Sea.Hydrus annulatus Gray, J.E. (1849). Cat. Spec. Snakes Coll. Brit. Mus., [p. 59]. Type data:holotype BMNH 1946.1.19.74. Type locality: Singapore.Hydrophis guntheri Theobald, W. (1868): Bull. Mus. R. Hist. Nat. Belg. 37: 67-74 [p. 69]. Typedata: presumed lost. Type locality: mouth of Hooghly River, India (as probably from theSandheads).Hydrophis granosa Anderson, J. (1871): Proc. Zool. Soc. Lond. 1871: 149-211 [p. 190]. Typedata: holotype IM. Type locality: mouth of Hooghly River, India (as Sand Heads).Hydrophis guttata Murray, J.A. (1887): J. Bombay Nat. Hist. Soc. 2: 32-35 [p. 33]. Type data:holotype BMNH 1946.1.19.73. Type locality: Makran coast, Iran and Pakistan (as MekranCoast).Description: Although there are longer species, this is usually a very large (if not the largest)species of sea snake, with a massive girth when mature, giving the body-form a very robustappearance. The head is usually not distinct from the thick neck, being long, deep, andsquarish (or weakly rounded) in shape when viewed from above, but with the snout slightly orsteeply angular in profile. There appears to be differences in head shape between the sexes,as well as some geographic variation. Overall the base body colour of mature specimens isusually creamish, yellowish-cream, pale brown or creamy-white, but occasionally muchdarker, more greyish to almost black specimens are found, with the head either creamishbrown,darker brown or black. In regards to dorsal pattern, specimens are sometimes withoutany pattern or with just an obscure reticulum of darker blotching along the lower lateral andventer. The largest (oldest?) specimens are often without any discernable pattern on auniform pale body. However, patterned specimens are the norm in any given population.There is usually a vertebral series of broad, dark leaden-grey or black transverse blotches,which are broadest along the mid-line and narrow to a point mid-laterally. Additionally, there isusually a smaller transverse blotch or group of blotches along the vertebral line between eachlarge blotch, and below this another smaller blotch on the upper lateral of the body. Thegeneral effect of this pattern is that of a series of broken transverse bands. In juveniles, thispattern is similar, but much more intense, with the base colour being almost white and thepattern being glossy black, with each large vertebral blotch in the shape of a sharply defineddiamond across the back, and so resulting in a prominent zig-zag pattern along the lateralzone. Occasionally, these dark blotches may coalesce along the vertebral and form a semicontinuousdarker dorsum. The smaller blotches between the larger markings are oftenrestricted to a single upper lateral spot or small blotch, but these markings are usually in aneat longitudinal alignment along the body. The ventral or lower body pattern usuallycomprises two series of small blotches in an alternating pattern, one along the mid-ventral lineand the other at the ventrolateral margin, but this pattern tends to be less distinct with age.Some significant features of this species' morphology are: head shields large andsymmetrical, with each shield much larger than adjacent nape scales; prefrontals present;nasals in contact; 1 preocular; 2 postoculars; 2 primary temporals; supralabials 10-11(occasionally divided); mental shield triangular in shape, wider than long, and a distinctmental groove; body scales relatively smooth, strongly imbricate, each with either a central24

Australian Biodiversity Record, 2007 (8): 1-124week keel or small line of tubercles, and in 46-63 rows at mid-body (with females having aslightly higher mid-body count than males); ventrals longitudinally divided, strongly imbricate,and numbering 226-286 (similar in males and females), each being only slightly wider thanthe adjacent body scales, and in mature specimens forming a distinct mid-line keel at thepoint of overlap of each ventral scale; preanal scales enlarged. It is said to attain a maximumtotal length of about 2 m., although 1.5 m. would be the usual maximum size - and femalestend to be somewhat larger than males, and have longer snout vent lengths. In Australianwaters a maximum snout-vent length of around 1.4 m. is achieved, with females being largerthan males which are seldom much over 1.2m. This is nevertheless an extremely largespecies, evidenced by an exceptional specimen 1.6 m. in total length that had a girth of 260mm. - making it probably the bulkiest of sea snakes. Variation in morphology suggests thatthis species may be composite.Distribution: Largely confined to tropical Australian seas, being found off the coasts ofQueensland, Northern Territory and Western Australia, and occasionally as far south as thewaters off Wollongong, New South Wales. Also widely known from the Arabian Gulf, acrossSouth East Asia, mainly from Pakistan, through Indonesia to New Guinea.Habitat: Mainly inhabits relatively shallow continental shelf areas near coral reefs with sandybottoms as well as around muddy river delta outflows and estuarine areas usually in watersfrom about 4m to 25m depth - but usually less than 10 metres.Biology/Ecology: This is both a diurnal and nocturnal species that feeds on a variety of smallfishes, with prey being actively hunted around coral. It may be often observed basking at thesurface in relatively turbid waters near rivers and is known to inhabit tidal waters somedistance up rivers as well. Specimens are occasionally found with growths of algae, barnaclesand bryozoans on its skin, and large infestations of nematodes may be carried in thestomach. It appears to have seasonal reproductive habits, but only limited data is presentlyavailable. Females carrying oviducal young have been found in mid-summer, and it ispresumed that the young are born in late Autumn. Litter sizes vary from 4 to 20, with anaverage of 10 to 14 being usual - juveniles are around 400 mm in length at birth.Toxicity: This is a potentially dangerous species as its venom is highly toxic to humans and itwill readily attempt to bite if handled or harassed. Caution should definitely be exerted withthis species, because on occasion it has been known to attack divers with little warning orprovocation. With its relatively large teeth and a large amount of venom, a wet-suit should notbe considered as adequate protection from penetration by the fangs of an adult snake.Similarly, specimens landed on trawlers vigorously bite anything within range, includingfishing gear, and those handled by researchers readily try to bite if restrained or handledroughly. If one is unfortunate enough to receive a bite from this powerful species, urgentmedical attention should be sought - even though to date there have been no known humanfatalities recorded against this species.Survival Status: Protected under the New South Wales National Parks and Wildlife Act (1974)but not listed in that State as a Threatened Species in any of the Schedules of the NSWThreatened Species Conservation Act (1995). Also protected under the Qld NatureConservation Act (1992), the Territory Parks and Wildlife Conservation Act (1998) and theWA Wildlife Conservation Act 1950 (as amended). Listed as a Marine Protected Speciesunder the Australian Environmental Protection and Biodiversity Conservation Act (1999).Probably not under threat, but its restricted distribution in Australian waters could potentiallymake it vulnerable in some parts of its range.Etymology: The name 'stokesii' honours English explorer and naturalist Admiral John LortStokes, who collected the species in 1840.25

Australian Biodiversity Record, 2007 (8): 1-124Genus Chitulia Gray, 1849Asturia Gray, J.E. (1842): In Gray, J.E. (ed.) Zoological Miscellany. [p. 55] [nomen nudum;date of publication, Apr. 1842].Aturia Gray, J.E. (1842): Monogr. Synop. Water Snakes…Hydridae. [p. 61] [date ofpublication, May 1842]. [preoccupied] Type species: Aturia ornata Gray, 1842 by subsequentdesignation [see Smith, M.A. (1926). Monogr. Sea Snakes]Chitulia Gray, J.E. (1849): Cat. Spec. Snakes Coll. Brit. Mus., [p. 36, 56]. Type species:Chitulia inornata Gray, 1849 by subsequent designation [see Smith, M.A. (1926). Monogr.Sea Snakes]Micromastophis Wall, F. (1921). Ophidia Taprobanica or the Snakes of Ceylon. [errata insert][nom. substit. pro Aturia Wall, 1921] [p. 344;? non Aturia Gray, 1842]. Type species: Hydrusfasciatus Schneider, 1799 (= Hydrus fasciatus Schneider, 1799) by monotypy.Diagnosis: A genus of moderate-sized snakes of the Family Hydrophiidae, readily diagnosedby the following combination of characters: head shields enlarged, symmetrical; width of headbetween the eyes less than half the distance from tip of snout to rear edge of parietals; nasalseparated from preocular by high 2nd supralabial; one or two supralabials contacting orbit;valvular nostrils and lingual fossa; mental groove present and distinct; mental triangular,broader than long, and not partially hidden in the shallow mental groove; anterior chin scaleslarge, and usually bordering mental groove; anterior part of maxilla not arched upwards, thetip of the fang projecting conspicuously below a line connecting the tips of the solidmaxillaries; less than 15 maxillary teeth following fang; body scales imbricate; small ventrals,about as wide as, or only slightly wider than adjacent body scales and never foliform;posterior ventrals distinct, mostly undivided; ventrals 195-340; tail strongly compressedvertically, paddle-like; small heart just behind anterior third of body; ovoviviparous. Etymology:At the time of the original description Chitulia was a village in British India (now Bangladesh).Content: Chitulia belcheri (Gray 1849); Chitulia inornata Gray 1849; Chitulia ornata (Gray1842); Chitulia ocellata (Gray, 1849); Chitulia sibauensis comb. nov. (Rasmussen, Auliya andBohme, 2001); Chitulia stricticollis Gunther 1864; and, Chitulia torquata Gunther 1864.Content in Australia: Chitulia belcheri (Gray, 1849); Chitulia inornata Gray, 1849; Chituliaocellata (Gray, 1849) and supposedly Chitulia ornata (Gray, 1842).Chitulia belcheri (Gray, 1849)Aturia belcheri Gray, J.E. (1849): Cat. Spec. Snakes Coll. Brit. Mus., [p. 46]. Type data:holotype BMNH 1946.1.1.97. Type locality: New Guinea.Hydrophis pachycercos Fischer, J.G. (1856): Abh. Geb. Naturw. Hamburg 3: 1-78 [p. 44, pl.2]. Type data: holotype ZMH 390. Type locality: Indian Seas.Hydrophis pachycerios Jan, G. (1859). Rev. Mag. Zool. (2)10 and (2)11-12, 1858 and 1859.[p. 24] [lapsus for Hydrophis pachycercos Fischer, 1856]Description: This is a moderate-sized species with an elongate body and small narrow head.The base body colour is pale to dark grey on the dorsum of the body. There is a series ofabout 60 blackish transverse bands on the body. The bands are widest at the vertebral, butgradually narrow on the lateral of the body, and fade to dark grey at the ventro-lateral zone.The venter is a paler grey than the dorsal base colour. Some significant features of thisspecies’ morphology are: head shields enlarged, symmetrical; anterior part of maxilla notarched upwards, the tip of the fang projecting conspicuously below a line connecting the tipsof the solid maxillaries; less than 10 maxillary teeth following fang; nasal separated frompreocular by high 2nd supralabial; usually only one supralabial contacting orbit; mental groovepresent and distinct; mental triangular, broader than long, and not partially hidden in theshallow mental groove; anterior chin scales large, and usually bordering mental groove; midbodyscales in 34 rows; small ventrals, about as wide as, or only slightly wider than adjacent26

Australian Biodiversity Record, 2007 (8): 1-124body scales and never foliform; posterior ventrals distinct, mostly undivided; ventrals 300-320.Attains a maximum total length of only around 1 m.Distribution: This sea snake has been frequently cited as part of the Australian faunahowever, it is actually an Asian species that occurs east as far as the island of New Guinea.Previous records of this species in Australian waters were actually based on specimens of asuperficially similar species that is now known as Leioselasma coggeri Kharin, 1984. Despitethe lack of firm specimen-based records for Chitulia belcheri in Australian waters, and the factthat resident populations are almost certainly confined to tropical seas around islands andreefs off the north-west coast of New Guinea, it may reach Australian territory on occasions.Its relative close proximity geographically, and the prevailing currents suggest that it couldpossibly occur in seas off northern Western Australia and to the north-west of the NorthernTerritory, so it is included here as tentatively part of our Hydrophiid fauna as well.Habitat: It inhabits reefs in relatively shallow continental waters around New Guinea, butelsewhere uncertain.Biology/Ecology: This is essentially a diurnal, fast-swimming species that feeds on a variety ofsmall fishes, such as eels. Reproductive biology unknown, but like other Hydrophiids it isovoviviparous, with a known litter of 4.Toxicity: It rarely attempts to bite under normal circumstances, but caution shouldnevertheless be exerted, because its venom is highly toxic to humans. Urgent medicalattention should always be sought in the event of a bite from this species.Survival Status: Protected under the Territory Parks and Wildlife Conservation Act (1998) andthe WA Wildlife Conservation Act 1950 (as amended). Listed as a Marine Protected Speciesunder the Australian Environmental Protection and Biodiversity Conservation Act (1999).Etymology: Named for Captain Sir Edward Belcher (1799-1877), commander of the shipcalled Sulphur, during the years 1836-1842. [Belcher was ordered to start his voyage at thePacific coast of South America and follow the Trans Pacific route to England. During thevoyage Belcher visited several islands such as the Solomon Islands, the Hawaiian Islands,the Society Islands, the Tonga Islands, the New Hebrides, and New Guinea - See Belcher, E.(1843). Narrative of a Voyage round the World, performed in Her Majesty's Ship Sulphur,during the years 1836-1842, including Details of the Naval Operations in China, from Dec.1840 to Nov. 1841. London, Henry Colburn, 1843. 2 volumes].Chitulia inornata Gray, 1849Chitulia fasciata Gray, J.E. (1849): Cat. Spec. Snakes Coll. Brit. Mus., [p. 56]. Type data:holotype BMNH 1946.1.1.59. Type locality: Indian Ocean.Chitulia inornata Gray, J.E. (1849): Cat. Spec. Snakes Coll. Brit. Mus., [p. 56]. Type data:holotype BMNH 1946.1.1.27. Type locality: Indian Ocean.Thalassophis schlegelii Schmidt, P. (1852): Beiträge zur ferneren kentniss derMeerschlangen. Abh. Geb. Naturw. Hamburg 2: 69-86 [p. 83, pl. 5]. Type data: syntypes ZMH393-395. Type locality: Java, Indonesia.Hydrophis manillae Owen, R. (1859): Descr. Cat. Spec. Nat. Hist. Mus. Roy. Coll. Surg.Vertebrata. [p. 77] [nom. nud.].Hydrophis longiceps Günther, A. (1864): Reptiles of British India. [p. 375, pl. 25 fig. 0][unnecessary replacement name for Chitulia fasciata Gray, 1849].Description: This is a rather small or diminutive species with a relatively slender and ratheruniform body-shape. Unlike many other adult sea snakes this species generally lacks anydistinctive dark blotching or banding. As its name suggests it is rather plain in pattern, being abasic pale bluish-grey on the dorsum with either no pattern at all, or if present, it is only a faint27

Australian Biodiversity Record, 2007 (8): 1-124darker blotching along the body. The venter is uniform whitish. Juveniles or immaturespecimens however, usually possess from about 50 to 65 black band-like bars over thedorsum - or even as complete rings around the body. Some significant features of thisspecies’ morphology are: head shields enlarged, symmetrical; width of head between theeyes less than half the distance from tip of snout to rear edge of parietals; 10-13 solidmaxillary teeth following fang; dentary teeth usually fewer than 23; 3rd and 4th supralabialscontacting orbit; mental groove present and distinct; mental triangular, broader than long, andnot partially hidden in the shallow mental groove; anterior chin scales large, and usuallybordering mental groove; nasal separated from preocular by high 2nd supralabial; bodyscales keeled, or with a slight tubercle, and imbricate in 35-48 rows at mid-body; smallventrals, each about as wide as, or only slightly wider than adjacent body scales and neverfoliform; ventrals about 195-295, those posterior distinct, mostly undivided; heart just behindanterior third of body. This species reaches a maximum total length of only around 70cm.Distribution: Widely distributed across South East Asia from the Philippines to Australasia, butseldom found in tropical Australian seas, being only known from the waters off the NorthernTerritory (Arafura Sea) and possibly Western Australia. Specimens have been rarely trawledin the south-eastern Gulf of Carpentaria, Queensland and stray specimens may occur as farsouth as the central coast in New South Wales.Habitat: More often found in turbid waters of river deltas, bays and gulfs on continental shelfareas. On occasions it may be found inhabiting tidal rivers, some kilometres upstream fromthe sea.Biology/Ecology: Very little is known of the biology and ecology of this apparently uncommonspecies. It is essentially diurnal and fast-swimming in habits, feeding on a variety of smallfishes which it captures in fairly shallow waters. It is known that 3 young have been producedin a brood, but it is believed that a litter size of around 8 may occur in this species.Toxicity: It rarely attempts to bite under normal circumstances, but caution shouldnevertheless be exerted, because its venom is highly toxic to humans. Urgent medicalattention should always be sought in the event of a bite from this species.Survival Status: Protected under the Territory Parks and Wildlife Conservation Act (1998), theWA Wildlife Conservation Act 1950 (as amended) and the Qld Nature Conservation Act(1992). Also protected under the New South Wales National Parks and Wildlife Act (1974) butnot listed in that State as a Threatened Species in any of the Schedules of the NSWThreatened Species Conservation Act (1995). Listed as a Marine Protected Species underthe Australian Environmental Protection and Biodiversity Conservation Act (1999). Probablynot under threat, but its restricted distribution in Australian waters could potentially make itvulnerable in some parts of its range.Etymology: The name 'inornata' means 'unadorned' and refers to the lack of a body pattern ofthe speciesChitulia ornata (Gray, 1842)Aturia ornata Gray, J.E. (1842): Monogr. Synop. Hydridae. Zoological Miscellany [p. 61]. Typedata: holotype BMNH 1946.1.23.72. Type locality: unknown.Hydrophis ocellata Gray, J.E. (1849): Cat. Spec. Snakes Coll. Brit. Mus., [p. 53]. Type data:holotype BMNH 1946.1.3.91. Type locality: Australia.Hydrophis laevis Lütken, C. (1863): Vidensk. Medd. Dan. Naturhist. Foren. 1862: 292-331[309, pl. 1, pl. 2 fig. 6]. Type data: syntypes (probable) whereabouts unknown. Type locality:easterly part of the globe.Hydrophis ellioti Günther, A. (1864): Reptiles Brit. India [p. 377, pl. 25 fig. n]. Type data:syntypes BMNH 1946.1.4.85, BMNH 1946.1.6.85, BMNH 1946.1.6.84. Type locality: Madras,India, Ceylon [Sri Lanka], provenance unknown.28

Australian Biodiversity Record, 2007 (8): 1-124Hydrophis godeffroyi Peters, W. (1872): Mber. K. Preuss. Akad. Wiss. Berl. 1872: 848-861 [p.856, pl. 1 fig. 3]. Type data: syntypes ZMB 7593 2 specimens, ZMB 28399 2 specimens. Typelocality: Gilbert Ils (as Kings-Mills Ils) [Kiribati].Distira andamanica Annandale, N. (1905): Bull. Mus. R. Hist. Nat. Belg. (ns)1: 173-176 [p.174]. Type data: holotype IM 15238 Type locality: Andaman Ils, India.Distira mjobergi Lönnberg, E. and Andersson, L.G. (1913). K. Sven. Vetensk.-Akad. Handl.52(3): 1-173 [p. 13]. Type data: holotype NHRM 2402. Type locality: Cape Jaubert, WA.Hydrophis lamberti Smith, M.A. (1917): J. Nat. Hist. Soc. Siam 2: 340-342 [p. 340]. Type data:holotype BMNH 1946.1.9.20. Type locality: mouth of the Mekong (as Meklong) River, Gulf ofSiam, Vietnam.Description: This species has a robust body-form along most of its length, being fairly uniformboth anteriorly and posteriorly (and only moderately compressed posteriorly). The head isrelatively large, deep and not distinct from the neck, the width of head between the eyesbeing at least half the distance from tip of snout to rear edge of parietals. The base bodycolour in mature specimens is usually bluish-grey or greyish on the dorsum and whitish-creamon the lower lateral area and venter. Pattern is usually confined to a series of 30-60 broadblackish bars or transverse blotches over the dorsum, below which there may be a series ofsmall dark blotches and dark-edged ocelli. The 'bands' may be absent or at best very obscurein aged individuals, with the upper body usually being uniform bluish-grey. Some significantfeatures of this species' morphology are: head shields large and symmetrical; 10-14 solidmaxillary teeth following fang; dentary teeth usually fewer than 23; heart just behind anteriorthird of body; supralabials 6-8; 3rd and 4th supralabials contacting orbit; nasal separated frompreocular by high 2nd supralabial; 1 preocular; 2-3 (usually 2) postoculars; 1-2 primarytemporals (occasionally 1 large secondary temporal); mental groove present and distinct;mental triangular, broader than long, and not partially hidden in the shallow mental groove;anterior chin shields large and usually bordering the mental groove; body scales smooth andimbricate, but juxtaposed dorsally with a short keel, and in 42-62 rows at mid-body (Australianspecimens tend to have higher mid-body scale counts than those from Asia - 45-62 vs 42-54);body scales 34-41 rows at neck (Australian specimens tend to have slightly higher neck scalecounts than those from Asia - 34-41 vs 31-44); ventrals small, with two short keels, eachventral about twice as wide as adjacent body scales and never foliform; ventrals 235-338,with those on the posterior body being entire (Australian specimens tend to have higherventral counts than those from Asia - 278-338 vs 235-294). Variation in morphology suggeststhat this species may be composite. Attains a mature snout-vent length of only around 85 cm.up to maximum of about 1m., however, on occasions specimens are found measuring around1.6 m. in total length. Males attain a larger size than females, and females are sexuallymature at around 80cm.Distribution: Widely distributed across South East Asia, from the Persian Gulf to India, SriLanka, Myanmar, Thailand, Malaysia, Vietnam, China, Japan, Philippines and Indonesia andacross to Australasia (including New Guinea) and the south west Pacific (Kiribati). In ourregion, it is known mainly from tropical and sub-tropical seas, being found off the coasts ofQueensland, Northern Territory and Western Australia to about Exmouth (but also as farsouth as Geographe Bay, WA as vagrants), and on occasions off New South Wales, Victoriaand even Tasmania as well (as vagrants). However, the Australian population is unique andendemic to this region, and I consider that it should be separately recognized as a differentspecies - see the use of Chitulia ocellata (Gray, 1849) in the taxonomic footnote below.Habitat: Known to live around coral reefs, as well as the turbid waters of river deltas, bays,estuaries and gulfs on continental shelf areas in depths ranging from around 4m. to around22m, although it is known from relatively deeper waters - sometimes exceeding 30 metres.Biology/Ecology: Essentially a diurnal species, its diet consists of a variety of small fishes.Gravid females have been found in Asia during the Dry Season (March-April), and it ispresumed that the young are born before around the early Wet or Summer Monsoon Season.29

Australian Biodiversity Record, 2007 (8): 1-124It produces anywhere from 1 to 17 young in a brood (with averages of about 6 in Australia,and only about 3 in Asia). Interestingly, larger specimens tend to have larger numbers ofyoung in the Australian part of the range, but in Asia the number of offspring is not related tofemale size.Toxicity: This is a fast swimming species that rarely attempts to bite under normalcircumstances, but caution should nevertheless be exerted, because its venom is highly toxicto humans. There have been a number of fatalities recorded from the bite of this species, sourgent medical attention should always be sought in the event of an envenomation. See forinstance Tamiya, Maeda, and Cogger (1983).Survival Status: Protected under the New South Wales National Parks and Wildlife Act (1974)but not listed in that State as a Threatened Species in any of the Schedules of the NSWThreatened Species Conservation Act (1995). Also protected under the Victorian Wildlife Act(1975) but not listed as threatened in the Victorian Flora and Fauna Guarantee Act (1988)],the Qld Nature Conservation Act (1992), the Territory Parks and Wildlife Conservation Act(1998), the Tasmanian National Parks and Wildlife Act (1970) and the WA WildlifeConservation Act 1950 (as amended). Listed as a Marine Protected Species under theAustralian Environmental Protection and Biodiversity Conservation Act (1999). Probably notunder threat, but its restricted distribution in Australian waters and the huge numbers of thisspecies killed as bycatch in fishing and prawn trawling operations could potentially make itvulnerable in some parts of its range.Etymology: The name 'ornata' means 'adorned' and refers to the body pattern of the species.The name ‘ocellata’ refers to the ocellate pattern along the lower lateral of the body in youngspecimens.Final Taxonomic Note: Storr, Smith and Johnstone (1986, 2002) used the name Hydrophisocellatus for this species, but this has generally been ignored. However, in my opinion theAustralian population long regarded as Hydrophis ornatus should be hereafter referred to thespecies called Hydrophis ocellata by Gray, in 1849. Consequently, this necessitates the useof the new combination of Chitulia ocellata (Gray, 1849) for the population in Australia.Genus Disteira Lacepede, 1804Disteira Lacépède, B.G.E. (1804) : Ann. Mus. Natl Hist. Nat. Paris 4: 184-211 [p. 210]. Typespecies: Disteira doliata Lacépède, 1804 by monotypy.Distira Boulenger, G.A. (1896): Cat. Snakes Brit. Mus. [p. 285] [invalid emend. pro DisteiraLacépède, 1804].Melanomystax Wall, F. (1921): Ophidia Taprobanica or the Snakes of Ceylon. [p. 381]. Typespecies: Hydrophis nigrocinctus Daudin, 1803 by monotypy.Diagnosis: A genus of marine snakes with four described species (of which two occur inAustralian waters) in the Family Hydrophiidae and readily separated from all other genera bythe following combination of characters: Head small, body form elongate to moderatelyrobust; head shields enlarged and symmetrical; less than 10 maxillary teeth following fang;valvular nostrils and lingual fossa; mental shield triangular, broader than long; mental groovepresent and shallow, but distinct; mental shield not partially hidden in the mental groove; bodyscales imbricate and keeled dorsally, but smooth laterally; ventrals small and only slightlylarger than body scales; ventrals mostly undivided, never as foliform scales and never forminga mid-ventral keel; tail strongly compressed vertically, paddle-like; prefrontal in contact withsecond supralabial; ovoviviparous. Content: Disteira kingii (Boulenger, 1896); Disteira major(Shaw, 1802); Disteira nigrocincta (Daudin, 1803); Disteira walli Kharin, 1989; Content inAustralia: Disteira kingii (Boulenger, 1896); Disteira major (Shaw, 1802).30

Australian Biodiversity Record, 2007 (8): 1-124Disteira kingii (Boulenger, 1896)Hydrophis kingii Boulenger, G.A. (1896): Cat. Snakes Brit. Mus. [p. 276]. Type data: holotypeBMNH 1946.1.10.10. Type locality: N Australia.Description: This is a very elongate species of sea snake with a small head that is barelydistinct from the neck. The base body colour is pale greyish-white, (more so ventrolaterally),and the head and throat is black with a distinctive white ring around the eye. The body and tailis conspicuously marked with a transverse series of about 45-50 large broad dark greyishbrownto black blotches or saddles. This pattern forms a broad banded appearance with paleinterspaces dorsally and this runs along the entire body and onto the tail. The most anterior ofthese broad dark bars is separated from the dark head by a narrow whitish bar or band at theneck. The darker blotches are more prominent on the upper lateral, as the lower area isgenerally paler (light creamish) than the pale dorsal interspaces. Sometimes these largertriangular lateral blotches merge to form a longitudinal series along the body; a similarlongitudinal series of smaller blackish spots or mini-blotches usually occurs along the lowerlateral as well. The ventral surface is dark, and this is formed by a blackish mid-ventral stripe.Some significant features of this species’ morphology are: head shields large andsymmetrical; 2-3 maxillary teeth following fang; supralabials 7-8; preocular present;postoculars 2-3; temporals 1+2; prefrontal in contact with second supralabial; anterior chinshields large and contacting mental groove (in Disteira major, the elongate infralabials contactthe mental groove, which results in the small chin shields either being completely excludedfrom contacting the mental groove, or at best allowing only point contact); body scalesstrongly imbricate and keeled dorsally, but weaker imbrication and smooth laterally; bodyscales at neck 22-29; mid-body scales in about 34-40 rows; ventrals range 299-337, small,mostly undivided and only about twice as wide as adjacent body scales; anal scales notenlarged. Attains a maximum total length of around 2.0 m., although a specimen 1.7m totallength would be very large.Distribution: Known mainly from tropical Australian seas (including New Guinea), being foundoff the coasts of Queensland, Northern Territory, Western Australia, commonly to aboutBarrow Island, with rare occurrences to about as far south as Perth. It is only known rarely inNew South Wales.Habitat: Inhabits deep water reefs as well as relatively deep continental shelf waters withsandy or muddy bottoms. Activity has been recorded in water depths ranging from around 2metres to 22 metres. Occasionally reported from inshore estuarine waters, sandy bays orfound beached after storms.Biology/Ecology: This is essentially a fast-swimming diurnal species that feeds on a widevariety of small fishes (mainly Anguilliformes). It appears to be a seasonally reproductivespecies, with gravid females being found in the Wet Season (January to March), and birthsapparently taking place during the Autumn or early Dry Season. Usually about 5 youngcomprise a litter (range 1-9). Occasional snakes are found encrusted in barnacles.Toxicity: This is species that rarely attempts to bite under normal circumstances, but cautionshould nevertheless be exerted, because its venom is highly toxic to humans. Urgent medicalattention should always be sought in the event of a bite from this species.Survival Status: Protected under the Qld Nature Conservation Act (1992), the Territory Parksand Wildlife Conservation Act (1998) and the WA Wildlife Conservation Act 1950 (asamended). Protected under the New South Wales National Parks and Wildlife Act (1974) butnot listed in that State as a Threatened Species in any of the Schedules of the NSWThreatened Species Conservation Act (1995). Listed as a Marine Protected Species underthe Australian Environmental Protection and Biodiversity Conservation Act (1999). Probablynot under threat, as it is rarely taken as by-catch in fishing operations, but its restricteddistribution in Australian waters could potentially make it vulnerable in some parts of its range.Etymology: The name 'kingii' honours British explorer and naturalist Admiral Philip ParkerKing (1791-1856) who collected the Type Specimen.31

Australian Biodiversity Record, 2007 (8): 1-124Disteira major (Shaw, 1802)Hydrus major Shaw, G. (1802): General Zoology, or Systematic Natural History. [p. 558, pl.124] [Note: other syntype, BMNH iii.9.1.a, referred to Astrotia stokesii]. Type data: lectotypeBMNH 1946.1.9.24. Subsequent designation: Cogger, H.G. (1983): Zool. Cat. Aust., 1.Amphibia and Reptilia. [see p. 246]. Type locality: Indian Ocean (as Indian Seas).Disteira doliata Lacépède, B.G.E. (1804): Ann. Mus. Natl Hist. Nat. Paris 4: 184-211 [199,210, pl. 57 fig. 2]. Type data: type status unknown (presumed lost). Type locality: Australia.Pelamis shavii Merrem, B. (1820): Tentamen Systematis Amphibiorum. [p. 139] [unnecessaryreplacement name for Hydrus major Shaw, 1802].Hydrophis mentalis Gray, J.E. (1842): Monographic Synopsis Water Snakes Hydridae. [p. 62].Type data: holotype BMNH 1946.1.9.24. Type locality: Indian Ocean.Disteira dumerilii Jan, G. (1859): Rev. Mag. Zool. (2)11: 148-157 [p. 149]. Type data: holotypeMNHP 7705. Type locality: Australia.Hydrophis lacepedei Jan, G. (1859) : Prodr. Iconogr. Ophidiens..Rev. Mag. Zool. (2)10 and(2)11-12, 1858 and 1859. [pl. D] [lapsus for Disteira dumerilii Jan, 1858]Distira nasalis De Vis, C.W. (1905): Ann. Qd Mus. 6: 46-52 pl. 15 [p. 48]. Type data: holotypeQM J203. Type locality: QLD coast.Description: This is a more robust-bodied species of sea snake than D. kingii, with a shortdeep head that is barely distinct from the neck. The base body colour is pale greyish-white toolive green, with the head tending to be a shade of olive green or olive-brown, with scattereddarker flecks. The body is boldly marked with a series of around 25-30 large broad darkbrown to black blotches or saddles across the body, each separated by a slightly wider paleinterspace. The large saddle-like blotches extend to about mid-way down the side of thebody. The lower lateral area is creamish-yellow. Within each pale interspace there is a thin(about 1 scale wide) black transverse ‘band’ in the centre, and each ‘band’ extends down thebody to about the mid-lateral area. This pattern forms a double banded appearance withalternating thick and thin dark blotches or bands along the body. In some the lower flanksmay be dark blotched as well, and these large lower blotches are below both the thininterspace bands and the dorsal saddle-like blotches. The overall pattern tends to be moreconspicuous in immatures and less defined in mature individuals. Ventrally, most specimensare dark grey, but some may have dark flecking on the ventrals as well. Some significantfeatures of this species’ morphology are: head shields large and symmetrical; 7-8 (rarely 9)maxillary teeth following fang; supralabials 7-8; preocular present; postoculars 1-2 (usually 2);prefrontal in contact with second supralabial; primary temporals usually 2; anterior chinshields barely contacting mental groove (usually prevented from contact by elongate firstinfralabial); body scales imbricate and bluntly keeled dorsally, but smooth laterally; bodyscales around neck 30-36; mid-body scales in about 33-45 rows; ventrals with two keels;ventrals around about 195-265, small, bicarinate, and only slightly larger than adjacent bodyscales; anal scales enlarged; Attains a maximum total length of around 1.6 m although anaverage adult would be around 1.4 m., and females tend to be somewhat longer than males.Females are sexually mature at 0.7m snout-vent length.Distribution: Largely confined to tropical Australian and New Guinean seas, being foundmainly off the coasts of New Guinea, Queensland, Northern Territory and Western Australia,usually to about Shark Bay, but rarely as far south as Bunbury. Also reaches New Caledonia.Rare sightings may occur in New South Wales waters as well.Habitat: Commonly found in deep (to about 22 metres) turbid waters in the vicinity of muddyor sandy river deltas, including relatively shallow tropical tidal rivers and creeks, as well as ingulf waters and deeper continental shelf waters.32

Australian Biodiversity Record, 2007 (8): 1-124Biology/Ecology: This is a diurnal fast swimming species that feeds on a wide variety of smallfishes, including catfishes. Gravid females have been found in the mid Wet Season (January-February), and it is believed that the young are born at the end of the Wet. From 2 to 10young are produced in a litter, with the average being around 5 or 6. Larger females tend toproduce more offspring in a litter, and females appear to be more abundant than males.Toxicity: Rarely attempts to bite under normal circumstances, but caution should neverthelessbe exerted, because its venom is highly toxic to humans. Although there have been no knownhuman fatalities arising from the bite of this species, urgent medical attention should alwaysbe sought in the event of an envenomation.Survival Status: Protected under the Qld Nature Conservation Act (1992), the Territory Parksand Wildlife Conservation Act (1998), the WA Wildlife Conservation Act 1950 (as amended).Protected under the New South Wales National Parks and Wildlife Act (1974) but not listed inthat State as a Threatened Species in any of the Schedules of the NSW Threatened SpeciesConservation Act (1995). Listed as a Marine Protected Species under the AustralianEnvironmental Protection and Biodiversity Conservation Act (1999). Probably not underthreat, but its restricted distribution in Australian waters and the common extent of itsoccurrence as by-catch in trawling operations in northern Australia could potentially make itvulnerable in some parts of its range.Etymology: The name 'major' means 'larger'.Genus Enhydrina Gray, 1849Enhydrina Gray, J.E. (1849). Cat. Spec. Snakes Coll. Brit. Mus. [p. 47]. Type species: Hydrusvalakadyn Boie, 1827 by subsequent designation [see Klemmer, K. (1963): Liste rezentenGiftschl.]Diagnosis: As presently defined, a genus containing two moderate-sized Hydrophiid snakeswith an elongate forebody and broadly compressed hindbody. The following combination ofcharacters readily distinguish this genus from all other Hydrophiid snakes: head scalesenlarged, symmetrical; valvular nostrils and lingual fossa; 3-4 maxillary teeth following fang;mental groove present and very distinct; mental scale long and narrow, splint or daggershaped,much longer than broad and partially hidden in the deep mental groove; anteriorinfralabials strongly elongate, each connected by skin which is capable of extensiveexpansion during feeding; body scales imbricate, and keeled; ventrals not enlarged, similar insize to adjacent body scales; anterior of body strongly compressed; tail strongly compressedvertically, paddle-like; ovoviviparous; diploid number of chromosomes: 32 in males and 33 infemales. Content in Australia: Enhydrina schistosa (Daudin, 1803); Enhydrina zweifeli Kharin,1985.Enhydrina schistosa (Daudin, 1803)Hydrophis schistosus Daudin, F.M. (1803): Histoire Naturelle, générale et particulière desReptiles. [p. 386] [description based on pl. 10 in Russell, P. (1801). A Continuation of anAccount of Indian Serpents. London: Shakespeare Press 15 pp., in which no type locality wasspecified]. Type data: holotype BMNH 1946.1.10.7. Type locality: Tranquebar, India.Hydrophis schistotus Jan, G. (1859). Prod. Iconogr. Ophidiens. Rev. Mag. Zool. (2)10 and(2)11-12, 1858 and 1859. [p. 23] [errore for Hydrophis schistosus Daudin, 1803].Disteira russelii Fitzinger, L.J. (1827): Isis Oken 20: 731-741 [p. 733]. Type data: holotype(probable) whereabouts unknown (described in pl. 11 in Russell, P. (1801). A Continuation ofan Account of Indian Serpents. London : Shakespeare Press 15 pp.). Type locality: Asia,Indian Ocean.Hydrus valakadyn Boie, F. (1827): Isis Oken 20: 508-566 [p. 554]. Type data: holotypewhereabouts unknown (described as pl. 11 in Russell, P. (1801). A Continuation of an33

Australian Biodiversity Record, 2007 (8): 1-124Account of Indian Serpents. London : Shakespeare Press 15 pp.). Type locality: Tranquebar,India.Hydrophis bengalensis Gray, J.E. (1842): Monogr. Synop. Water Snakes Hydridae [p. 62].Type data: holotype BMNH 1946.1.1.73. Type locality: Bengal, India and Bangladesh.Hydrophis subfasciata Gray, J.E. (1842): Monogr. Synop. Water Snakes Hydridae [p. 62].Type data: holotype BMNH 1946.1.1.90. Type locality: Bengal, India and Bangladesh.Thalassophis werneri Schmidt, P. (1852): Abh. Geb. Naturw. Hamburg 2: 69-86 [84, pl. 6].Type data: holotype ZMH 392. Type locality: Samarang, Java, Indonesia.Description: A moderate-sized species that has a very slender forebody, and the posteriorthird of the body large and broadly compressed. The colouration and patterning seems to varyconsiderably across its range, and the Australasian population appears to be morphologydistinct from those of Asia. Kharin has applied the name Enhydrina zweifeli to this palecoloured/darkblotched population. However, it appears that both species of Enhydrina mayoccur in Australian waters, so the following description is largely composite until morematerial can be examined. In Australian waters, adults are usually a uniform grey dorsally onthe head and body, and creamish-white ventrally. In some individuals, there may be a seriesof large dark bars or blotches on the dorsum, and these can be about the same width as thepaler interspaces, or in some much larger. These large blotches extend across the dorsum, toabout mid-laterally. There is no suborbital stripe on the side of the head in E. schistosa.Juveniles have around 45-55 blackish transverse bars over the body, in marked contrast tothe greyish rings of juveniles from Asia. Some characteristic features of this species are: headsmall and barely distinct from the neck, with enlarged symmetrical shields; 3-4 maxillary teethfollowing fang; 5-6 palatine teeth; palatine teeth larger than pterygoid teeth; mental groovepresent and very distinct; mental scale long and narrow, splint or dagger-shaped, muchlonger than broad and partially hidden in the deep mental groove; anterior infralabials veryelongate; throat capable of considerable extension due to extensile skin structure; anterior ofbody slender, but posterior deep and strongly compressed; body scales imbricate, in 49-66rows at mid-body; body-scale rows at neck 40-55; body scales each with a short, low keel,with the keels being slightly larger in males; ventrals about 240-320 (females do not appear tohave a higher ventral count than that of males); ventrals not enlarged, similar in size toadjacent body scales; preanal scales slightly enlarged. The largest specimens have beenreported in Asian populations (up to 1.4m) but those from Australia are not particularly large.In Australia, this is species only attains a maximum snout-vent length of only around 85 cm.,with exceptional specimens reaching around 1.1 m in total length. Females are generallylarger than males, and males have a slightly longer tail than females.Distribution: This species as presently defined, occurs as many scattered populations acrossa vast area of tropical waters from south of the Seychelles and Madagascar, the Arabian Seaand Persian Gulf (off Oman), the seas off South Asia (Pakistan, India and Bangladesh),Southeast Asia (Myanmar (formerly Burma), Thailand, Vietnam), and through westernIndonesia to Australasia. In our region, it is largely confined to tropical seas between Australiaand New Guinea, and is generally thought to be either extremely rare or absent fromAustralian territorial waters. However, I have observed this species being commonly trawledin Queensland and Northern Territory continental shelf waters as bycatch from prawn trawlingoperations, so it may be more widespread here - at least seasonally.Habitat: Throughout its range, it mainly inhabits relatively shallow water depths from about 4to 22 metres depths over muddy or sandy bottoms. It inhabits river deltas, estuaries, bays,harbours and gulf waters, and near-shore reefs as well as the shallower continental shelfwaters. Also sometimes observed well-up tidal sections of rivers, and in parts of Asia it hasbeen found in rivers many kilometres upstream from the sea.Biology/Ecology: Although this is the commonest species of sea snake in parts of Asia, inAustralian waters it is relatively uncommon compared to Hydrophis species. It appears to bean essentially nocturnal species that is occasionally observed foraging near the surface afterdark. It feeds on a variety of small fishes, such as catfish and puffer-fishes, and also34

Australian Biodiversity Record, 2007 (8): 1-124consumes prawns. In general, the prey size is related to the growth stage of the snake, withsmaller snakes tending to feed on smaller prey such as prawns and smaller fish, and largersnakes preferring larger fish. This could explain the unique morphology of its highlyelasticised anterior throat skin that would facilitate the swallowing of the very large fishes thathave been found in the stomachs of this species. Generally, larger snakes do not eat smallprey, indicating that their foraging methods may be less effective in securing this resource.Prey is actively hunted in the dark while the snake is swimming near the bottom, and fishesare apparently detected by tactile means such as touch by the snake or perhaps pressurewaves from the swimming fish rather than by sight or smell. Once a fish is seized, it is heldfirm in the jaws until it stops moving, and the prey is not constricted. During feeding, thesnake may orientate its body in the current to reduce its effect as it manipulates the prey toease swallowing, or on occasion it may press the prey against a hard object so as to get abetter grip. Prey is usually swallowed head-first, and after swallowing, snakes tend to swimaround actively or even twist their body into a knot so as to facilitate the movement of the preyinto the stomach. During the Wet Season it can be more often found in the estuarine (tidal)parts of both large rivers and smaller creeks - particularly in shallow waters of mangroveforests - rather than in more open waters. Males are known to practice combat behaviour, butwhether this is the result of competition for mates, prey or territoriality is unclear. Thereproductive biology is poorly known in this widespread species, and there is some notablevariation both geographically and seasonally. The sex ratio for the species does not differsignificantly from parity over the species’ range. In Asia, gravid females have been found fromaround August to January (late Dry Season, through to the Wet Season), with births occurringin February-March. Depending upon the area, it would appear that a gestation period of atleast 3 or 4 months occurs, and the timing of births is correlated with the end of the Monsoonor Wet Season. In Australia, mating has been recorded in May (early Dry Season). This is anovoviviparous species, with recorded litters ranging from 3 to 34, although the averagenumber of offspring in a litter is around 7 to 18 (larger females tend to produce larger litters).Juveniles are around 230-240 mm in length at birth in Australian waters, but they appear tobe somewhat larger in Asia, at around 300-340mm when born. It has a life span of around 4years. This species is known to be occasionally infested with parasitic worms, and onoccasions may carry small colonies of bryozoans attached to its skin.Toxicity: Although it has been long regarded as a relatively inoffensive species, bites tohumans are well-known in Asia, and on occasion have resulted in fatalities - indeed, mostfatalities from all recorded sea snake bites have been from this species. I have observed thatthis species will readily try to bite if handled or harassed, so extreme caution should beexerted when approaching or handling this snake. Venom studies have indicated that this is adangerously venomous species. As there have been human fatalities arising from the bite ofthis species elsewhere, urgent medical attention should always be sought in the case of anenvenomation.Survival Status: Protected under the Qld Nature Conservation Act (1992) and the TerritoryParks and Wildlife Conservation Act (1998). Listed as a Marine Protected Species under theAustralian Environmental Protection and Biodiversity Conservation Act (1999). Probably notunder threat, but its apparently restricted distribution in Australian waters could potentiallymake it vulnerable in some parts of its range to large scale fishing operations.Etymology: The name ‘schistosa’ means very divided. The name ‘zweifeli’ honours Americanherpetologist Richard Zweifel.Additional Comment: This is a highly variable species that has sometimes been placed withinthe genus Disteira, but it is herein considered distinctive enough to warrant separate genericrecognition as has long been maintained by Cogger and other authorities. Its morphologysuggests that there may be additional species in this genus awaiting formal recognition inAsia, and that both Enhydrina schistosa and Enhydrina zweifeli may occur in Australianterritorial waters. The northern and eastern Australian population is almost certainlyEnhydrina zweifeli Kharin, 1985 [see Kharin, V.E (1985): Zool. Zh., 64(5): 785-787. Typelocality: ‘off mouth of Sepik River, PNG’] and those of the far north-west and Indian Oceanwaters Enhydrina schistosa.35

Australian Biodiversity Record, 2007 (8): 1-124Genus Hydrophis Sonnini and Latreille, 1802Hydrophis Sonnini de Manoncourt, C.S. and Latreille, P.A. (1802): Hist. Nat. Reptiles [p. 193].Type species: Hydrophis laticauda Sonnini and Latreille, 1802 by subsequent designation[see Smith, M.A. (1926). Monogr. Sea Snakes]Dolichodira Wall, F. (1921). Ophidia Taprobanica or the Snakes of Ceylon [p. 399]. Typespecies: Hydrophis diadema Günther, 1864 (= Hydrophis torquata Günther, 1864) bymonotypy.Polypholophis Wall, F. (1921). Ophidia Taprobanica or the Snakes of Ceylon. [p. 380]. Typespecies: Hydrophis neglectus Wall, 1906 (= Hydrophis stricticollis Günther, 1864) bymonotypy.Porrecticollis Wall, F. (1921). Ophidia Taprobanica or the Snakes of Ceylon. [p. 335]. Typespecies: Hydrophis obscurus Daudin, 1803 by monotypy.Diagnosis: A genus of small to moderate-sized Hydrophiid snakes with a small head notdistinct from the narrow cylindrical neck, and the anterior part of body usually much moreattenuate than the posterior which is robust, deep and highly compressed. Readilydistinguished from all other genera by the following combination of characters: head shieldsenlarged, symmetrical; less than 9 maxillary teeth following fang; in all but one species, theanterior part of maxilla is arched upwards, the tip of the longer fang on a line connecting thetips of the solid maxillaries (in the exception, Hydrophis elegans, the anterior part of maxillanot arched upwards, the tip of the fang projects slightly below a line connecting the tips of thesolid maxillaries); parietal bone not contacting prefrontal; valvular nostrils and lingual fossa;rostral normal, triangular, broader than high; 3rd and 4th supralabials contacting orbit; firstinfralabial normal, about as long as pregenial; nasal separated from preocular by high 2ndsupralabial; mental groove present and distinct; mental triangular, broader than long, and notpartially hidden in the shallow mental groove; anterior chin scales large, and usually borderingmental groove; body scales imbricate, with those on neck and posterior of body each with acentral tubercle or short keel; usually 25 or more scale rows on neck; mid-body scales in 35-50 rows; ventrals small, about 325-515; posterior ventrals distinct, mostly undivided, neverfoliform and barely wider than adjacent body scales; tail strongly compressed vertically,paddle-like; ovoviviparous. Etymology: The name Hydrophis means ‘water snake’, and isderived from the Greek ‘hydros’ - water, and ‘ophis’ - snake. Content: Hydrophis atricepsGunther 1864; Hydrophis bituberculatus Peters 1872; Hydrophis brooki Gunther 1872;Hydrophis elegans (Gray 1842); Hydrophis fasciatus (Schneider 1799); Hydrophis klossiBoulenger 1912; Hydrophis laboutei Rasmussen and Ineich 2000; Hydrophis lamberti Smith1917; Hydrophis lapemoides (Gray 1849); Hydrophis mcdowelli Kharin 1983; Hydrophismelanosoma Gunther 1864; Hydrophis obscurus Daudin 1803; Hydrophis parviceps Smith1935; and, Hydrophis vorisi Kharin 1984. Content in Australia: Hydrophis atriceps Gunther,1864; Hydrophis elegans (Gray, 1842); Hydrophis mcdowelli Kharin, 1984; Hydrophismelanosoma Gunther, 1864; Hydrophis vorisi Kharin, 1984.Hydrophis atriceps Gunther, 1864Hydrophis atriceps Günther, A. (1864): Reptiles of British India. [p. 371, pl. 25 fig. 1]. Typedata: syntypes BMNH 1946.1.2.62, BMNH 63.9.29.50. Type locality: Thailand.Hydrophis alcocki Wall, F. (1906): Mem. Asiat. Soc. Beng. 1: 277-299 [p. 288, pl. 15 fig. 3].Type data: holotype IM 14470 [see Smith, M.A. (1926). Monogr. Sea Snakes (Hydrophiidae)].Type locality: Puri, Orissa, India.Disteira cincinnatii van Denburgh, J. and Thompson, J.C. (1908): Proc. Calif. Acad. Sci. 3: 41-48 [42, pl. 1]. Type data: holotype CAS 15016. Type locality: 1 mi[les] NE of Cavite, ManilaBay, Philippines.Description: This species has a relatively small head which is barely distinct from the neck.The anterior part of body is much more slender than the posterior which is deep and highly36

Australian Biodiversity Record, 2007 (8): 1-124compressed. The head is black, even in adults, and sometimes has a yellow spot behind eyeand/or nostril. The base body colour is essentially pale ranging from yellow through variousshades of lighter brown. Young specimens are prominently marked with series of 50 to 70dark blackish transverse bands across the dorsum, with some completely encircling the body.However, this banding is less distinct with age, with adults having more of a blotched pattern.The ventrals are black anteriorly, but progressively becoming paler posteriorly to more of adark grey on the last third of the venter and tail. It is readily distinguished by the followingcombination of characters: head shields enlarged, symmetrical; anterior part of maxilla archedupwards, the tip of the longer fang on a line connecting the tips of the solid maxillaries; 5-6maxillary teeth following fang; parietal bone not contacting prefrontal; rostral normal,triangular, broader than high; 3rd and 4th supralabials contacting orbit; first infralabial normal,about as long as pregenial; nasal separated from preocular by high 2nd supralabial; mentalgroove present and distinct; mental triangular, broader than long, and not partially hidden inthe shallow mental groove; anterior chin scales large, and usually bordering mental groove;body scales imbricate, with those on neck and posterior of body each with a central tubercleor short keel; usually 25-30 scale rows on neck; mid-body scales in 35-49 rows; ventralssmall, about 320-515; posterior ventrals distinct, mostly undivided, never foliform and barelywider than adjacent body scales. A maximum total length of only around 1.2 m. is attained; atbirth the young are around 300mm in length. Females are sexually mature at around 650 mmin total length, and females tend to be somewhat larger than males and reach a longermaximum length. Males tend to have longer tails than females.Distribution: This species is known mainly from South East Asia - such as Singapore,Thailand, Vietnam and China - although it extends into the waters of Indonesia and thePhilippines to New Guinea and Australia. In Australian waters it is only known from theArafura Sea between the Northern Territory, Queensland and New Guinea.Habitat: Inhabits reefs in relatively shallow continental waters. It has been observed in wateraround 20 metres deep.Biology/Ecology: This is a fast swimming, essentially diurnal species that feeds on a variety ofsmall fishes such as eels near the bottom. When ascending to the surface to breath duringthe day, its movements are swift and direct to gain a quick breath of air before rapidlyreturning to the bottom. However, at night it has been observed casually swimming near thesurface. The reproductive biology various somewhat across this species enormous range,with oviducal young being present in the late Dry Season (May-August) to early Wet Season(July-December) in Indonesia. However, in the Philippines, oviducal young have beenrecorded in the early Dry season (January). This implies that young are born during the earlyWet Season to early Dry Season. Litter size ranges from 1 to 10 (usually 3 or 4).Toxicity: This species rarely attempts to bite under normal circumstances, but caution shouldnevertheless be exerted, because its venom is highly toxic to humans. Urgent medicalattention should always be sought in the event of a bite from this species.Survival Status: Protected under the Qld Nature Conservation Act (1992) and the TerritoryParks and Wildlife Conservation Act (1998). Listed as a Marine Protected Species under theAustralian Environmental Protection and Biodiversity Conservation Act (1999). Probably notunder threat, but its restricted distribution in Australian waters could potentially make itvulnerable in some parts of its range.Etymology: The name 'atriceps' means 'dull black head', and refers to the head colouration ofthe species.37

Australian Biodiversity Record, 2007 (8): 1-124Hydrophis elegans (Gray, 1842)Aturia elegans Gray, J.E. (1842). Monogr. Synop. Water Snakes [p. 61]. Type data: holotypeBMNH 1946.1.3.89. Type locality: Port Essington, NT.Distira grandis Boulenger, G.A. (1896). Cat. Snakes Brit. Mus [p. 293]. Type data: syntypesBMNH 1946.1.17.10-11, BMNH 70.11.30.71. Type locality: Malay Archipelago and QLD,QLD.Description: This is large species of Hydrophiid snake with a small head not distinct from theneck, and the anterior part of body much more slender than the posterior - which is robust,deep and highly compressed. The colouration is somewhat variable in this species. Adults arepale brown with only a faint pattern of darker blotches usually evident. Sometimes adults haveprominent dark banding that is more intense anteriorly, and often with secondary markingsbetween the main bands, and two alternating rows of dark spots along the flanks - one alongthe mid-lateral placing a dark spot below a pale interspace, and another along the lowerlateral of the body placing a spot below a dark transverse bar or blotch. The pale interspacesub-pattern comprises either rows of dark scales, spots or blotches within the paleinterspaces between the dark bands, and this is usually more obvious about the neck region,or along the posterior of the body and tail. However, juveniles and immatures are prominentlymarked with tan brown on the body, a black head and about 35-55 distinct black bands on thebody; these bands are broadest on the dorsum and venter, but somewhat narrow orincomplete on the flanks. The tip of the tail is greyish, and this is a paler grey than that of thedarker cross-bands - when bands are present. The ventral colour anteriorly is black, butbecoming paler posteriorly with alternating dark barring. Some significant features of thisspecies morphology are: head shields enlarged, symmetrical; anterior part of maxilla notarched upwards, and the tip of the fang projecting conspicuously below a line connecting thetips of the solid maxillaries; 6-7 maxillary teeth following fang; nasal separated from preocularby high 2nd supralabial; preocular single; postoculars 1-2 (mostly 2); one large primarytemporal; supralabials 6-7; usually only 3rd and 4th supralabial contacting orbit; mentalgroove present and distinct; mental triangular, broader than long, and not partially hidden inthe shallow mental groove; anterior chin scales large, and usually bordering mental groove;anterior part of body much more slender than the posterior, which is very deep and highlycompressed; dorsal body scales weakly imbricate, and slightly keeled or smooth; body scales25-30 rows at neck; mid-body scales in 35-49 rows; ventrals about 345-430 (females have ahigher ventral count than males); ventrals small and smooth and never foliform; posteriorventrals distinct, mostly undivided, and scarcely wider than adjacent body scales (exceptanteriorly, because ventrals here are often entire). Reaches a maximum total length of around2.2m, although a large specimen would be around 1.7 m. Females are sexually mature atabout 1.2m and tend to be noticeably larger than males.Distribution: This sea snake is known mainly from tropical and sub-tropical Australian seas,being found off the coasts of Queensland and the Northern Territory. It occurs commonly inWestern Australia, down to about Shark Bay or more rarely even as far south as GeographeBay in the extreme south-west. On occasions in summer, specimens occur off New SouthWales, Victoria and even Tasmania. Also known from the waters around southern NewGuinea.Habitat: Occurs in a variety of conditions, but more often found around deep water reefs (upto around 80 metres in depth). It has also been observed over soft muddy or sandy sedimentsin relatively shallow, turbid waters of river deltas, bays and gulfs on continental shelf areas(ranging from around 2m to 20m depths), and is known to enter tidal stretches of tropicalrivers several kilometres upstream from the sea.Biology/Ecology: Essentially a diurnal and fast swimming species that feeds on small squidand octopuses. It has also been reported to feed on a variety of small fishes - principally eels.Ovoviviparous, producing up to 33 (usually about 12) young in a brood during the late WetSeason to early Dry Season (March to May). The new-born young are around 400 to 500 mmin total length. It would seem that not all females are reproductive in any one year, implying38

Australian Biodiversity Record, 2007 (8): 1-124that individual females may not reproduce every year. On occasions, specimens are foundwith encrustations of barnacles, hydrozoans and bryozoans on the skin.Toxicity: This species rarely attempts to bite under normal circumstances, but caution shouldnevertheless be exerted, because when fishermen accidentally capture the species, it hasbeen observed to readily bite nets and other fishing gear. Studies have revealed that itsvenom is highly toxic to humans, so urgent medical attention should always be sought in theevent of a bite from this species - but there have been no recorded fatalities in Australia todate.Survival Status: Protected under the New South Wales National Parks and Wildlife Act (1974)but not listed in that State as a Threatened Species in any of the Schedules of the NSWThreatened Species Conservation Act (1995). Also protected under the Victorian Wildlife Act(1975) but not listed as threatened in the Victorian Flora and Fauna Guarantee Act (1988)],the Qld Nature Conservation Act (1992), the Territory Parks and Wildlife Conservation Act(1998), the Tasmanian National Parks and Wildlife Act (1970) and the WA WildlifeConservation Act 1950 (as amended). Listed as a Marine Protected Species under theAustralian Environmental Protection and Biodiversity Conservation Act (1999). Probably notunder threat, due to its great abundance but its scattered distribution in Australian waters andhigh frequency of accidental capture during trawling operations could potentially make itvulnerable in some parts of its range.Etymology: The name ‘elegans’ means elegant, and probably refers to the gracile body formof the species.Hydrophis mcdowelli Kharin, 1984Hydrophis macdowelli Kharin, V.E. (1983): Zool. Zh. 62(11): 1751-1753 [p.1751]. Type data:holotype ZIL 19678. Type locality: Northern Australian Shelf.Description: This is a medium-sized seasnake with a very small narrow head that is notdistinct from the neck, an extremely elongate forebody and a deep, strongly compressedposterior body. The base body colour is pale creamish dorsally, gradually becoming moreyellowish on the lower ventro-lateral and ventral surfaces. Adults possess a series of blackishdorsal blotches on the dorsum, with each blotch terminating about the upper to mid-lateral ofthe body. In some specimens the pale interspaces can possess a blackish narrow line orseries of broken bars or spots as well. There are three separate longitudinal rows of darkermarkings along the flanks as follows: an upper lateral line of dark greyish inverted triangle orelongate rhomboid-shaped markings with each below a pale interspace; a reduced series ofirregular blotches along the lower lateral zone, with each blotch being positioned below a darkdorsal blotch; a third series of tiny blackish or greyish spots or bars, each below a paleinterspace. The tail is usually prominently marked with 4 or 5 wide but faint greyish or blackishtransverse bands, with a narrow blackish line within each pale interspace. The dorsum of thehead is dark olive or black, and the throat and anterior venter is black fading to dark grey, withthe rest of the ventrals yellowish. Some significant features of this species’ morphology are:Head shields enlarged and symmetrical; 5-6 maxillary teeth following fang; preocular 1;postoculars 2; supralabials 7, with 3rd and 4th contacting orbit; primary temporals 1; bodyscales smooth, imbricate but keeled on neck; 23-26 body scale rows on neck; mid-bodyscales in 35-42 rows; ventrals smooth, about 243-274 and mostly entire, and wider thanadjacent body scales; Usually attains a maximum total length of only around 80 cm., althoughit can reach a total length of nearly 1.2m.Distribution: Known only from tropical Australian seas, off the coasts of Queensland, NorthernTerritory and Western Australia to about as far south as the Pilbara coast. Also known from asmall part of the southern coast of New Guinea, and across to New Caledonia.Habitat: More often found in shallow (about 7-25 metres deep) waters over flat open sandybottoms of bays and gulfs, but also known from turbid conditions of estuaries and river deltas,but also rarely in deeper waters on continental shelf areas.39

Australian Biodiversity Record, 2007 (8): 1-124Biology/Ecology: This is a very common species that mainly occurs in near coastal situations.It appears to be an essentially diurnal species that feeds on a variety of small fishes such aseels which they hunt by actively probing the sandy bottoms. It produces up to 5 (usually 2-3)living young in a brood (ovoviviparous).Toxicity: This is a fast swimming species that rarely attempts to bite under normalcircumstances, but caution should nevertheless be exerted, because its venom is highly toxicto humans. Urgent medical attention should always be sought in the event of a bite from thisspecies.Survival Status: Protected under the Qld Nature Conservation Act (1992), the Territory Parksand Wildlife Conservation Act (1998) and the WA Wildlife Conservation Act 1950 (asamended). Listed as a Marine Protected Species under the Australian EnvironmentalProtection and Biodiversity Conservation Act (1999). Probably not under threat, but itsrestricted distribution in Australian waters could potentially make it vulnerable in some parts ofits range.Etymology: The name 'mcdowelli' honours American herpetologist Samuel B. McDowell.Hydrophis melanosoma Gunther, 1864Hydrophis melanosoma Günther, A. (1864): Reptiles British India. [p. 367, pl. 25 fig. e]. Typedata: holotype BMNH 1946.1.10.6. Type locality: unknown.Hydrophis floweri Boulenger, G.A. (1898): Proc. Zool. Soc. Lond. 1898: 106-107 [106, pl. 9].Type data: syntypes BMNH 1946.1.9.48-51. Type locality: Brunei Bay, Borneo.Description: This is a small species, with a moderate head size that is barely distinct from theneck. The body-form is not noticeably as slender anteriorly as other Hydrophis, and onlymoderately compressed posteriorly. The base body colour is pale - creamish to yellowish -with adults possessing a black head and 50-70 black transverse bands that are about twicethe width of the pale interspaces. Adults tend to be less prominently marked as they age.Juveniles also have a black head, and are greyish on the upper body with prominent blackbands that taper to a point about mid-laterally; ventrally juveniles are blackish. Some of thesignificant features of this species’ morphology are: head shields enlarged, symmetrical;nasal separated from preocular by high 2nd supralabial; supralabials 6-7, with 3rd and 4thsupralabials contacting orbit; mental groove present and distinct; mental triangular, broaderthan long, and not partially hidden in the shallow mental groove; anterior chin scales large,and usually bordering mental groove; parietal bone contacting the prefrontal; anterior part ofmaxilla arched upwards, the tip of the longer fang on a line connecting the tips of the solidmaxillaries; 6-8 maxillary teeth following fang; body scales on neck and posterior of bodyeach with a weak central tubercle or short keel; usually 25 (range 21 to 28) scale rows onneck; mid-body scales in 37-43 rows; body scales weakly imbricate anteriorly; ventrals 260-370, small ventrals, each about as wide as, or only slightly wider than adjacent body scalesand never foliform; posterior ventrals distinct, mostly undivided. Attains a maximum totalsnout-vent length of only around 1.1 m. although larger specimens around 1.5m. are known.Distribution: Widely distributed across South East Asia to about the Moluccas in easternIndonesia. In Australian waters only known from reefs off north-western Western Australia,principally Ashmore Reef and Scott Reef. It reputedly occurs in Northern Territory waters aswell, and has been recorded in Torres Strait, Queensland.Habitat: Largely restricted to reefs and sea-grass beds in relatively shallow continentalwaters.Biology/Ecology: Essentially a fast-swimming diurnal species that feeds on a variety of smallfishes - principally eels. Ovoviviparous, producing about 6 living young in a brood.Toxicity: This species rarely attempts to bite under normal circumstances - and then onlyspontaneously, but caution should nevertheless be exerted, because its venom is highly toxic40

Australian Biodiversity Record, 2007 (8): 1-124to humans. Urgent medical attention should always be sought in the event of a bite from thisspecies, although there have been no known fatalities recorded.Survival Status: Protected under the Qld Nature Conservation Act (1992), the Territory Parksand Wildlife Conservation Act (1998) and the WA Wildlife Conservation Act 1950 (asamended). Listed as a Marine Protected Species under the Australian EnvironmentalProtection and Biodiversity Conservation Act (1999). Probably not under threat, but itsrestricted distribution in Australian waters could potentially make it vulnerable in some parts ofits range.Etymology: The name 'melanosoma' refers to the black bands on the body.Hydrophis vorisi Kharin, 1984Hydrophis vorisi Kharin, V.E. (1984): Zool. Zh. 63(4): 630-632. Type data: holotype AMNH58869. Type locality: eastern bank of Fly River, opposite Sturt Island, Western Province,Papua New Guinea.[Note: Previous records of this species in Australian waters were misidentifications of thesoutheast Asian species Hydrophis obscurus Daudin, 1803]Description: This is a medium-sized seasnake with a very small narrow head that is notdistinct from the neck, a very elongate forebody and a deep, compressed posterior body. Thebase body colour is pale greyish to whitish, with the head darker. There is prominenttransverse banding that encircles the entire body and tail, and there are usually no othermarkings within the pale interspaces along the body, with the exception of the extremeposterior portion where small blackish lines or flecking may be also present. The large series(70-80) of dark, almost black, crossbands are wider on the dorsum than the pale interspaces,but laterally, they taper somewhat to be about the same width as the paler interspaces. Insome specimens the dark bands are broken or interrupted along the vertebral line on thebody. The tail may have in addition to the banding pattern, scattered black marks within thepale interspaces. Some of the significant features of this species’ morphology are: headshields enlarged, symmetrical; parietal bone contacting the prefrontal; anterior part of maxillaarched upwards, the tip of the longer fang on a line connecting the tips of the solid maxillaries;6 maxillary teeth following fang; nasal separated from preocular by high 2nd supralabial; 3rdand 4th supralabials contacting orbit; mental groove present and distinct; mental triangular,broader than long, and not partially hidden in the shallow mental groove; anterior chin scaleslarge, and usually bordering mental groove; fewer than 25 body scale rows at neck; mid-bodyscales in 29-37 rows; body scales smooth and weakly imbricate anteriorly, but keeledposteriorly; ventrals 330-350, small, each about as wide as, or only slightly wider thanadjacent body scales, and never foliform; posterior ventrals distinct and mostly undivided.Attains a maximum total length of only around 0.6 m.Distribution: Known only from a relatively small part of Australia and New Guinea, centred onthe Torres Strait area of Queensland and adjacent parts of near coastal Papua New Guinea.Habitat: This species has been detected living mainly around coral reefs in shallowcontinental waters, but has also been recorded from inshore estuarine conditions as well.Biology/Ecology: This appears to be at least a partly diurnal species that feeds on a variety ofsmall fishes. Ovoviviparous.Toxicity: This is a small species that rarely attempts to bite under normal circumstances, butcaution should nevertheless be exerted, because its venom is likely toxic to humans. Urgentmedical attention should always be sought in the event of a bite from this species.Survival Status: Protected under the Qld Nature Conservation Act (1992). Listed as a MarineProtected Species under the Australian Environmental Protection and BiodiversityConservation Act (1999). Probably not under threat, but its restricted distribution in Australianwaters could potentially make it vulnerable in some parts of its range.41

Australian Biodiversity Record, 2007 (8): 1-124Etymology: The name ‘vorisi’ honours American herpetologist Harold K. Voris.Genus Leioselasma Lacepede, 1804Leioselasma Lacépède, B.G.E. (1804): Ann. Mus. Natl Hist. Nat. Paris 4: 184-211 [seeKharin, V.E. (1984): Zool. Zh. 63(10): 1535-1546 for the formal definition and re-instatementof the genus Leioselasma]Leioselasmus Fischer (1808): Tabl. Syn. Zoog., ed. 2, Rept. Col., (7) [nom. nov. proLeioselasma Lacepède 1804]Lielasma Agassiz (1846): Nomen. Zool. Index Univ. [emend. pro Leioselasma Lacepède1804]Diagnosis: A genus of moderately large species of Hydrophiid snakes readily identified by thefollowing combination of characters: anterior part of body much more slender than theposterior, which is deep and moderately compressed; head shields enlarged, symmetrical;valvular nostrils and lingual fossa; 5-8 maxillary teeth following fang; supralabials 6-7; 3rd and4th supralabials contacting orbit; preocular 1; postoculars 1-2 (usually 1); 1 large primarytemporal; nasal separated from preocular by high 2nd supralabial; mental groove present anddistinct; mental triangular, broader than long, and not partially hidden in the shallow mentalgroove; anterior chin scales large, and usually bordering mental groove; body scales weaklyimbricate anteriorly, juxtaposed posteriorly; body scales 21-28 rows at neck, 29-36 rows atmid.body; posterior ventrals distinct, mostly undivided; ventrals 275-320; small ventrals onlyslightly wider than adjacent body scales and never foliform; tail strongly compressedvertically, paddle-like; ovoviviparous. Content: Leioselasma coggeri Kharin 1984;Leioselasma cyanocinctus (Daudin 1803); Leioselasma czeblukovi Kharin 1984; Leioselasmamamillaris (Daudin 1803); Leioselasma melanocephala Gray 1849; Leioselasma pacifica(Boulenger 1896); Leioselasma semperi (Garman 1881); and, Leioselasma spiralis (Shaw1802). Content in Australia: Leioselasma coggeri Kharin, 1984; Leioselasma czeblukoviKharin, 1984; Leioselasma pacifica (Boulenger, 1896). [Note: The Asian species Leioselasmamelanocephala (Gray, 1849) is no longer considered part of the Australian herpetofauna, theprevious records of this species being based on what is now known as Leioselasma coggeriKharin, 1984].Note: As to the identity of Leioselasma striata Lacépède, B.G.E. (1804): Ann. Mus. Natl Hist.Nat. Paris 4: 184-211 [p. 198, 210] there is some confusion. The decision by McDowell (1972)to regard this species as a synonym of Hydrophis cyanocinctus Daudin, 1803 [see McDowell,S.B. (1972): Trans. Zool. Soc. Lond. 32: 189-247 [p. 215] was rejected by Cogger (1983) ascyanocinctus was not known from Australian waters. McDowell suggested that the specimenrepresented by pl. 18, figs 4-5 in Schlegel, H. (1837). (ex Merrem ms). [see Essai sur laPhysionomie des Serpens. 2. Partie descriptive. Kips and Stockum, La Haye] be made theNeotype of Leioselasma striata Lacépède,1804]. Type data: Holotype MNHP [presumed lost[Boie, F. (1827): Isis Oken 20: 508-566 (see p. 553)]. Type locality: Australia.Leioselasma coggeri Kharin, 1984Leioselasma coggeri Kharin, V.E. (1984): Zool. Zh. 63(10): 1535-1546 [p. 1538]. Type data:holotype ZIL 19681. Type locality: Port Suva [Viti Levu], Fiji.Description: Specimens in Australian waters once regarded as Hydrophis melanocephalusand Hydrophis belcheri should be referred to Leioselasma coggeri, as melanocephalusdefinitely does not occur here, and belcheri has never been positively identified in Australia,although it can be found in nearby regions. Leioselasma coggeri has a conspicuously smallhead that is not distinct from the neck. It is a rather elongate species that has the anterior partof body much more slender than the posterior, which is deep and moderately compressed.The base body colour is pale greenish to olive-grey with scattered black flecking in adults,and the head is black. There is usually a series of around 28-40 dark blackish cross-bands inadult specimens that are more or less equal in intensity in colour and width along body, with42

Australian Biodiversity Record, 2007 (8): 1-124some bands sharply tapering mid-laterally, before expanding again ventrally. The anterior ofthe venter is black due to the continuation of the transverse black banding over the ventrals.There is no pattern of secondary bands or blotches in the pale interspaces between the darkbands, but the bands tend to be less distinct in larger specimens (age?). Juveniles tend to bepale yellowish with a series of very distinct black bands, with the black head colour beingstrikingly marked with a yellowish line that forms a vague horse-shoe shape across theprefrontals, the snout, orbit, postoculars and primary temporals. The subcaudal area andposterior of the tail is black in juveniles. Some significant features of this species’ morphologyare: head shields enlarged, symmetrical; anterior part of maxilla not arched upwards, the tipof the fang projecting conspicuously below a line connecting the tips of the solid maxillaries;5-8 maxillary teeth following fang; supralabials 6-7; 3rd and 4th supralabials contacting orbit;preocular 1; postoculars 1-2 (usually 1); 1 large primary temporal; nasal separated frompreocular by high 2nd supralabial; mental groove present and distinct; mental triangular,broader than long, and not partially hidden in the shallow mental groove; anterior chin scaleslarge, and usually bordering mental groove; body scales weakly imbricate anteriorly,juxtaposed posteriorly; body scales 21-29 rows at neck, 29-36 rows at mid-body; posteriorventrals distinct, mostly undivided; ventrals 275-360; small ventrals only slightly wider thanadjacent body scales and never foliform. May attain a maximum total length of only around 1m., although a specimen 1.3 m. has been recorded. Variation in morphology suggests thatthis species may be composite.Distribution: Widely known from the Indian Ocean, South East Asia through Indonesia's TimorSea and Australia, across the Arafura Sea to New Guinea waters, and into the Coral Sea andSouth West Pacific, including Vanuatu, New Caledonia, the Loyalty Islands and Fiji. InAustralian seas, it is commonly found off the coasts of Queensland, including the outer reefsof the Coral Sea, the Northern Territory and north-western Western Australia, about AshmoreReef and Scott Reef.Habitat: Apparently inhabits a variety of tropical marine conditions ranging from the deeperwaters at the edges of coral reefs, gulf seagrass beds, mangrove flats, river deltas, coral reefsto estuaries - in relatively shallow continental shelf waters with sandy or muddy bottoms fromaround 25 to 45 metres in depth.Biology/Ecology: This is a fast-swimming species that is believed to be mainly diurnal in itsactivity, although it has also been observed at the surface during night-time. Its diet comprisesburrowing eels (Ophichthidae and Congridae) which it hunts on the sandy bottoms, by slowlysearching in the tunnels inhabited by the eels. When an eel is seized, it usually swallowedtail-first. Ovoviviparous, producing up to 8 living young in a brood.Toxicity: Although this species is reluctant to bite without some sort of serious provocation -like being seized by a human and hauled from the water - it will bite if harassed, so cautionshould be exerted. Although there are no known human fatalities arising from the bite of thisspecies, its venom is highly toxic to humans, so urgent medical attention should always besought if someone is bitten.Survival Status: Protected under the Qld Nature Conservation Act (1992), the Territory Parksand Wildlife Conservation Act (1998) and the WA Wildlife Conservation Act 1950 (asamended). Listed as a Marine Protected Species under the Australian EnvironmentalProtection and Biodiversity Conservation Act (1999). Probably not under threat, but itsapparently patchy distribution in Australian waters could potentially make it vulnerable insome parts of its range.Etymology: The name ‘coggeri’ honours Australian herpetologist Harold George Cogger.43

Australian Biodiversity Record, 2007 (8): 1-124Leioselasma czeblukovi Kharin, 1984Leioselasma czeblukovi Kharin, V.E. (1984): Zool. Zh. 63(10): 1535-1546 [p. 1542]. Typedata: holotype ZIL 19800. Type locality: Arafura Sea.Hydrophis geometricus Smith, L.A. (1986): Rec. West. Aust. Mus. 13(1): 151-153 [p. 151].Type data: holotype WAM R76484. Type locality: 84 km NW of Dampier, WA [20º10'S116º06'E] - [see Rasmussen and Smith (1997)-The taxonomic status of the sea snakesHydrophis czeblukovi (Kharin, 1984) from north-west Australian waters. Amphibia-Reptilia,18: 419-425]Description: This is a medium-sized seasnake with a very small narrow head that is notdistinct from the neck, an extremely elongate forebody and a deep, strongly compressedposterior body. The base body colour is greyish-black, with a distinctive geometrical pattern.The pattern is formed from a series of narrow white bars dorsally, which divide on the flanksto form the apices of some 34-38 dorsal hexagons and lateral pentagons; the lower flankshave pale mottling forming a more yellowish or greyish appearance on the lower ventrolateral.The tail is yellowish with a series of greyish elliptical blotches, each separated by paleyellowish or whitish barring. Some significant features of this species’ morphology are: Headshields enlarged and symmetrical; nasal cleft absent; preocular 1; postoculars 2-3;supralabials 7-8, with 3rd, 4th and 5th supralabials contacting orbit; primary temporals 1 or 2;dorsal body scales juxtaposed, each with a short keel; 31-34 body scale rows on neck; midbodyscales in 51-58 rows; ventrals smooth, about 288-304 and mostly entire, although insome occasionally divided posteriorly; ventrals wider than adjacent body scales. Usuallyattains a maximum total length of about 1.2m.Distribution: Known mainly from tropical Australian and southern New Guinea seas. It hasbeen recorded off the coast of the Northern Territory (Arafura Sea) and the Indian Ocean offnorth-western Western Australia.Habitat: Occurs in waters about 100m in depth in continental shelf areas.Biology/Ecology: This is essentially a diurnal species that feeds on a variety of small fishes.Ovoviviparous, producing about 4 young in a litter..Toxicity: This is a fast-swimming species that rarely attempts to bite under normalcircumstances, but caution should nevertheless be exerted, because its venom could behighly toxic to humans. Urgent medical attention should always be sought in the event of abite from this species, but there have been no recorded fatalities.Survival Status: Protected under the WA Wildlife Conservation Act 1950 (as amended) andthe Territory Parks and Wildlife Conservation Act (1998). Listed as a Marine ProtectedSpecies under the Australian Environmental Protection and Biodiversity Conservation Act(1999). Probably not under threat, but its limited distribution in Australian waters couldpotentially make it vulnerable.Etymology: The name ‘czeblukovi' was bestowed in honour of Vladimir P. Czeblukov, acolleague of the Russian herpetologist Vladimir Kharin.Leioselasma pacifica (Boulenger, 1896)Hydrophis pacificus Boulenger, G.A. (1896). Cat. Snakes Brit. Mus. [p. 278]. Type data:holotype BMNH 1946.1.10.14. Type locality: New Britain, Papua New Guinea.Distira macfarlani Boulenger, G.A. (1896). Cat. Snakes Brit. Mus. [p. 294, pl. 17 fig. 1]. Typedata: syntypes BMNH 1946.1.3.92-93. Type locality: Murray Ils, Torres Strait, QLD.Description: Anteriorly, this is a very elongate but robust species with a head that is relativelylarge and deep, and not distinct from the neck. The posterior of the body is even more robustwith fairly deep lateral compression. The base body colour of mature specimens is dark slaty44

Australian Biodiversity Record, 2007 (8): 1-124grey dorsally and whitish ventrally, although the ventral area of the neck may be blackish;mid-laterally, there is a zone where the darker upper, and paler lower body base colourationssmoothly merge. Patterning in adults is confined to a series (about 50-70) of dark - almostblack - transverse bands on the body, and these are broadest and darkest vertebrally (wherethey create a wide blotched pattern to the dorsum). The body bands gradually narrow andfade laterally to form continuous greyish banding to the ventrals. Sometimes the body bandsmay be split or completely separated at the vertebral line, and the banding of the neck iscontinuous with the black ventrals. The dark banding of adults tends to be also more intenseanteriorly, often with secondary bands or blotches between the main bands. The tail is oftendark with banding less pronounced, with the tip greyish, and this is a paler colour than thedarker cross-bands, when bands are present on the tail. The dorsum of the head is dark greyfleckedwith pale whitish flecks. The juvenile or immature pattern is much more pronounced,with distinctive black bands encircling the body; the anterior of the venter is black. The headand throat of juveniles is black, with pale flecking or tiny spotting on the snout, and there is ayellowish spot on each temporal area (just behind each eye). Some significant features of thisspecies’ morphology are: head shields enlarged, symmetrical; anterior part of maxilla notarched upwards, the tip of the fang projecting conspicuously below a line connecting the tipsof the solid maxillaries; 5-8 maxillary teeth following fang; nasal separated from preocular byhigh 2nd supralabial; usually the 3rd, 4th, and 5th supralabials contacting orbit; mental groovepresent and distinct; mental triangular, broader than long, and not partially hidden in theshallow mental groove; anterior chin scales large, and usually bordering mental groove; bodyscales imbricate, in 39-49 rows at mid.body; posterior ventrals distinct, mostly undivided;ventrals 320-430, small, about as wide as, or only slightly wider than adjacent body scalesand never foliform. Attains a maximum total length of around 1.5m.Distribution: Known from tropical Australian and New Guinea seas, being mainly found in theGulf of Carpentaria and Torres Strait in Queensland and the Arafura Sea near the NorthernTerritory.Habitat: This species lives mainly around coral reefs, but also found in muddy waters of riverdeltas, bays, estuaries and gulfs on continental shelf areas.Biology/Ecology: This is a fast-swimming, diurnal species, which feeds on a variety of smallfishes. Very little has been recorded on the reproductive biology of this species, oviducaldevelopment occurs in the Dry Season, so it likely that the young are born during or followingthe Wet Season. A gravid female carrying 17 developing young has been reported.Toxicity: This species rarely attempts to bite under normal circumstances, but caution shouldnevertheless be exerted, because its venom is highly toxic to humans. Urgent medicalattention should always be sought in the event of a bite from this species.Survival Status: Protected under the Qld Nature Conservation Act (1992) and the TerritoryParks and Wildlife Conservation Act (1998). Listed as a Marine Protected Species under theAustralian Environmental Protection and Biodiversity Conservation Act (1999). Probably notunder threat, but its restricted distribution in Australian waters could potentially make itvulnerable in some parts of its range.Etymology: The name ‘pacifica’ probably refers to the Pacific geographic area, but this wouldbe somewhat of a misnomer as this species is only known from the edge of the region.Genus Microcephalophis Lesson, 1834Microcephalophis Lesson, R.P. (1834). Reptiles. [Pp. 289-336]. In: Bélanger, C. (Editor):Voyage aux Indes-Orientales, par le Nord de l'Europe pendant les Années 1825, 1826, 1827,1828 et 1829. Paris: Arthus Bertrand xxxix 535 pp. [p. 320] [?non Hydrus gracilis Shaw,1802]. Type species: Microcephalophis gracilis Lesson, 1834 by monotypy. AlthoughMicrocephalophis was based on illustrations in Russell (1801), none refer to Hydrus gracilis ofShaw (1802). The determination of the Type Species was done by Smith (1926), and laterfollowed by McDowell (1972) [see Cogger (1983): Zool. Cat. Aust., 1 Amphibia Reptilia]45

Australian Biodiversity Record, 2007 (8): 1-124Liopala Gray, J.E. (1842): Monogr. Synop. Water Snakes…Hydridae. [p. 60]. Type species:Hydrus gracilis Shaw, 1802 by subsequent designation [see Gray, J.E. (1849): Cat. Spec.Snakes Coll. Brit. Mus.]Liopola Gray, J.E. (1842): Monogr. Synop. Water Snakes…Hydridae. [p. 60] [errore proLiopala Gray, 1842]Diagnosis: A genus of two species of sea snakes in the family Hydrophiidae (one of whichoccurs in Australia), and readily separated from all other genera by the following combinationof characters: Head shields large and symmetrical; valvular nostrils and lingual fossa; 3rd and4th supralabials suborbital; 3rd supralabial contacts prefrontal; nasal separated frompreocular by high 2nd supralabial; 5-6 maxillary teeth follow fang; anterior chin shields largeand bordering the mental groove; mental groove present and distinct; mental triangular,broader than long, and not partially hidden in the shallow mental groove; head very small andnot distinct from neck; elongate body-form, with the neck and anterior body extremely slender;posterior body deep and strongly compressed laterally; tail strongly compressed vertically,paddle-like; body scales juxtaposed and either possessing prominent tubercles or large keels,the lower laterals noticeably larger than the dorsals; mid-body scales in 29-37 rows in M.gracilis and 41-48 rows in M. cantoris (with females of both having a slightly higher mid-bodycount than males); ventrals small, posterior ventrals indistinct, all divided by a longitudinalfissure; ventrals about as wide as, or only slightly wider than, adjacent body scales and neverfoliform; ovoviviparous. Attains a maximum snout-vent length of only around 90 cm., although70 cm is mature, and females are larger than males. Etymology: The name Microcephalophismeans ‘very small snake head’, and is derived from ‘micro’ - very small, ‘cephale’ - head,‘ophis’ - snake. Content: Microcephalophis cantoris (Gunther, 1864); Microcephalophisgracilis (Shaw, 1802). Content in Australia: Microcephalophis gracilis (Shaw, 1802).Microcephalophis gracilis (Shaw, 1802)Hydrus gracilis Shaw, G. (1802): General Zoology, or Systematic Natural History. Pt. 2. [p.560]. Type data: holotype BMNH 1946.1.17.37. Type locality: unknown.Hydrus kadellnagam Boie, F. (1827): Isis Oken, 20: 508-566 [p. 554] [as Hydrus Kadell-Nagam]. Type data: holotype whereabouts unknown [described as pl. 13 in Russell, P. (1801)A Continuation of an Account of Indian Serpents. London: Shakespeare Press 15 pp.]. Typelocality: unknown.Hydrophis leprogaster Duméril, A.M.C. and Bibron, G. (1856): Abh. Geb. Naturw. Hamburg 3:1-78 [p. 53] [nomen nudum; introduced in synonomy].Thalassophis microcephala Schmidt, P. (1852): Abh. Geb. Naturw. Hamburg 2: 69-86 [78, pl.2]. Type data: syntypes ZMH 396, ZMH 400 [see Smith, M.A. (1926). Monograph on the SeaSnakes. (Hydrophiidae). London : British Museum xvii 130 pp.]. Type locality: coast of Java,Indonesia.Hydrophis guntherii Murray, J.A. (1884): Vert. Zool. Sind. [p. 396]. Type data: holotype BMNH1946.1.17.38. Type locality: Karachi, Pakistan.Hydrophis rostralis Smith, M.A. (1917): J. Nat. Hist. Soc. Siam 2: 340-342 [p. 340]. Type data:holotype 'The Federated Malay States Museum, Kuala Lumpur 1102' [see Smith, M.A. (1926).Monograph on the Sea Snakes. (Hydrophiidae). London : British Museum xvii 130 pp.]. Typelocality: Kuala Larut, Perak, Malaysia.Description: Although this species is rather small, it has a greatly elongate body-form, with theneck and anterior body very slender, and the posterior body deep and strongly compressedlaterally. The head is very small and not distinct from the neck. The base body colour inmature specimens is usually greyish on the dorsum and whitish ventrally. Pattern is usuallyconfined to a series of obscure pale bars over the dorsum, but the keels of the lower lateralbody are usually very dark brown or blackish giving that part of the body a dark fleckedappearance. Immature or juvenile specimens are more boldly coloured and patterned, with a46

Australian Biodiversity Record, 2007 (8): 1-124series of 40-60 whitish 'bands' on a blackish body; the head and neck is black. These bandsare most distinct and complete on the posterior, deeper part of the body, but are reduced to aseries of large oval spots on the lateral part of the very slender anterior body. As specimensage, the pattern becomes far less distinct. Some significant features of this species'morphology are: head shields enlarged, symmetrical; nasal separated from preocular by high2nd supralabial; 3rd and 4th supralabials suborbital; 5-6 maxillary teeth follow fang; mentalgroove present and distinct; mental triangular, broader than long, and not partially hidden inthe shallow mental groove; anterior chin scales large, and usually bordering mental groove;neck scales in 17-21 rows; body scales in 29-37 rows at mid-body, with females tending tohave slightly higher mid-body counts than males; body scales juxtaposed and those of thelower lateral being larger than dorsals, and either possessing prominent tubercles or largekeels, which are most prominent in males; small ventrals, about as wide as, or only slightlywider than adjacent body scales and never foliform; posterior ventrals indistinct, but alldivided by a longitudinal fissure; ventrals about 220-290 (females have a higher ventral countthan males). Variation in morphology suggests that this species may be at the very leastpolytypic. Attains a maximum total length of around 1m., with a snout-vent length of onlyaround 70-90 cm. Females grow larger than males, and are sexually mature at around 70cm.At birth this species is relatively large, at 330 mm to 380 mm in total length.Distribution: This is a widely distributed species, being found from the Persian Gulf,throughout South East Asia, across Indonesia to Australasia and well into the Coral Sea,although it does not reach New Caledonia. However, in Australian waters, this species hasonly been recorded (uncommonly) from the Torres Strait of Queensland. It is also knownrarely from the southern Great Barrier Reef part of south-east Queensland - probably asvagrants.Habitat: Although largely restricted to coral reefs in relatively shallow continental shelf watersover sandy substrates to a depth of around 30m, occasionally specimens are found inmangrove estuaries.Biology/Ecology: A fast-swimming, diurnal species that feeds on a variety of small fishes -mainly eels, which it presumably hunts by burrowing into sandy substrates. This is aovoviviparous species, that produces from 1 to 6 (average 3) young in a brood. There isconsiderable variation in the timing of reproduction across this species’ range, but it wouldappear that females are gravid during Spring and give birth in Summer. For example, in thePersian Gulf, they are gravid in Spring. In various parts of Asia, gravid females may occur inboth the inter-monsoonal period (April) as well as just after the end the summer monsoonseason (June-July). A gravid female has also been found in Australia during the early WetSeason (December).Toxicity: This snake rarely attempts to bite under normal circumstances, but caution shouldnevertheless be exerted, because its venom is highly toxic to humans. Urgent medicalattention should always be sought in the event of a bite from this species.Survival Status: Protected under the Qld Nature Conservation Act (1992). Listed as a MarineProtected Species under the Australian Environmental Protection and BiodiversityConservation Act (1999). Probably not under threat, but its restricted distribution in Australianwaters could potentially make it vulnerable in some parts of its range.Etymology: The name 'gracilis' means 'slender', and refers to the body-form of the species.Genus Polyodontognathus Wall, 1921Polyodontognathus Wall, F. (1921). Ophidia Taprobanica or the Snakes of Ceylon. [p. 374].Type species: Hydrus caerulescens Shaw, 1802 by monotypy.Diagnosis: A monotypic genus of Hydrophiid snakes readily identified by the followingcombination of characters: Head shields enlarged, symmetrical; 12 or more solid maxillaryteeth following fang; dentary teeth 22 or more; heart about midway along body; valvularnostrils and lingual fossa; nasal separated from preocular by high 2nd supralabial; 3rd and 4th47

Australian Biodiversity Record, 2007 (8): 1-124supralabial contacting orbit; anterior chin scales large, and usually bordering mental groove;mental groove present and distinct; mental triangular, broader than long, and not partiallyhidden in the shallow mental groove; body scales imbricate, strongly keeled in males and in35-54 rows at mid-body (females have a slightly higher mid-body count than males); ventralssmall, distinct, mostly undivided, barely wider than adjacent body scales and never foliform;ventrals about 253-334; males and females do not have significantly different ventral counts;tail strongly compressed vertically, paddle-like; ovoviviparous. Attains a maximum snout-ventlength of only around 0.8 m. in total length, with males being slightly larger than females.Etymology: The name Polyodontognathus is from Greek roots - ‘poly’ - many, ‘dont’ - tooth,‘gnathus’ refers to the skull, and was bestowed in recognition of the high number of maxillaryteeth in its only included species. Content: Polyodontognathus caerulescens (Shaw, 1802).Polyodontognathus caerulescens (Shaw, 1802)Hydrus caerulescens Shaw, G. (1802): General Zoology, or Systematic Natural History. [p.561] [label on holotype cites type locality as 'Indian Ocean : Vizagapatam']. Type data:holotype BMNH 1946.1.3.90. Type locality: Indian Ocean (as East-Indian) [see Smith, M.A.(1926). Monogr. Sea Snakes. (Hydrophiidae)]Polyodontes annulatus Lesson, R.P. (1834): Reptiles. [Pp. 289-336]. In: Bélanger, C. (Editor):Voyage aux Indes-Orientales, par le Nord de l'Europe [p. 325, pl. 4] [synonymy doubtful - seeSmith, M.A. (1926). Monogr. Sea Snakes. (Hydrophiidae)]. Type data: syntypes (probable)whereabouts unknown (presumed lost). Type locality: Malabar coast, India and Gulf ofMartaban, Burma.Hydrophis hybrida Schlegel, H. (1844): Abbildungen neuer oder unvollständing bekannterAmphibien [p. 115, pl. 37]. Type data: holotype RMNH 1492 [see Smith, M.A. (1926). Monogr.Sea Snakes. (Hydrophiidae)]. Type locality: Moluccas, Indonesia.Hydrophis protervus Jan, G. (1859): Rev. Mag. Zool. (2)11: 148-157 [p. 148] [see also Jan, G.(1859). Rev. Mag. Zool. (2)10 and (2)11-12, 1858 and 1859. (pl. D), as H. protervus]. Typedata: holotype (probable) MNHP [see Smith, M.A. (1926). Monogr.SeaSnakes.(Hydrophiidae)].Type locality:China Sea (?).Hydrophis frontalis Jan, G. (1863): Elenco Sistematico Degli Ofidi [see Jan, G. and Sordelli,F. (1881). Iconographie Générale des Ophidiens. Atlas (p. 110, livr. 39 pl. 5 fig. 2); as H.frontalis]. Type data: holotype RMNH 1477. Type locality: Indian Ocean.Hydrophis polydonta Jan, G. (1863): Elenco Sistematico Degli Ofidi [see also Jan, G. andSordelli, F. (1881). Iconographie Générale des Ophidiens. Atlas (p. 110, livr. 41 pl. 1); as H.polydonta]. Type data: holotype MSNM [see Smith, M.A. (1926). Monogr. Sea Snakes.(Hydrophiidae). Type locality: unknown.Hydrophis caerulescens thai Smith, M.A. (1920): J. Fed. Malay States Mus. 10(1): 1-63 [p.18]. Type data: holotype BMNH 1921.2.11.83. Type locality: Hua Hin, Gulf of Siam, Thailand.Description: This is a relatively, small, elongate sea snake with a short deep head that is notdistinct from the neck. The overall base body colour is pale blue to greyish-blue, with a blackhead and a series of 41-43 black bands on the body; the tail may be black with about 4 paleblotches, or similar to the body with another 7-9 bands. The bands over the body may becomplete, or broken about the vertebral, and so create an alternating pattern of black andblue on the dorsum. Head shields enlarged, symmetrical; 12 or more solid maxillary teethfollowing fang; dentary teeth 22 or more; heart about midway along body; nasal separatedfrom preocular by high 2nd supralabial; 3rd and 4th supralabial contacting orbit; anterior chinscales large, and usually bordering mental groove; mental groove present and distinct; mentaltriangular, broader than long, and not partially hidden in the shallow mental groove; bodyscales imbricate, strongly keeled in males and in 35-54 rows at mid-body (females have aslightly higher mid-body count than males, and Australian specimens appear to have lowermid-body counts than elsewhere); ventrals small, distinct, mostly undivided, barely wider thanadjacent body scales and never foliform; ventrals about 253-334 (although Australian48

Australian Biodiversity Record, 2007 (8): 1-124specimens appear to have lower ventral counts than those of Asia); males and females do nothave significantly different ventral counts. Attains a maximum snout-vent length of onlyaround 0.8 m. in total length, with males being slightly larger than females. Females aresexually mature at about 0.6m. in total length.Distribution: In Australian waters only known from the Gulf of Carpentaria in Queensland, andthe Northern Territory. However, this species’ total range is from southern India to China,including most of South East Asia, to about the Moluccas in eastern Indonesia, across to NewGuinea and into parts of northern Australia.Habitat: In Australia it inhabits similar relatively shallow continental shelf and gulf watersusually in about 2 m. to 20 m. depths; it apparently favours habitats fairly close to coastlinesor inshore waters. In Asia it is largely restricted to shallow mangrove flats, gulf waters, riverdeltas and estuaries.Biology/Ecology: This is a fast swimming, diurnal species that is active throughout the year.Its diet comprises small fishes, principally of the family Gobiidae, as well as eels, which arehunted along muddy bottoms of relatively shallow waters. The reproductive biology is poorlyknown, but fragmentary observations suggest that females are gravid during the intermonsoonalperiod, and give birth from between 3-13 young (usually about 6-7) prior to theonset of the monsoon season. Larger (heavier) females produce higher numbers of offspringin a litter.Toxicity: It rarely attempts to bite under normal circumstances, but caution shouldnevertheless be exerted, because its venom is highly toxic to humans. Urgent medicalattention should always be sought in the event of a bite from this species.Survival Status: Protected under the Qld Nature Conservation Act (1992) and the TerritoryParks and Wildlife Conservation Act (1998). Listed as a Marine Protected Species under theAustralian Environmental Protection and Biodiversity Conservation Act (1999). Probably notunder threat, but its restricted distribution in Australian waters could potentially make itvulnerable in some parts of its range.Etymology: The name ‘caerulescens’ refers to the bluish colouration of this species.Family LaticaudidaeThe Family Laticaudidae comprises two genera - Laticauda with 6 species - viz: Laticaudacolubrina (Schneider 1799); Laticauda crockeri Slevin 1934; Laticauda guineai Heatwole,Busack and Cogger 2005; Laticauda laticaudata (Linnaeus 1758); Laticauda saintgironsiCogger and Heatwole 2005; and Laticauda schistorhynchus (Gunther 1874) of which only twohave been recorded from Australia (as vagrants only) - plus Pseudolaticauda semifasciata(Reinwardt 1837) (which does not occur in Australia).Genus Laticauda Laurenti, 1768Laticauda Laurenti, J.N. (1768). Synopsin Reptilium. [p. 109]. Type species: Laticaudascutatus Laurenti, 1768 by subsequent designation [see Smith, M.A. (1926). Monogr. SeaSnakes. (Hydrophiidae)].Platurus Sonnini de Manoncourt, C.S. and Latreille, P.A. (1802). Histoire e Naturelle desReptiles...[p.183]. Type species: Platurus fasciatus Sonnini and Latreille, 1802 by subsequentdesignation [see Smith, M.A. (1926). Monogr. Sea Snakes. (Hydrophiidae)].Platyurus Agassiz, L. (1846). Nomenclatoris Zoologici Index Universalis. [p. 297] [emend. proPlaturus Sonnini and Latreille, 1802].Pseudolaticauda Kharin, V.E. (1984): Trans. Zool. Inst. Acad. Sci. USSR 124: 128-139 [p.134]. Type species: Platurus semifasciatus Reinwardt, 1837 by original designation.49

Australian Biodiversity Record, 2007 (8): 1-124Diagnosis: A genus of six species proteroglyphic snakes variously placed with the true seasnakes or as a separate family, but nevertheless closely related to the Asian ‘Elapidae’, andreadily identified by the following combination of characters: Head shields large andsymmetrical; maxillary extends forward, beyond the palatine; nostrils laterally positioned;nasals separated by internasals; infralabials partly fragmented; body scales smooth andimbricate; ventral scales much broader than wide, being about half as wide as body; analdivided; subcaudals divided; tail strongly compressed vertically, paddle-like; oviparous.Etymology: The name Laticauda means ‘broad tail’. Content: Content in Australia: Laticaudacolubrina (Schneider, 1799); Laticauda laticaudata (Linnaeus, 1758).Laticauda colubrina (Schneider, 1799)Hydrus colubrinus Schneider, J.G. (1799). Historiae Amphibiorum Naturalis et Literariae. [p.238]. Type data: holotype ZMB 9078. Type locality: unknown [see Smith, M.A. (1926).Monogr. Sea Snakes. (Hydrophiidae)].Coluber platycaudatus Oken, L. (1836). Allgemeine Naturgeschichte. Amphibien [p. 566][unnecessary replacement name for Hydrus colubrinus Schneider, 1799].Description: A large, robust, and round-bodied snake, with a short deep head not distinct fromthe neck, and a laterally compressed tail barely higher than the body. The base body colour iscyan blue to dark bluish-grey, with a prominent pattern of 20-65 broad black transverse bandsthat encircle the body. The venter is cream or yellow between the dark bands. The dorsal partof the head has a large black patch extending from about the prefrontals and over theparietals, and extending down behind the temporals to the throat. The snout is yellowishextending to about the internasals with blackish smudging, then brighter yellow over thesupraoculars to the temporals stopping at the downward extension of the black head patch.One the side of the head, there is a thin black line running from the snout through the eye, tomeet with the temporal extension of the black head patch also. The labials are yellowish -hence the common name of Yellow-lipped Sea Krait. Head shields large and symmetrical;maxillary extends forward, beyond the palatine; nostrils laterally positioned; nasals separatedby internasals; prefrontals separated by large azygous shield; body scales smooth andimbricate; mid-body scales in 21-25 rows; ventral scales much broader than wide, beingabout half as wide as body; ventrals 210-250; anal divided; subcaudals divided (25-35 infemales, 36-50 in males). This species has been redefined in the recent revisions by Coggerand Heatwole (2006) and Heatwole, Busack and Cogger (2005). Attains a maximum totallength of around 1.8 m., and a snout-vent length of about 1.6 m., with females being largerthan males. Males are about one third smaller than females, and only weigh about half theweight of females. The record size for this species is reputedly 3.6 m. in total length, but thiswould be exceptional.Distribution: Widely known from the waters of South East Asia, through Indonesia to NewGuinea and throughout much of the South West Pacific Ocean, to as far south as NewZealand as vagrants. In the Australian region it is known mainly from tropical and sub-tropicalseas. Occurrence in all Australian waters are however, regarded as vagrant, as there are noknown resident or breeding populations. It has on occasions been reported as part of thefauna of Queensland (mainly Torres Strait), and the Northern Territory, and on extremely rareoccasions for New South Wales, Victoria and Tasmania as well, but the evidence isinconclusive, because the few records are rare or very old. It is particularly common in partsof the South West Pacific and its islands.Habitat: Known to forage around tropical coral reefs in shallow seas, amongst estuaries andmangrove swamps, as well as in the open sea. It may be also found active in the sub-littoral,littoral, supralittoral zones, as well as on beaches, rocky islets, exposed reefs and theshorelines of major islands. Permanent populations are not known from continental landmasses.Biology/Ecology: A diurnal, nocturnal or crepuscular species, that is both aquatic andterrestrial in habits (truly amphibious). In the islands of the South West Pacific Ocean, thisspecies is also found on the land along the shoreline, or even well away from the water -50

Australian Biodiversity Record, 2007 (8): 1-124sometimes many hundreds of metres from the sea. At any time it would appear that only asmall percentage of the population is actually on land at one time, with approximately 75%being at sea and 25% on land. Specimens may stay at sea for a few hours, but many snakesforage for days, and some weeks before returning to land to digest prey. Most specimensfound on land are males. Most movements onto the land appear to coincide with high-tideconditions at night, although specimens have also been observed apparently basking onexposed reefs and amongst coral debris on reefs left exposed at low tide. At certain times ofthe year this species may form large aggregations on land presumably for breeding, andmales actively seek out females as they come ashore, for mating. It is also possible that largeaggregations may form for migration between breeding areas, and there appears to be somekind of homing behaviour practiced also. On suitable islands and basking sites, largenumbers may co-exist - large dens of over 100 individuals can be formed, with severalspecimens sharing quite small pieces of cover. This aggregative behaviour althoughobviously related to mating, is probably a response to the high diurnal temperatures on theirisland homes, so all available cover would be needed for thermoregulation. This speciesfeeds almost exclusively on eels, which are captured on reefs in shallow water at night.Females and males tend to eat different species of eels in different parts of the marineenvironment. The larger females spend longer periods at sea, forage in deeper water at theedges of fringing reefs, and eat mainly the larger-headed Conger Eels. Males, being muchsmaller, tend to forage in the shallow water near-shore reefs, and eat the small-headed MorayEels. Generally, most food items taken are quite large when compared to the size of thesnake, being about 20-40% of the snake’s mass. When foraging, the snakes usually quicklyreturn to the surface every few minutes, but if necessary, they can remain submerged for upto an hour at times. At dawn, specimens come ashore to presumably rest and digest theirprey, before returning again to the sea at dusk. When on land they shelter in crevices, cavesas well as beneath ground vegetation and debris, such as dried coconut fronds on theground. Unlike the “true” sea snakes (Hydrophiidae), all species of Laticauda are oviparous.They reproduce seasonally in more temperate parts of their range, but more often in tropicalareas. They lay up to 20 eggs (usually about 4-8 eggs) in a clutch on land above the high tidemark, and are thought to secrete their eggs deep in rock crevices or even in limestone‘grottos’ beneath the island’s surface. The eggs hatch after an incubation lasting about 2months. The only confirmed predators on the adults are White-breasted Sea Eagles, althoughcrabs and larger predatory fishes would prey on juveniles. In parts of Asia, this species isheavily exploited by humans for its skin.Toxicity: This is an inoffensive species that only very rarely attempts to bite, but cautionshould nevertheless be exerted, because is venom is highly toxic to humans. Urgent medicalattention should always be sought in the event of a bite from this species.Survival Status: Protected under the New South Wales National Parks and Wildlife Act (1974)but not listed in that State as a Threatened Species in any of the Schedules of the NSWThreatened Species Conservation Act (1995). Also protected under the Qld NatureConservation Act (1992) and the Territory Parks and Wildlife Conservation Act (1998). Listedas a Marine Protected Species under the Australian Environmental Protection andBiodiversity Conservation Act (1999). Probably not under threat, but its restricted distributionin Australian waters could potentially make it vulnerable in some parts of its range. Thisspecies however, has been reported to be abundant in the south-west pacific and parts ofAsia but is seldom seen in significant numbers in Australian waters.Etymology: The name ‘colubrina’ is Latin for serpent.Laticauda laticaudata (Linnaeus, 1758)Coluber laticaudata (part.) Linnaeus, C. (1758): Classis III. Amphibia. Systema Naturae [p.222]. Type data: syntypes NHRM one of two extant syntypes. Type locality: 'Indiis'.Laticauda scutata Laurenti, J.N. (1768): Synopsin Reptilium.[p. 109]. Type data: holotype (pl.16 in Linnaeus, C. (1754). Museum S.R.M. Adolphis Friderici Regis…]. Type locality: 'Indiis'.51

Australian Biodiversity Record, 2007 (8): 1-124Platurus laurenti Rafinesque, C.S. (1817): Am. Month. Mag. 1: 431-436 [432] [non Platuruslaurenti Daudin, 1803. Note : Smith, M.A. (1926). Monogr. Sea Snakes. (Hydrophiidae).regards P. laurenti as a substitute name (??)]. Type data: type status unknown. Type locality:unknown.Platurus fischeri Jan, G. (1859): Rev. Mag. Zool. (2)11: 148-157 [p. 149]. Type data: syntypesMSNM* (not found), MNHP* (not found). Type locality: Indian Ocean.P[laturus] affinis Anderson, J. (1871): Proc. Zool. Soc. Lond. 1871: 149-211 [p. 190]. Typedata: holotype IM 8289 (see Smith, M.A. (1926). Monogr. Sea Snakes. (Hydrophiidae)]. Typelocality: Tolly's Nullah (as Fallah's Nullah), a tidal stream, Calcutta India.Platurus muelleri Boulenger, G.A. (1896). Catal. Snakes Brit. Mus. [p. 309] [ex Müller MS].Type data: type status unknown* (not found). Type locality: South Pacific.Description: Similar in most respects to the preceding sea snake, this species althoughrobust, and round-bodied, doesn’t reach anywhere near the size of L. colubrina. The basebody colour is bright blue to dark bluish-grey, with a prominent pattern of 25-70 broad blacktransverse bands that do not generally encircle the body, as most or all of the bands areincomplete on the venter. The venter is creamish or yellowish. The dorsal part of the head isblack and the snout is yellowish or bluish, extending to about the internasals and over thesupraoculars. The labials are brownish. Some significant features of this species’ morphologyare: Head shields large and symmetrical; maxillary extends forward, beyond the palatine;infralabials partly fragmented; nostrils laterally positioned; nasals separated by internasals;prefrontals not separated by large azygous shield; body scales smooth and imbricate; neckscale rows 19; mid-body scale rows 19; ventral scales much broader than wide, being abouthalf as wide as body; ventrals 225-243; anal divided; subcaudals 25-50, divided (males havemore subcaudals than females). Attains a maximum snout-vent length of about 1.1 m., withthe average being only around 80cm in total length (females are larger than males).Distribution: In Australian waters being mainly found off the coasts of Queensland, and theNorthern Territory, and on occasions off New South Wales as well. Occurrences in Australianwaters however, are regarded as vagrant, as there are no known resident or breedingpopulations and most records are either old or lacking verification (no specimen-basedrecords). Widely known elsewhere from the waters of South East Asia - including India,Thailand, Malaysia, Taiwan, China and Japan - through Indonesia and the Philippines to NewGuinea and throughout much of the South West Pacific Ocean. In our region, known mainlyfrom tropical and sub-tropical seas, in the South West Pacific and its islands, to about NewCaledonia.Habitat: Known to live around tropical coral reefs in shallow seas, amongst estuaries andmangrove swamps, as well as open sea. In the Pacific islands, this species also lives on theland along the shoreline, sometimes being found well inland from the sea.Biology/Ecology: A diurnal, nocturnal or crepuscular species, that is both aquatic andterrestrial in habits. Oviparous, laying around 7 eggs in a clutch on land above the high tidemark, usually in sand under fallen palm fronds and other litter. It feeds on a variety of smallfishes.Toxicity: This is an inoffensive species that is extremely reluctant to bite, but caution shouldnevertheless be exerted, because its venom is highly toxic to humans. Urgent medicalattention should always be sought in the event of a bite from this species.Survival Status: Protected under the Qld Nature Conservation Act (1992) and the TerritoryParks and Wildlife Conservation Act (1998). Listed as a Marine Protected Species under theAustralian Environmental Protection and Biodiversity Conservation Act (1999). Probably notunder threat, but its restricted distribution in Australian waters could potentially make itvulnerable in some parts of its range.Etymology: The name ‘laticaudata’ means ‘broad tail’.52

Australian Biodiversity Record, 2007 (8): 1-124Family PelamiidaeI include within the family Pelamiidae the genera Kolpophis, Lapemis, Pelamis andPraescutata. Kolpophis annandalei (Laidlaw, 1901) and Praescutata anomalus (Schmidt,1852) do not occur in the Australian region and so are not discussed below.Genus Lapemis Gray, 1835Lapemis Gray, J.E. (1835): Illustrations of Indian Zoology [pl. 87 fig. 2]. Type species:Lapemis hardwickii Gray, 1835 by monotypy.Diagnosis: A genus of small to moderate-sized Hydrophiid snakes with a large deep head notdistinct from the thick neck, and robust body-form. Readily distinguished from all other generaby the following combination of characters: head shields enlarged, symmetrical; 3-6 maxillaryteeth following fang; valvular nostrils and lingual fossa; preocular 1; postoculars 1 or 2;primary temporals 2 or 3; rostral normal, triangular, broader than high; supralabials 7-8; 3rdand 4th supralabials contacting orbit; nasal separated from preocular by high 2nd supralabial,or a separate scale that results from a horizontal split in the 2nd supralabial (not a loreal); adistinct mental groove; anterior chin shields reduced, in only narrow contact with mentalgroove, and separated by first infralabials; mental normal, triangular, broader than long, andnot partially hidden in the shallow mental groove; body scales juxtaposed and hexagonal orsquarish in shape; lower laterals much larger than dorsals and with enlarged tubercles orspines in males; mid-body scales in 23-45 rows; ventrals not distinct, no larger than adjacentbody scales, about 110-240 (females have a higher ventral count than males); tail stronglycompressed vertically, paddle-like; ovoviviparous. Etymology: The name ‘Lapemis’ is derivedfrom an anagram of ‘Pelamis’. Content: Lapemis curtus (Shaw, 1802) (cf. Smith, 1926);Lapemis hardwickii Gray, 1835.Lapemis curtus (Shaw, 1802)*Hydrus curtus Shaw, G. (1802): General Zoology, or Systematic Natural History. [p. 562].Type data: holotype BMNH III.2.1.a. Type locality: provenance unknown.Hydrophis pelamidoides Schlegel, H. (1837) : Essai sur la Physionomie des Serpens. [p. 521,pl. 18 figs 16-17]. Type data: type status unknown. Type locality: unknown.Hydrophis propinquus Jan, G. and Sordelli, F. (1872): Iconographie Générale des Ophidiens[Pp. 39-42].Hydrophis pelamidoides unimaculatus Peters, W. (1876): Mber. K. Preuss. Akad. Wiss. Berl.1876: 528-535 [1877 on title page].Lapemis hardwickii Gray, J.E. (1835): Illustrations of Indian Zoology [pl. 87]. Type data:holotype BMNH 1946.1.18.39. Type locality: India.Lapemis loreatus Gray, J.E. (1843): Ann. Mag. Nat. Hist. 11: 46 [p. 46]. Type data: holotypeBMNH 1946.1.17.48. Type locality: ? Borneo [see Smith, M.A. (1926). Monogr. Sea Snakes.(Hydrophiidae)].Hydrophis (Pelamis) pelamidoides annulata Fischer, J.G. (1856): Abh. Geb. Naturw.Hamburg, 3: 1-78 [67, pl. 3]. Type data: syntypes ZMH 419-420 [see Smith, M.A. (1926).Monogr. Sea Snakes. (Hydrophiidae)]. Type locality: Java, Indonesia.Hydrophis problematicus Jan, G. (1859): Rev. Mag. Zool. (2)11: 148-157 [p. 150] [as H.problematicus]. Type data: syntypes MNHP (not found), MSNM (not found). Type locality:Manila, Philippines.Hydrophis abbreviatus Jan, G. (1863): Elenco Sistematico Degli Ofidi Descritti e Disegnati perl'Iconografia Generale. [see also Jan, G. (1859). Rev. Mag. Zool. (2)10 and (2)11-12, 1858and 1859. (p. 109, pl. d); as H. abbreviatus]. Type data: holotype MCZ 20647. Type locality:Manila, Philippines.53

Australian Biodiversity Record, 2007 (8): 1-124Hydrophis brevis Jan, G. (1863): Elenco Sistematico Degli Ofidi Descritti e Disegnati perl'Iconografia Generale. [p. 109] [as H. brevis]. Type data: holotype MSNM not found. Typelocality: Manila, Philippines.Hydrophis fayeriana Anderson, J. (1871): Proc. Zool. Soc. Lond. 1871: 149-211.Hydrophis pelamoides Hilgendorf, F. (1876): Mitt. Dt. Ges. Nat.-U. Volkerk. Ostasiens, 1: 29-34 [31] [? lapsus for Hydrophis pelamidoides Schlegel, 1837].*It is presently believed that Lapemis hardwickii of Asia is synonymous with Lapemis curtus ofAustralasia. Therefore the following description includes the characteristics of both species,although I am personally not entirely convinced that we are dealing with only a single variablespecies in the genus Lapemis.Description: This is a small to moderate-sized sea snake with a large deep head not distinctfrom the thick neck, and a robust body-form Mature specimens have a fairly plain overallcolouration with either shades of orange-brown, olive-grey, or pale greenish dorsally and acreamish to yellowish venter. There is an irregular line of demarcation between these coloursabout mid-laterally, and often this takes the form of a wavy or zig-zag line which is an aftereffect of the loss of the juvenile pattern. Juveniles and occasionally some young adults aremore brightly coloured with a series of about 30 to 55 small dark grey brown band-likeblotches that run down the back of the body to the tail; in most juvenile specimens theblotches are quite distinct extending about mid-laterally where smoothly taper to a roundedtriangular shape. Additionally, some specimens may also have a small black spot or splotch inthe pale interspace between the large blotches, while in others the large blotches maycoalesce at the vertebral somewhat to form a dark wavy pattern dorsally. This immaturepattern is usually lost with the onset of maturity. Some of the distinctive morphologicalcharacters are: head shields enlarged, symmetrical; preocular 1; postoculars 1 or 2; primarytemporals 2 or 3; rostral normal, triangular, broader than high; supralabials 7-8; 3rd and 4thsupralabials contacting orbit; nasal separated from preocular by high 2nd supralabial, or aseparate scale that results from a horizontal split in the 2nd supralabial (not a loreal); adistinct mental groove; parietals entire or partly fragmented; anterior chin shields reduced, inonly narrow contact with mental groove, and separated by first infralabials; mental normal,triangular, broader than long, and not partially hidden in the shallow mental groove; bodyscales juxtaposed and hexagonal or squarish in shape; lower laterals much larger thandorsals and with well-developed (enlarged) tubercles, spines or keels in males (and also insome females); body scales around neck in 23-35 rows; mid-body scales in 23-45 rows (withfemales tending to have slightly higher mid-body counts than males); ventrals not distinct, nolarger than adjacent body scales, about 110-240 (females have a higher ventral count thanmales, and Australian populations may have slightly more ventrals than populations ofLapemis hardwickii in Asia). This species attains a maximum snout-vent length of only around85 cm. (about 1m. in total length). Females are about the same size as males, or at best onlymarginally larger in total length, but significantly larger than males in snout-vent length.Females also appear to be slightly bulkier than males as well. The largest specimen knownfrom Australian waters was about 1.3m. Females can be sexually mature at only around0.45m SVL, although this would appear exceptional, with mature females of around 0.7m SVLbeing more usual. The young are relatively large at birth when compared to the mother’slength across the species range. Juveniles are nearly half the mother’s length at around 33cm- 35cm in snout-vent length when born in the Asian region, but appear to be slightly smaller inlength than those from Indonesia. Interestingly, mature Asian specimens (Lapemis hardwickii)are smaller than those from Australia, seldom exceeding 0.85m. in length and they alsoappear to have somewhat less bulk than Australian specimens (Lapemis curtus?). Materialfrom Australia often reaches around 1.0m. to 1.3m in length and are much heavier in build aswell. Given that bulk and length appear to developmentally determined in this species (seeSmith, 1926) this would appear to negate an ecological explanation for this variation.Distribution: In Australian waters, this species mainly occurs in tropical areas, being regularlyfound off the coasts of Queensland, Northern Territory and Western Australia. It is widelyknown elsewhere from the Persian Gulf, and much of South East Asia, including Myanmar,54

Australian Biodiversity Record, 2007 (8): 1-124Pakistan, India, Sri Lanka, Thailand, Malaysia, Vietnam, China, Taiwan, and Japan. It rangesacross most of the Philippines and Indonesia, through to Australia and New Guinea to aboutas far east as the Coral Sea and New Caledonia.Habitat: Essentially lives on continental shelf habitats in water depths varying from around4m. to 55m and with a variety of substrate types. Occurs around coral reefs with sandybottoms in water depths as shallow as a few metres, up to around 25m. It is also an inhabitantof the turbid waters of river deltas, bays, estuaries and gulfs with muddy and/or sandybottoms. This species may be also found in the tidal stretches of tropical rivers, and can occurmany kilometres upstream from the sea, however it avoids freshwater conditions as suchwater can be fatal to its metabolism. Also known from the open sea, where it may occur inlarge numbers along current lines.Biology/Ecology: This is a relatively bulky, slow-swimming diurnal species that feeds on agreat variety of small fishes; it will also take small squid. As would be expected with a speciesoccurring across a wide geographic range, its reproductive conditions may vary dependingupon the location, but it would appear that development occurs during the Dry Season(winter), with births occurring during the Wet Season (summer). Mature females of Lapemishardwickii from the Persian Gulf and Asia appear to reproduce every year, with up to 7(usually about 2 to 4) young born in a brood. In Australia the brood size varies from 1 to 15(usually about 4 to 9). This would appear to be somewhat higher than brood sizes in Asianpopulations of Lapemis hardwickii. Similarly, larger females tend to produce more young in abrood in the Australian population, but brood-size and female size appears to be independentin some Asian populations of Lapemis hardwickii, but not in others. Despite its abundancevery little of this species’ habits in the wild is well understood. For instance it has beenobserved to gather in huge numbers along current lines on the open sea and such slicks ofsea snakes appear to extend for kilometres at times. Similarly, in a local context, smallgatherings of several individuals can be occasionally observed entwined together in a ball onthe surface of the sea - a puzzling behaviour that has also been observed in other species ofsea snakes as well. This species may also harbour small colonies of bryozoans, barnacles,small bivalves and hydrozoans attached to its skin, as well as a range of internal parasitictrematode and other unidentified worms associated with the skin and digestive system, soperhaps this balling behaviour is a response to the presence of attached bryozoans and thelike. Other Hydrophiids have been observed twisting themselves into knots, so as to scrapeoff organisms by abrading the body against itself as it crawls through such a knot; severalspecimens might collaborate in such behaviour as so form one of these sea snake balls.Toxicity: It may readily attempt to bite if harassed, so caution should be exerted whenhandling this species. Its venom has been the subject of considerable research and is knownto be highly toxic to humans. Urgent medical attention should always be sought in the eventof a bite from this species as there have been a number of human fatalities recorded from itsbite.Survival Status: Protected under the Qld Nature Conservation Act (1992), the Territory Parksand Wildlife Conservation Act (1998) and the WA Wildlife Conservation Act 1950 (asamended). Listed as a Marine Protected Species under the Australian EnvironmentalProtection and Biodiversity Conservation Act (1999). In Asia vast numbers are harvested forthe leather trade and this is undoubtedly having a serious impact on populations. Probably notunder threat overall at this stage as it is among the most abundant of all species of seasnakes, but its restricted distribution in Australian waters could potentially make it vulnerablein some parts of its range here given its rate of accidental capture in trawling operations.Etymology: The name ‘curtus’ is from the Latin, meaning ‘short’; the name 'hardwickii'honours the British naturalist General Hardwicke who collected the Type Specimen.Further Notes: There is some confusion about the taxonomic status of Lapemis hardwickii,with recent opinion favouring the submergence of this species with Lapemis curtus (Shaw,1802) due to the apparent unreliability of the fragmented vs unfragmented parietal conditionsthat have been traditionally used to separate these species (see references). Given that thedifferent ventral scale conditions, as well as numbers of mid-body scale rows vs number of55

Australian Biodiversity Record, 2007 (8): 1-124ventrals between the two taxa appear to have some significant geographical basis (i.e.developmentally derived), I am reluctant at this time to support the belief that there is only asingle highly variable species of Lapemis. I consider that the Australian population is in partreferrable to Lapemis curtus (Shaw, 1802) (cf. Smith, 1926), but it is not at all inconceivablethat Lapemis hardwickii Gray, 1835 could also occur in the Indian Ocean territory of Australia.In my opinion Lapemis needs to be thoroughly studied across its entire range, before aconvincing case for a single polymorphic species can be made.Genus Pelamis Daudin, 1803Pelamis Daudin, F.M. (1803). Histoire Naturelle….des Reptiles. [p. 357]. Type species:Pelamis bicolor Schneider, 1799 by subsequent designation, see Gray, J.E. (1825): AnnalsPhilos. (2)10: 193-217.Pelamydrus Stejneger, L. (1910): Proc. R. Soc. Vic. 38: 91-114 [p. 111]. Type species: Anguisplatura Linnaeus, 1766 by original designation.Diagnosis: As presently defined, a monotypic genus of sea snakes in the family Pelamiidaewith a pan-global distribution. The head is rather long and deep, the body-form is long andhigh (strongly compressed laterally) but robust. It can be immediately separated from all othergenera by the following combination of characters: Head shields enlarged, symmetrical; 7-11maxillary teeth following fang; valvular nostrils and lingual fossa; preoculars 1-2 (mostly 1);postoculars 2-3 (mostly 2); primary temporals 2 or 3; supralabials 6-8 (usually 7 or 8); 3rdsupralabial contacting preocular; no distinct mental groove; body scales small andjuxtaposed, either triangular or hexagonal in shape, and in 47-69 rows at mid-body; bodyscales at the ventro-lateral margin usually with 2 or 3 small tubercles; ventrals about 240 to406, small (barely discernable), usually divided and about as wide as, or just wider than,adjacent body scales; tail strongly compressed vertically, paddle-like; diploid karyotype 38 (20macrochromosomes and 18 microchromosomes); ovoviviparous. Etymology: The namePelamis is derived from the Greek ‘pelamys’ for fish. Content: Pelamis platurus (Linnaeus,1766).Pelamis platurus (Linnaeus, 1766)Anguis platura Linnaeus, C. (1766): Systema Naturae [p. 391]. Type data: type statusunknown presumed lost. Type locality: unknown.Hydrus bicolor Schneider, J.G. (1799): Historiae Amphibiorum Naturalis et Literariae. [p. 242].Type data: type status unknown. Type locality: Tahitian seas.Pelamis schneideri Rafinesque, C.S. (1817): Am. Month. Mag. 1: 431-436 [p. 432][unnecessary replacement name for Hydrus bicolor Schneider, 1799].Hydrophis pelamis Schlegel, H. (1837): Essai sur la Physionomie des Serpens. [p. 508, pl. 18figs 13-15] [as H. Pelamis]. Type data: type status unknown. Type locality: unknown.Pelamis ornata Gray, J.E. (1842): Monogr. Synopsis Water Snakes…Hydridae. [p. 60]. Typedata: holotype BMNH 1946.1.17.51. Type locality: Borneo (as India) [see Smith, M.A. (1926).Monogr. Sea Snakes. (Hydrophiidae)]Pelamis variegata Duméril, A.M.C., Bibron, G. and Duméril, A. (1854): Erpétologie Générale[p. 1337]. Type data: holotype MNHP 3976. Type locality: Macassar, Celebes Ils, Indonesia.Pelamis bicolor sinuata Duméril, A.M.C., Bibron, G. and Duméril, A. (1854): ErpétologieGénérale [p. 1338]. Type data: syntypes (probable) MNHP 8683 5 specimens. Type locality:unspecified locality.Hydrophis (Pelamis) bicolor alternans Fischer, J.G. (1856): Abh. Geb. Naturw. Hamburg 3: 1-78 [63] [unnecessary replacement name for Pelamis variegata Duméril, Bibron and Duméril,1854].56

Australian Biodiversity Record, 2007 (8): 1-124Hydrophis bicolor maculata Jan, G. (1863). Elenco Sistematico Degli Ofidi. [p. 109] [?nomennudum].Hydrophis bicolor maculata Jan, G. and Sordelli, F. (1872). Iconographie Ophidiens. Atlas.[livr. 40, pl. 3 fig. 1]. Type data: syntypes (probable) MNHP 7710, MNHP 3975. Type locality:Indian Ocean.Description: Probably the most widespread as well as one of the most beautiful species ofsea snakes. This species can be immediately recognized by its bold colour pattern of blackdorsum and golden yellow venter - although there is considerable variation in colour andpatterning both between and within the different ‘populations’ across its range. In mostspecimens there is a clear mid-lateral line of demarcation between the dorsal and ventralcolour anteriorly, but on the posterior of the body and the tail, this line may become verywavy. The head is uniform black with the labials yellowish. The dorsal colouration varies fromone individual to another with some being purplish black, or uniform black, or even darkbrown. The ventral colour is more often yellow, but some can be brownish throughout theventer and this extends onto the lower lateral zone where it is separated from the darkdorsum colour by a narrow yellow line mid-laterally. The tail is a similar yellow to the venter,with the addition of a black barring pattern, and a lower ventro-lateral row dark spots orblotches - which may coalesce into a longer pattern in places. This unique colouration andpatterning suits a lay-and-wait predator, as the lighter yellowish ventral colour and darkbluish-black dorsum provides excellent camouflage amongst the yellowish-green and darkblue-black of drifting seaweeds that it lives amongst. Some of the more distinctive features ofthis species’ morphology are as follows: The head is rather long and deep, the body-form islong but robust, and is strongly compressed laterally. It can be immediately separated from allother genera by the following combination of characters: Head shields enlarged, symmetrical;preoculars 1-2 (mostly 1); postoculars 2-3 (mostly 2); primary temporals 2 or 3; supralabials6-8 (usually 7 or 8); 3rd supralabial contacting preocular; no distinct mental groove; bodyscales small and juxtaposed, either triangular or hexagonal in shape; body scales in 27-34rows at neck; body scales in 47-69 rows at mid-body (with females having a slightly highermid-body count than males); body scales at the ventro-lateral margin usually with 2 or 3 smalltubercles in mature specimens, and these appear to be slightly more prominent in males;ventrals about 240 to 406 (females have a higher ventral count than males); ventrals small(barely discernable from adjacent body scales), usually divided and about as wide as, or justwider than, adjacent body scales. There is a positive correlation between the number of midbodyscale rows and the number of ventrals that appears to have a developmental rather thanan ecological basis. It reaches a maximum size of about 0.85m. in total length, althoughspecimens around 1.1 metre in total length are known. Females tend to have a greater svlthan males, are sexually mature at around 63cm svl. Australian specimens also appear toreach a larger size than populations elsewhere in the world and when one also considers thedistinctive reproductive pattern in Australian waters, this may indicate that Pelamis platuruscould comprise more than one species. During the last 240 years since its originaldescription, a number of synonyms have been described, and numerous studies on otheraspects of its biology have appeared, but no adequate taxonomic revision has beenundertaken on the species across its entire range. A global-wide genetic and morphologicalstudy could reveal surprises that may enrich our understanding of the evolutionary history ofthis species as well as its relations with other proteroglyphic snakes. Research on thetaxonomy of Pelamis could also reveal new insights into other fields such as ecology,physiology and toxicology for instance that may have broader implications for ourunderstanding and conservation of marine ecosystems and their functioning. The taxonomy ofPelamis platurus would be an ideal project for an international research collaborationinvolving various universities and museums.Distribution: This species has the largest distribution of any sea snake, mainly occurringthroughout tropical and sub-tropical parts of both the Indian and Pacific Oceans, but not in theAtlantic Ocean. In Australia, it occurs widely throughout tropical, sub-tropical and onoccasions even in temperate waters, being found off the coasts of Queensland, NorthernTerritory and Western Australia, and on frequent occasions in New South Wales; it has beenonly rarely observed in the waters of Victoria and Tasmania. The species’ occurrence in57

Australian Biodiversity Record, 2007 (8): 1-124temperate waters through drift in major currents is thought to result in it temporarily occupyingnon-feeding and non-reproductive habitats such as southern Australia, Africa and NewZealand, but observations also suggest the possibility that this species can have a long-termpresence in such southern waters. At the broader global scale, it ranges from the southernCape of Africa throughout the Indian Ocean to India and across most of Asia and Australasia.It is widespread throughout the Pacific Ocean and although it occurs in both hemispheres, it isapparently more successful south of the Equator. In the Northern Hemisphere it has beenrecorded as far north as Possiet Bay, Russia in the western Pacific Ocean. The distributionextends right across to the western coasts of the Americas where it has been recorded fromabout as far north as California in the USA, and south to about Ecuador. In the south Pacific itoccurs in the Solomon Islands, Vanuatu, New Caledonia, Fiji and numerous other reefs andisland territories and states, to as far south as New Zealand.Habitat: This is a pelagic, or surface-dwelling species, well-known from open seas andoceans in tropical, subtropical and even temperate areas. It drifts worldwide amongst theflotsam of debris and seaweeds that gather along major current lines and most wouldprobably spend all of their lives on the open sea. Such drifts represent a unique marinehabitat for fishes and other marine life which utilise the debris fields as shelter and as asource of pelagic micro-organisms. Generally their presence in continental shelf waters isunusual, and probably the result of storms or unusual currents in their primary open-seahabitat.Biology/Ecology: Essentially a nocturnal species that drifts beneath or amongst flotsam suchas drifting seaweed where they lay in wait for surface-dwelling fishes. Although they tend tobe far less active than other sea snakes (indeed, they can lay motionless on the surface forhours at a time) for the most part they just go with the flow unless they are disturbed. Ifnecessary, they swim quite quickly by powerful lateral undulations and can also swim inreverse if necessary. Pelamis are essentially float-and-wait foragers, ambushing a variety ofsmall pelagic fishes by swimming backwards to seize fish that have been unknowingly usingthe snakes’ body as shelter. They will also actively chase fishes and seize them either headonor by a quick sideways lunge. In captivity they will eat fish that alive or dead, as well aspieces of fish and a captive Pelamis can get quite excited during feeding, rapidly lunging andbiting even other snakes in the frenzy to secure a fish; frogs have also been taken by captivespecimens as well. Specimens have lived just over 2 years in captivity, but their lifespan isunknown. The reproductive biology of Pelamis is poorly known, and varies somewhat acrossits range. It produces from 2 to 6 young in a brood (usually about 3), following a 6-8 monthgestation period. The size of young at birth range from 220mm. to 280mm in total length, andcan feed on small fish on the same day they are born. Apparently the number of offspring isnot related to female body-length. In Australian waters (New South Wales) it reproducesduring the winter months (May-July) in contrast to the pattern elsewhere over the species’range. Whereas near Costa Rica in the Eastern Pacific gravid females are found during theDry Season (December-April), with births appearing to occur around March (in nearbyPanama). In Asia (Sri Lanka) gravid females have been found in March (during the intermonsoonperiod). Off Africa in the Indian Ocean gravid females have been recorded from lateAutumn through Winter to Spring (May to October). Huge populations can accumulate inareas of converging currents, and on occasions thousands of snakes have been observedsheltering amongst the flotsam over many kilometres of drift. This species can also beoccasionally found stranded on beaches following periods of strong onshore winds. As suchlarge populations of Yellow-bellied Sea Snakes drift with flotsam in the major sea currents, itcan be expected that many are washed ashore during severe storms. When such eventsoccur, they appear to be able to temporarily seek shelter on the land in stormy weather, andon occasions can be found sheltering under debris above the high tide mark along beachesand mangroves. Their capacity for temporary survival out of water is in marked contrast tomost other sea snakes where it can be very quickly fatal. This probably indicates that thelower pressure of surface-dwelling or pelagic habits of Pelamis have resulted in a blood-flowphysiology less sensitive to otherwise hazardous terrestrial air pressure conditions. However,their extreme laterally-compressed body-form makes movement land extremely difficult, withthe snake being forced to crawl on its sides (laterally rather than ventrally). One of thepossible reasons for this species’ abundance could be its toxic flesh - which would make itpotentially unpalatable or even dangerous to most carnivores - although sharks, some large58

Australian Biodiversity Record, 2007 (8): 1-124fish and even the Leopard Seal have been rarely recorded eating this species. A number ofinternal parasites have been recorded (nematodes and cestodes) but their effects on thisspecies (if any) are unknown. Epizoans such as bryozoans and barnacles are occasionallyfound attached to the skin and no doubt help to disguise this species from small fishes thatmake up its diet. However, the snakes periodically tie themselves in knots to not only sloughtheir skin (which they do quite regularly) - but also to eliminate such growths that may havebecome uncomfortable.Toxicity: This is a species that will attempt to bite if sufficiently harassed, so caution should beexerted, because its venom is highly toxic to humans. There have been human fatalitiesarising from the bite of this species, so urgent medical attention should always be sought inthe event of an envenomation. Additionally, severe human illness has resulted from eveneating its flesh.Survival Status: Protected under the New South Wales National Parks and Wildlife Act (1974)but not listed in that State as a Threatened Species in any of the Schedules of the NSWThreatened Species Conservation Act (1995). Also protected under the Victorian Wildlife Act(1975) but not listed as threatened in the Victorian Flora and Fauna Guarantee Act (1988)],the Qld Nature Conservation Act (1992), the Territory Parks and Wildlife Conservation Act(1998), the Tasmanian National Parks and Wildlife Act (1970) and the WA WildlifeConservation Act 1950 (as amended). Listed as a Marine Protected Species under theAustralian Environmental Protection and Biodiversity Conservation Act (1999). Probably notunder threat, but its restricted distribution in Australian waters could potentially make itvulnerable in some parts of its range. Historical records suggest that the species may havebeen once more common in New South Wales than at present.Etymology: The name Pelamis is from the Greek ‘Pelamis’ which literally translates to ‘tunnyfish’, a possible reference to their appearance or perhaps it referred to their diet. The name'platurus' was derived from the Greek ‘platys’ meaning ‘flat’ and ‘oura’ meaning ‘tail’ andliterally means 'tail like the blade of an oar', in reference to the broad tail of the species.ReferencesNote: This is not a complete bibliography of the sea snakes, but it does include a wide rangeof material on many aspects of their biology and ecology - and many of these referencescontain other citations to sea snakes as well. Most of the more important works are citedbelow that clearly demonstrates the morphological, physiological, toxicological and ecologicaladaptations that warrant the recognition of the sea snakes as separate Families to theterrestrial Proteroglypha.Ackman, R.G., MacPherson, E.J. and O'Dor, R.K. 1991 Fatty acids of the depot fats from theblue-banded sea snake (Laticauda colubrina) and its principal food the conger eel (Congercinerus). Comparative Biochemistry and Physiology B Comparative Biochemistry, 98 (2-3):423-425Adnagulov, E.V., Tarasosv, I.G. and Gorobeiko, V.V. 2000 New data on amphibians andreptiles distribution in the Russian Far East. Russian Journal of Herpetology, 7 (2): 139-154Agassiz, L. 1846 Nomenclatoris Zoologici Index Universalis. Jent et Gassman, Soloduri [Pp.1-393; see p. 9 for Aepyurus; see p. 297 for use of Platyurus - emend. pro Platurus Sonniniand Latreille, 1802]Aird, S.D. 2004 Taxonomic distribution and quantitative analysis of free purine and pyrimidinenucleosides in snake venoms. Comparative Biochemistry and Physiology, BAlderslade, P.N. et al [contributors] 1984 Reader’s Digest Book of the Great Barrier Reef.Reader’s Digest, Sydney [Pp. 1-384]Allen, M. and Tu, A.T. 1985 The effect of tryptophan modification on the structure andfunction of a sea snake neurotoxin. Molecular Pharmacology, 27(1): 79-8559

Australian Biodiversity Record, 2007 (8): 1-124Allen, R. 1975 Marine parks: The Cinderella of conservation. New Scientist, 14 August: 366-369Alvarez-Leon, R. and Hernandez-Camacho, J.I. 1998 Notas sobre la ocurrencia de Pelamisplaturus (Reptilia: Serpentes: Hydrophiidae) en el Pacifico Colombiano. Caldasia, 20(2): 93-102Anderson, J. 1871 On some Indian reptiles. Proc. Zool. Soc. Lond. 1871: 149-211 [see fordescription of Hydrophis fayeriana; see p. 190 for original descriptions of P[laturus] affinis andHydrophis granosa]Angel, F. 1946 Reptiles and Amphibians. Feder. Fran. Soc. Sci. Naturelle, Paris [Faune deFrance, No 45]Angel, F. 1949 Petit Atlas des Amphibiens et Reptiles. Boubee and Co., Paris [Volume 1: 1-121; Volume 2: 1-141]Annandale, N. 1905 Additions to the collection of oriental snakes in the Indian Museum. Part2. Specimens from the Andamans and Nicobars. Bull. Mus. R. Hist. Nat. Belg. (ns)1: 173-176[see p. 174 for original description of Distira andamanica]Anonymous nd A Glimpse of Poisonous and Venomous Marine Life. SandozPharmaceuticals, Sydney [Pp. 1-47]Anonymous 1931 [No title]. Sydney Morning Herald, 11 March: 14Anonymous 1940 Giant snakes of the coral seas. Walkabout, 6: 3-4 [by 'P']Anonymous 1951 The real sea serpent. Wild Life [Melbourne], 13 (6): 508-512Anonymous 1954 New danger in surf at Bondi. Third sea snake at beach. Daily Telegraph[Sydney], 4 OctoberAnonymous 1962 Man, 39, killed by sea snake. Daily Telegraph [Sydney], 12 MarchAnonymous 1962 Sea snake on beach. Sun [Sydney], 21 NovemberAnonymous 1962 Died after bite by sea snake. Sydney Morning Herald, 12 MarchAnonymous 1962 Deadly sea snake on beach. Sydney Morning Herald, 26 MarchAnonymous 1963 Second sea snake at Manly Beach. Sydney Morning Herald, 3 JanuaryAnonymous 1963 Poisonous Snakes of the World. United States Office of Naval IntelligenceStudy, No 3-62 [Washington]Anonymous 1968 Poisonous Snakes of the World. Department of the Navy, GovernmentPrinting Office, Washington DC [Pp. i-viii, 1-203]Anonymous 1970 The Venomous Australians. Paul Hamlyn, Sydney [Pp. 1-47]Anonymous 1971 Channel for sea snakes. Nature 232: 11-12Anonymous 1975 How sea snakes may avoid the bends. Sea Frontiers, 21 (6): 358Anonymous 2005 Policy on Managing Activities that include the Direct Take of a ProtectedSpecies from the Great Barrier Reef Marine Park. Great Barrier Reef Marine Park Authority,Townsville [Pp. 1-43]Antram, F. 1989 Australian sea snake utilization - an update. Traffic Bulletin, 10(3-4): 3160

Australian Biodiversity Record, 2007 (8): 1-124Aplin, K.P. and Smith, L.A. 2001 Checklist of the frogs and reptiles of Western Australia.Records of the Western Australian Museum, Supplement No 63: 51-74ASIH Committee on Herpetological Common Names 1956 Common Names for NorthAmerican Amphibians and Reptiles. Copeia, 1956 (3): 172-185Averianov, A.O. 1997 Paleocene sea snakes from the eastern part of Tethys. Russian Journalof Herpetology, 4 (2): 128-142Avolio, C., Shine, R. and Pile, A. 2006 Sexual dimorphism in scale rugosity in sea snakes(Hydrophiidae). Biological Journal of the Linnean Society, 89 (2): 343–354Avolio, C., Shine, R. and Pile, A. 2006 The adaptive significance of sexually dimorphic scalerugosity in sea snakes. American Naturalist, 167 (5): 728-738Ayala, S.C. 1978 Checklist, host index, and annotated bibliography of Plasmodium fromreptiles. Journal of Protozoology, 25 (1): 87-100Bacolod, P.T. 1983 Reproductive biology of two sea snakes of the genus Laticauda fromcentral Philippines. Philippine Scientist, 20: 39-56Bacolod, P.T. 1984 Notes on sea snake fishery on Gato Islet, Cebu Island, Philippines and aproposal for a conservation and management program. Philippine Scientist, 21: 155-163Bacolod, P.T. 1990 The biology of some commercially important species of sea snakes(Hydrophiidae) in the Visayas Sea. Philippine Scientist, 27: 61-88Baird, J. and Labuc, G. 1978 Fijian sea snakes. Victorian Herpetological Society Newsletter,10: 6-7Bal, D.V. and Navathe, K.V. 1949 The circulatory system of Hydrophis caerulescens (Shaw).Journal of the University of Bombay, 17(ns): 1-14Baldwin, J. and Seymour, R.S. 1977 Adaptation to anoxia in snakes: Levels of glycolyticenzymes in skeletal muscle. Australian Journal of Zoology, 25 (1): 9-13Banks, C.B. 1989 Management of fully aquatic snakes. International Zoo Yearbook. 28: 155-163Barbour, T. 1912 A contribution to the zoogeography of the East Indian islands. Memoirs ofthe Museum of Comparative Zoology, 44:1-203Barbour, T. 1921 Reptiles and amphibians from the British Solomon Islands. Proceedings ofthe New England Zoological Club, 7: 91-112Barbour, T. 1926 Reptiles and Amphibians. Their Habits and Adaptations. Houghton MifflinCo., London [Pp. 1-125]Barbour, T. 1934 Reptiles and Amphibians. Their Habits and Adaptations. Houghton MifflinCo., Cambridge, Mass. [Revised Edition; pp i-xx, 1-129]Barme, M. 1963 Venomous sea snakes of Viet Nam and their venoms. [Pp. 373-378]. In:Keegan, H.L. and MacFarlane, W.V. (Editors): Venomous and Poisonous Animals andNoxious Plants of the Pacific Region. Pergamon, OxfordBarme, M. 1968 Venomous sea snakes (Hydrophiidae). [Pp. 285-308]. In: Bucherl, W.,Buckley, E.E. and Deulofeu, V. (Editors): Venomous Animals and their Venoms. Volume 1.Venomous Vertebrates. Academic Press, New York [Pp. 1-707]61

Australian Biodiversity Record, 2007 (8): 1-124Barme, M., Huard, M. and Nguyen, X.M. 1962 Preparation d’un serum antivenind’hydrophiides. Premier essais therapeutiques. Ann. Inst. Pasteur [Paris], 102: 497Barrett, C.L. 1950 Reptiles of Australia: Crocodiles, Snakes and Lizards. Cassell, Sydney [Pp.i-xiii, 1-168]Bauer, A. and Devaney, K.D. 1987 Convergence and mimicry in sea snakes and other NewCaledonian reef organisms. [Pp. 43-48]. In: J.J. van Gelder, H. Strijbosch and P.J.M. Bergers,(Editors): Proceeedings of the 4th Ordinary General Meeting of the Societas EuropeaHerpetologica. Faculty of Sciences Nijmegen. Societas Europea Herpetologica, Faculty ofSciences, University of Nijmegen.Bavay, A. 1869 Catalogue des reptiles de la Nouvelle-Calédonie et description d'espècesnouvelles. Mém. Soc. Linn. Normandie 15: 1-37 [see p. 33 for original description ofAipysurus duboisii; see p. 34 for original description of Aipysurus chelonicephalus].Baverstock, P.R. and Schwaner, T.D. 1985 Phylogeny of Australian Elapid snakes: Thegenetic approach. [Pp. 159-164]. In: Grigg, G., Shine, R. and Ehmann, H. (Editors): Biology ofAustralasian Frogs and Reptiles. Surrey Beatty and Sons, Sydney [Note that the publicationdate is erroneously given as August, 1985 - actually published not earlier than November1985]Baxter, E.H. and Gallichio, H.A. 1974 Cross-neutralization by Tiger Snake (Notechis scutatus)antivenene and sea snake (Enhydrina schistosa) antivenene against several sea snakevenoms. Toxicon, 12: 273-278Baxter, E.H. and Gallichio, H.A. 1976 Protection against sea snake envenomation:Comparative potency of four antivenenes. Toxicon, 14 (5): 347-355Becak, M.L. 1975 Chromosome Atlas: Fish, Amphibians, Reptiles, and Birds. Berlin [Volume3; Pp. 1-73]Becak, W. 1968 Karyotypes, sex chromosomes, and chromosomal evolution in snakes. [Pp.53-95]. In: Bucherl, W., Buckley, E.E. and Deulofeu, V. (Editors): Venomous Animals andtheir Venoms. Volume 1. Venomous Vertebrates. Academic Press, New YorkBecke, L. 1925 Some sea snakes. Zoological Society Bulletin [New York], 28 (5): 131-133Behler, J.L. 1979 Animal Kingdom reports: operation sea snake. Animal Kingdom [New York],82 (6):Bell, T. 1843 Reptiles. [Pp. i-vi, 1-51]. In: Darwin, C. (Editor): The Zoology of the Voyages ofH.M.S. Beagle, under the Command of Captain Fitzroy, R.N., during the years 1832-1836.Volume 3. Part 5. Smith, Elder and Co., LondonBellairs, A.d'A. 1957 Reptiles. Hutchinson's University Library, London [Pp. 1-195]Bellairs, A.d'A. 1968 Reptiles. Hutchinson's University Library, London [2nd Edition]Bellairs, A.d'A. 1969 The Life of Reptiles. Weidenfeld and Nicholson, London [Volume 1: Pp.1-282]Bellairs, A.d'A. 1970 The Life of Reptiles. Weidenfeld and Nicholson, London [Volume 2: Pp.283-590]Bellairs, A.d'A. 1971 La Grande Encyclopedie de la Nature. Volume 11. Les Reptiles. EditionsRencontre, Lausanne [Pp. 385-768]62

Australian Biodiversity Record, 2007 (8): 1-124Bellairs, A.d'A. 1972 Comments on the evolution and affinities of snakes. [Pp. 157-172]. In:Joysey, K.A. and Kemp, T.S. (Editors): Studies in Vertebrate Evolution. Oliver and Boyd,Edinburgh [Pp. 1-285]Bellairs, A.d'A. and Carrington, R. 1969 The World of Reptiles. Chatto and Windus, London[Pp. 1-154]Bellairs, A.d'A. and Cox, C.B. (Editors) 1976 Morphology and Biology of Reptiles. AcademicPress, London [Linnean Society Symposium Series, No 3: 1-290]Bellairs, A.d'A. and Underwood, G. 1951 The origin of snakes. Biological Reviews, 26 (2):193-237Benyajati, S., Yokota, S.D. and Dantzler, W.H. 1985 Renal function in sea snakes 2. Sodium,potassium, and magnesium excretion. American Journal of Physiology, 249(2): R237-R245Bergman, A.M. 1943 The breeding habits of sea snakes. Copeia, 1943 (3): 156-160Bergman, R.A.M. 1949 The anatomy of Lapemis hardwickii Gray. I -II. Proceedings of theKoninklijke Nederlandsche Akademie van Wetenschappen [Amsterdam], 52(8): 882-898Bergman, R.A.M. 1962 The anatomy of Hydrophis fasciatus atriceps. Biologisch Jaarboek,30: 389-416Berridge, W.S. 1935 All about Reptiles and Amphibians. London [Pp. 1-271]Berry, P.Y. 1973 Amphibians, reptiles and fishes. Malayan Nature Journal, 25: 45-61Billard, J.B. and Dale, B. 1970 Panama - Link between oceans and continents. NationalGeographic, 137 (3): 440Biswas, S. and Sanyal, D.P. 1980 A report on the Reptilia fauna of Andaman and NicobarIslands in the collection of Zoological Survey of India. Records of the Zoological Survey ofIndia, 77: 255-292Blackburn, D.G. 1993 Chorioallantoic placentation in squamate reptiles: structure, function,development, and evolution. Journal of Experimental Zoology, 266: 414-430Blackburn, D.G. 1998 Structure, function, and evolution of the oviducts of squamate reptiles,with special reference to viviparity and placentation. J. Exper. Zool., 282 (4/5): 560-617 [seealso Herp. Review, 30(1): 9]Blakeslee, S. 1984 Mysterious sea snakes lure a research team. Chicago HerpetologicalSociety Newsletter, 1984 (June): 1pBlanc, C.P. 1972 Le Reptiles de Madagascar et des iles voisines. [Pp. 501-614]. In: Battistini,R. and Richard-Vindard, G. (Editors): Biogeography and Ecology of Madagascar. Junk, TheHagueBlanc, C.P., Ineich, I. and Blanc, F. 1983 Composition et distribution de la faune des Reptilesterrestres en Polynesie Francaise. Bulletin Soc. etudes Oceaniennes, No 223, 18 (12): 1323-1335Bleeker, P. 1858 Eene Zeeslang (Aipysurus margaritophorus). Natuurkundig Tijdschrift voorNederlandsch Indie, (4)16: 49-50 [see p. 49 for original description of Aipysurusmargaritophorus].Bleeker, P. 1859 Reptilien van Nieuw-Guinea. Natuurkundig Tijdschrift voor NederlandschIndie, 16: 420-42563

Australian Biodiversity Record, 2007 (8): 1-124Boettger, O. 1888. Uber aussere Geschlechtscharactere bei den Seeschlangen. ZoologischeAnzeiger, 284: 395-398Boettger, O. 1890 On a monstrous specimen of Hydrus platurus, L. with two pairs of nostrils.Bericht des Senckenbergischen Gesellschaft, 1890: lxxivBoettger, O. 1892 Listen von Kriechtieren und Lurchen aus dem tropischen Asien und ausPapuasien. Bericht des Offenbacher Vereins fur Naturkunde, 29-32: 65-164Boettger, O. 1898 Katalog der Reptilien-Sammlung im Museum der SenckenbergischenNaturforschenden Gesellschaft in Frankfurt am Main. II. Teil (Schlangen). SenckenbergischenNaturforschenden Gesellschaft (Gebrader Knauer), Frankfurt am Main [Pp. i-ix + 1-160]Bogert, C.M. 1954 Amphibians and Reptiles of the World. [Pp. 1189-1390]. In: Drimmer(Editor): The Animal Kingdom. Doubleday, New YorkBoie, F. 1827 Bemerkungen über Merrem's Versuch eines Systems der Amphibien. IsisOken, 20: 508-566 [see p. 553 in regards to Leioselasma striata; see p. 554 for originaldescription of Hydrus valakadyn - but see pl. 11 in Russell, P. (1801); see p. 554 for originaldescription of Hydrus kadellnagam - as Hydrus Kadell-Nagam ; note: description based onPlate 13 in Russell, P. (1801)]Bolanos, R., Flores, A., Taylor, R.T. and Cerdas, L. 1974 Color patterns and venomcharacteristics in Pelamis platurus. Copeia, 1974 (4): 909-912Bory de Saint-Vincent, J.B.G.M. 1828 Resume d'Erpetologie, ou d'Histoire Naturelle desReptiles. Bachelier, Paris [Pp. 1-292]Boulenger, E.G. 1893 Reptiles and Batrachians. Dent and Sons, LondonBoulenger, E.G. 1904 Reptiles and Batrachians. Dent and Sons, London [Another edition]Boulenger, E.G. 1914 Reptiles and Batrachians. Dent and Sons, London [Another edition]Boulenger, G.A. 1890 The Fauna of British India; including Ceylon and Burma. Reptilia andBatrachia. Taylor and Francis, London [Pp. i-xviii, 1-541]Boulenger, G.A. 1896 Catalogue of Snakes in the British Museum (Natural History). 3. BritishMuseum, London [Pp. i-xiv, 1-727+25 plates; see p. 269 for erection of Acalyptophis nom.substit. pro Acalyptus Duméril, 1853; see p. 270 and pl. 12, fig. 1 for original descriptions ofHydrelaps and Hydrelaps darwiniensis; see p. 276 for original description of Hydrophis kingii;see p. 278 for original description of Hydrophis pacificus; see p. 285 for use of Distira; see p.293 for original description of Distira grandis; see p. 294, pl. 17 fig. 1 for original description ofDistira macfarlani; see p. 309 for original description of Platurus Muelleri - ex Müller MS]Boulenger, G.A. 1898 Description of a new sea-snake from Borneo. Proc. Zool. Soc. Lond.1898: 106-107 [see p. 106, pl. 9 for original description of Hydrophis floweri]Boulenger, G.A. 1898 On a little-known sea-snake from the South Pacific. [Pp. 57-58]. In:Zoologial results based on material from New Britain, New Guinea, Loyalty Islands andelsewhere during the years 1895, 1896 and 1897. Volume 1. Cambridge University Press,CambridgeBoulenger, G.A. 1908 Note on the ophidian genus Emydocephalus. Annals and Magazine ofNatural History, (8)1: 111-115Boulenger, G.A. 1912. A Vertebrate Fauna of the Malay Peninsula from the Isthmus of Kra toSingapore Including the Adjacent Islands. Taylor and Francis, London [Pp. 1-294]64

Australian Biodiversity Record, 2007 (8): 1-124Bourret, R. 1935 Les serpents marins de l'Indochine francaise. Institut Oceanographiquel'Indochine [Hanoi], 25: 1-69Bourret, R. 1936 Les serpents de l'Indochine. 1. Ãstudes sur la fauna. Henri Basuyau & Cie,Toulouse [Pp. 1-141]Branch, W.R. 1979 The venomous snakes of southern Africa. Part 2: Elapidae andHydrophiidae. Snake, 11: 199-225Brazaitis, P. 1992 Snakes of the World. Crescent Boooks, New York [Pp. 1-176]Brazenor, C.W. 1936 The Reptiles and Amphibians of Victoria. [Pp. 25-38]. In: Gawler, O.(Editor): Victorian Yearbook for 1934-35. Government Printer, MelbourneBreen, J.F. 1974 Encyclopedia of Reptiles and Amphibians. TFH Publications, Neptune City,New Jersey [Pp. 1-576]Broad, A.J., Sutherland, S.K. and Coulter, A.R. 1979 The lethality in mice of dangerousAustralian and other snake venom. Toxicon, 17: 664-667Bromham, L. 2002 Molecular clocks in reptiles: Life history influences rate of molecularevolution. Molecular Biology and Evolution, 19 (3): 302-309 [see data on Laticaudidae]Brongersma, L.D. 1931 Resultats scientifiques du voyage aux Indes Orientales Neerlandaisesde le Prince et la Princesse Leopold de Belgique. Reptiles. Memoires Musee Royald'Histoire Naturelle de Belgique, 5 (2): 3-39Brongersma, L.D. 1938 Het belang van anatomisch onderzoek voor de systematiek. E.J. Brill,Leiden [11pp.]Brongersma, L.D. 1956 Notes on New Guinean reptiles and amphibians. V. Proceedings ofthe Koninklijke Nederlandse Akademie van Wetenschappen, Series C, Biological and MedicalSciences, 59: 599-610 [Hydrelaps darwiniensis]Brongersma, L.D. 1958 The Animal World of Netherlands New Guinea. J.B. Wolters,GroningenBrongersma, L.D. 1961 Zoological exploration of Netherlands New Guinea. Proceedings ofthe Ninth Pacific Science Congress, 19 (Zoology): 68-71Brown, J.H. 1973 Toxicology and Pharmacology of Venoms from Poisonous Snakes. CharlesC. Thomas, Springfield, Illinois [Pp. i-xiv, 1-184]Brown, J.N.B. 1989 Sea snakes of the Arabian Gulf. Wildlife Views, 4(11): 2-5Bruun, A. (Editor) 1956 The Galathea Deep Sea Exploration, 1950-1952. Macmillan, NewYorkBucherl, W. and Buckley, E.E. (Editors) 1971 Venomous Animals and their Venoms. Volume2. Venomous Vertebrates. Academic Press, New YorkBucherl, W., Buckley, E.E. and Deulofeu, V. (Editors) 1968 Venomous Animals and theirVenoms. Volume 1. Venomous Vertebrates. Academic Press, New York [Pp. i-xxii, 1-707]Buddle, R. 1929 Snakes of Singapore Island. Kelly and Walsh Ltd, Singapore [Pp. 1-48]Buden, D.W. 1995 Reptiles, birds, and mammals of Mokil and Pingelap Atolls, easternCaroline Islands. Micronesica, 28: 9-2365

Australian Biodiversity Record, 2007 (8): 1-124Burger, W.L. and Natsuno, T. 1974 A new genus for the Arafura smooth sea snake andredefinitions of other sea snake genera. Snake, 6 (2): 61-75 [see p. 65 for original descriptionof Parahydrophis]Burns, B. 1969 An Oral Sensory System in Sea Serpents (Hydophiidae): Histological andMorphological Considerations. MSc Thesis, University of Waterloo [Pp. 1-22]Burns, B. 1969 Oral sensory papillae in sea snakes. Copeia, 1969 (3): 617-619Burns, B. and Pickwell, G.V. 1972 Cephalic glands in sea snakes (Pelamis, Hydrophis andLaticauda). Copeia, 1972 (3): 547-559Burns, G.W. 1984 Aspects of the Population Movements and Reproductive Biology ofAipysurus laevis, the Olive Sea Snake. PhD Thesis, University of New England, ArmidaleBurns, G.W. 1985 The female reproductive cycle of the Olive Sea Snake, Aipysurus laevis(Hydrophiidae). [Pp. 339-341]. In: Grigg, G., Shine, R. and Ehmann, H. (Editors): Biology ofAustralasian Frogs and Reptiles. Surrey Beatty and Sons, Chipping Norton [Sydney] [Notethat the publication date is erroneously given as August, 1985 - actually published not earlierthan November 1985]Burns, G.W. 1985 Aspects of population movements and reproductive biology of Aipysuruslaevis, the Olive Sea Snake. Australian Journal of Ecology, 10 (3): 368-369 [PhD Thesisabstract only]Burns, G.W. and Heatwole, H. 1998 Home range and habitat use of the Olive Sea Snake,Aipysurus laevis, on the Great Barrier Reef, Australia. Journal of Herpetology, 32(3): 350-358Burns, G.W. and Heatwole, H. 2000 Growth, sexual dimorphism, and population biology ofthe Olive Sea Snake, Aipysurus laevis, on the Great Barrier Reef of Australia. Amphibia-Reptilia, 21(3): 289-300Burton, M. 1978 Snakes, Crocodiles and Lizards - The World of Reptiles and Amphibians.Paul Hamlyn, SydneyBurton, R. 1978 Venomous Animals. Stafford Pemberton Publishing, Crows Nest (Sydney)[Pp. 1-64]Bush, A.O. and Holmes, J.C. Plicatrium visayanensis new species (Digenea: Hemiuridae)from Hydrophis ornatus (Serpentes: Hydrophiidae) from the Visayan Sea, Philippines.International Journal of Parasitology, 14(1): 35-38Bush, A.O. and Holmes, J.C. 1979 Sterrhurus carpentariae nov. sp. (Digenea: Hemiuridae)from Lapemis hardwickii (Serpentes: Hydrophiidae) from the eastern Gulf of Carpentaria,Australia. International Journal of Parasitology, 9(3): 189-192Bush, B., Maryan, B., Browne-Cooper, R. and Robinson, D. 1995 Reptiles and Frogs of thePerth Region. University of Western Australia Press, Perth [Pp. i-xiv. 1-226]Bussarawit, S., Rasmussen, A.R. and Andersen, M. 1989 A preliminary study on sea snakes(Hydrophiidae) from Phuket Harbour, Phuket Island, Thailand. Natural History Bulletin of theSiam Society, 37 (2): 209-225Cadle, J.E. and Gorman, G.C. 1981 Albumin immunological evidence and the relationships ofsea snakes. Journal of Herpetology, 15 (3): 329-334Cagle, F.R. 1957 Reptiles. [Pp. 275-358]. In: Blair, W.F., et al (1957): Vertebrates of theUnited States.66

Australian Biodiversity Record, 2007 (8): 1-124Calmette, A. 1908 Venoms, Venomous Animals and Antivenomous Serum - Therapeutics.John Bale, Sons and Danielsson, London [Pp. i-xvi, 1-403]Calnan, T. 2006 Rainforest to Reef just 40 km from Darwin. An assessment of theconservation values of the Gunn Peninsula/Vernon Islands area and the impacts of theproposed Glyde Point heavy industry and residential estate. Report to the EnvironmentCentre of the Northern Territory, and the Australian Marine Conservation Society, Darwin [Pp.1-58]Cameron, E.E. and Cogger, H.G. 1992 The herpetofauna of the Weipa Region, Cape YorkPeninsula. Technical Reports of the Australian Museum, 7: 1-200Campbell, C.H. 1964 Venomous snake bite and its treatment in the Territory of Papua andNew Guinea. Papua New Guinea Medical Journal, 7(1):1-11Campbell, C.H. 1976 Snake Bite, Snake Venoms and Venomous Snakes of Australia andNew Guinea. An annotated bibliography. School of Public Health and Tropical Medicine,University of Sydney, Service Publication No 13: i-ix, 1-222Campbell, J.A. and Lamar, W.W. 1989 The Venomous Reptiles of Latin America. ComstockPublications, Cornell University Press, Ithaca [Pp. i-xi, 1-425]Campden-Main, S.M. 1970 A Field Guide to the Snakes of South Vietnam. United StatesNational Museum, Division of Reptiles and Amphibians, Washington [Pp. 1-114]Candor, S., Puffer, H. and Tamiya, N. 1975 Immunological aspects of venom of sea snakesfrom the Indo-Pacific. Toxicon, 13 (2): 89 [Abstract only]Cantor, T.E. 1841 Observations upon pelagic serpents. Transactions of the ZoologicalSociety of London 2(4): 303-313Cantor, T.E. 1847 Catalogue of Reptiles inhabiting the Malay Peninsula and Islands. Journalof the Asiatic Society of Bengal, 16 (2): 897-902, 1026-1078 [Facsimile reprint published in1966 by Asher and Co., Amsterdam]Capocaccia, L. 1961 Catalogo dei tipi di Rettili del Museo Civico di Storia Naturale di Genova.Ann. Mus. Civ. Stor. Nat. 'Giacomo Doria' 72: 86-111 [see treatment of Pelagophis lubricus]Caras, R. 1964 Dangerous to Man. Chilton Books, PhiladelphiaCaras, R. 1978 Dangerous to Man. Penguin, Harmondsworth (UK) [Another Edition]Carey, J.E. and Wright, A.S. 1960 The toxicity and immunological properties of some seasnake venoms with particular reference to that of Enhydrina schistosa. Transactions of theRoyal Society of Tropical Medicine and Hygiene, 52: 50-67Carey, J.E. and Wright, E.A. 1961 The site of action of the venom of the sea snake Enhydrinaschistosa. Transactions of the Royal Society of Tropical Medicine and Hygiene, 55: 153-160Carlson, B.A. 2000 Husbandry of the Yellow-bellied Sea Snake, Pelamis platurus. AmericanZoo and Aquarium Association Annual Conference Proceedings 2000: 27-29Casas-Andreu, G. 1990 Pelamis platurus (Yellow-belly Sea Snake). Herpetological Review,21(2): 41Challen, R.G. 1975 Venomous Australian snakes and antivenenes. Australian Journal ofHospital Pharmacy, 5 (1): 4-16Chapgar, B.F. and Kewalramani, H.G. 1967 Mating and other observations on sea snakes incaptivity. Journal of the Bombay Natural History Society, 64 (3): 563-56567

Australian Biodiversity Record, 2007 (8): 1-124Chazeau, J. 1995 Bibliographie Indexee de la Faune Terrestre de Nouvelle-Caledonie.Systematique, Ecologie et Biogeographie. Editions Orstrom, Noumea [Pp. 1-95]Chetty, N., Du, A., Hodgson, W.C., Winkel, K. and Fry, B.G. 2004 The in vitro neuromuscularactivity of Indo-Pacific sea-snake venoms: Efficacy of two commercially available antivenoms.Toxicon, 44: 193-200Chippaux, J.P., Williams, V., White, J., 1991 Snake venom variability: Methods of study,results and interpretation. Toxicon, 29: 1279-1303Chubb, I.W. and Greffen, L.B. (Editors) 1979 Neurotoxins: Fundamental and ClinicalAdvances. Adelaide University Union Press, AdelaideCiliento, R.W. 1944 Some poisonous plants, sea and land animals of Australia and NewGuinea. Smith and Robertson, BrisbaneCloudsley-Thompson, J.L. 1971 The Temperature and Water Relations of Reptiles. Merrow,Watford, Herts [Pp. 1-159]Coborn, J. 1991 Atlas of Snakes of the World. TFH Publications, New JerseyCochran, D.M. 1943 Poisonous Reptiles of the World: A Wartime Handbook. SmithsonianInstitution, Washington ['War Background Studies No 10'; Publication No 3727, Pp. i-v, 1-37]Cochran, D.M. 1944 Dangerous Reptiles. Annual Report of the Smithsonian Institution[Washington DC], for 1943: 275-323 [Smithsonian Institution, Publication No 3753]Cochran, D.M. and Goin, C.J. 1970 The New Field Book of Reptiles and Amphibians.Putnam, New York [Pp. 1-359]Cogger, H.G. 1959 Sea snakes. Australian Museum Magazine, 13 (2): 34-41Cogger, H.G. 1962 Snakes, Lizards and Chelonians. In: The Natural History of Sydney.Australian Museum, SydneyCogger, H.G. 1963 Sea snakes. In: International convention on life saving techniques(Sydney, March 11-20, 1960) - "B" Group - Scientific Section. The Hazards of DangerousMarine Creatures. Sydney [Pp. 5-6; see also "Discussion" Pp. 6-7]Cogger, H.G. 1967 Australian Reptiles in Colour. Reed, Sydney [Pp. 1-112]Cogger, H.G. 1967 The snakes of Australia. [Pp. 152-158]. In: McMichael, D.F. (Editor):Treasury of Australian Wildlife. Ure Smith, SydneyCogger, H.G. 1971 The venomous snakes of Australia and Melanesia. [Pp. 35-77]. In:Bucherl, W. and Buckley, E.E. (Editors): Venomous Animals and Their Venoms. Volume 2.Venomous Vertebrates. Academic Press, New York [Pp. 1-687]Cogger, H.G. 1972 Keys to the frogs and reptiles of the central coast of New South Wales.Part II. Lizards and Snakes. Herpetofauna, 5 (3): 9-15Cogger, H.G. 1972 Reptiles. [Pp. 1012-1013]. In: Ryan, P.A. (General Editor): Encyclopaediaof Papua and New Guinea. Volume 2. Melbourne University Press, MelbourneCogger, H.G. 1972 Australian Reptiles in Colour. Reed, Sydney [Revised Edition; Pp. 1-112]Cogger, H.G. 1972 Snakes, Lizards and Chelonians. [Pp. 35-38]. In: The Natural History ofSydney. Australian Museum, Sydney [2nd Edition; Pp. 1-63]68

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Australian Biodiversity Record, 2007 (8): 1-124Gilboa, I. and Dowling, H.G. 1972 A bibliography on chromosomes of amphibians andreptiles. HISS Publications in Herpetology, No 4: 1-33Gill, B.J. 1986 Collins Handguide to the Frogs and Reptiles of New Zealand. Collins,Auckland [Pp. 1-112]Gill, B.J. 1997 Records of turtles and sea snakes in New Zealand, 1837-1996. New ZealandJournal of Marine and Freshwater Research, 31(4): 477-486Glasby, C.J., Ross, G.J.B. and Beesley, P.L. (Editors) 1993 Fauna of Australia. Volume 2A.Amphibia and Reptilia. Australian Government Publishing Service, Canberra [Pp. 1-439]Glauert, L. 1928 The vertebrate fauna of Western Australia, Part 1. Reptilia (Testudinata,Ophidia). Journal of the Royal Society of Western Australia, 14: 61-77Glodek, G.S. and Voris, H.K. 1982 Marine snake diets: prey composition, diversity andoverlap. Copeia, 1982 (3): 661-666Golay, P. 1985 Checklist and Keys to the Terrestrial Proterogyphs of the World (Serpentes:Elapidae - Hydrophiidae). Elapsoidea, fondation culterelle vivarium, Geneva [Pp. i-ix, 1-91;available from the author, 1 Vieux-Moulin 1213 Onex, Switzerland]Golay, P., Smith, H.M., Broadley, D.R., Dixon, J.R., McCarthy, C., Rage, J.-C., Schatti, B. andToriba, M. (Editors) 1993 Endoglyphs and Other Major Venomous Snakes of the World: AChecklist. Azemiops, Geneva, Switzerland [Pp. i-xv + 1-478]Gonzalez-Fernandez, J.E. 1997 La coleccion de Batracolgia y Herpetologia del MuseoNacional de Ciencias Naturales (CSIC). Graellsia, 53 : 95-100 [collection includes specimensof Hydrophiidae and Laticaudidae]Gopalakrishnakone, P. 1985 Structure of the spinous scales of Lapemis hardwickii (1). Light,transmission electron and scanning electron microscopic studies. Snake, 17(2): 149-155Gopalakrishnakone, P. 1985 Structure of the venom glands of sea snakes (Hydrophiidae).Toxicon, 23(4): 571Gopalakrishnakone, P. (Editor) 1990 A Colour Guide to Dangerous Animals. SingaporeUniversity Press, Singapore [Pp. 1-156]Gopalakrishnakone, P. and Kochva, E. 1990 Venom glands and some associated muscles insea snakes. Journal of Morphology, 205: 85-96Gopalakrishnakone, P. and Kochva, E. 1993 Histological features of the venom apparatus ofsea snake Lapemis curtus. Snake, 25(1): 27-37Gorman, G.C. 1981 The chromosomes of Laticauda and a review of karyotypic evolution inthe Elapidae. Journal of Herpetology, 15 (2): 225-233Gorman, G.C., Licht, P., McCollum, F. 1981 Annual reproductive patterns in three species ofmarine snakes from the central Phillippines. Journal of Herpetology, 15 (3): 335-354Gow, G.F. 1976 Snakes of Australia. Angus and Robertson, Sydney [Pp. 1-88]Gow, G.F. 1977 Snakes of the Darwin area. Northern Territory Museum Publication, Darwin[Undated first Edition- see Anon. (1977); Note: of the original 5000 produced, 4650 weredestroyed at the printers - see also review by Wells, R.W. (1978)]Gow, G.F. 1978 Snakes of the Darwin area. Northern Territory Museum Publication, Darwin[2nd Edition, dated 1977, but not published until 1978]78

Australian Biodiversity Record, 2007 (8): 1-124Gow, G.F. 1981 Checklist of reptiles and amphibians of the northern sector of the NT.Northern Territory Naturalist, No 4: 16-19Gow, G.F. 1982 Australia's Dangerous Snakes. Angus and Robertson, London [Pp. 1- 80 +plates]Gow, G.F. 1983 Snakes of Australia. Angus and Robertson, London [2nd Edition (the'Completely Revised Edition')]Gow, G.F. 1989 Graeme Gow's Complete Guide to Australian Snakes. Angus and Robertson,Sydney [Pp. 1-171]Graham, J.B. 1974 Aquatic respiration of the sea snake Pelamis platurus. RespirationPhysiology, 21: 1-7Graham, J.B. 1974 Body temperatures of the sea snake Pelamis platurus. Copeia, 1974 (2):531-533Graham, J.B., Gee, J.H. and Robinson, F.S. 1975 Hydrostatic and gas exchange functions ofthe lung of the sea snake, Pelamis platurus. Comparative Biochemistry and Physiology, 50A:477-482Graham, J.B., Gee, J.H., Motta, J. and Rubinoff, I. 1987 Subsurface buoyancy regulation bythe sea snake Pelamis platurus. Physiological Zoology, 60 (2): 251-261Graham, J.B., Lowell, W.R., Rubinoff, I. and Motta, J. 1987 Surface and subsurfaceswimming of the sea snake Pelamis platurus. Journal of Experimental Biology, 127: 27-44Graham, J.B., Rubinoff, I. and Hecht, M.K. 1975 Temperature physiology of the sea snakePelamis platurus. An index to its colonization potential in the Atlantic Ocean. Proceedings ofthe National Academy of Sciences, 68: 1360-1363Grasse, P.-P. (Editor) 1970 Traite de Zoologie. Tome XIV. Reptiles. Masson et Cie, ParisGray, J.E. 1825 A synopsis of the genera of reptiles and Amphibia, with a description of somenew species. Annls. Philos. (2)10: 193-217 [see designation of Pelamis bicolor Schneider,1799 as Type Species for Pelamis]Gray, J.E. 1835 Illustrations of Indian Zoology; Chiefly Selected from the Collection of Major-General Hardwicke. London [Vol. 2 see Plate 87 fig. 2 for original description of Lapemis; seep. 19-20 pl. 87 for original description of Lapemis hardwickii]Gray, J.E. 1841 A catalogue of the species of Reptiles and Amphibia hitherto described asinhabiting Australia, with a description of some new species from Western Australia, andsome remarks on their geographical distribution. [Pp. 422-449]. In: Grey, G. (1841): Journalsof Two Expeditions of Discovery in North-west and Western Australia, during the years 1837,38 and 39, under the Authority of Her Majesty's Government describing many newlydiscovered, important, and fertile districts, with Observations on the Moral and PhysicalConditions of the Aboriginal Inhabitants &c, &c. T. and W. Boone, London [volume 2;Appendix E; see p. 433 for use of Aspisurus]Gray, J.E. 1842 Description of some hitherto unrecorded species of Australian reptiles andbatrachians. [Pp. 51-57]. In: Gray, J.E. (Editor): Zoological Miscellany. Treuttel, Würz and Co.,London [Date of publication, April 1842; see p. 55 for use of Asturia - nomen nudum]Gray, J.E. 1842 Monographic synopsis of the water snakes, or the family Hydridae. [Pp. 59-68]. In: Gray, J.E. (Editor): Zoological Miscellany. Treuttel, Würz and Co., London [Date ofpublication: May 1842; see p. 60 for original description of Pelamis ornate; see p. 60 fororiginal description of Liopala; also for Liopola errore pro Liopala; see p. 61 for original79

Australian Biodiversity Record, 2007 (8): 1-124description of Aturia elegans and for original description of Aturia; see p. 61 for originaldescription of Aturia ornate; see p. 62 for original description of Hydrophis mentalis; see p. 62for original descriptions of Hydrophis bengalensis and Hydrophis subfasciata]Gray, J.E. 1843 Description of two new species of reptiles from the collection made during thevoyages of H.M.S. Sulphur. Ann. Mag. Nat. Hist. 11: 46 [see p. 46 for original description ofLapemis loreatus]Gray, J.E. 1845 The reptiles of Australia. [Pp. 1-8 + plates]. In: Richardson, J. and Gray, J.E.(Editors): The Zoology of the Voyage of H.M.S. Erebus and Terror, under the Command ofCapt. J. C. Ross during 1839-1845. Volume 2. Reeve Brothers, London [E.W. Janson,London]Gray, J.E. 1846 A new genus of sea-snake from Port Essington. Ann. Mag. Nat. Hist. 18: 284[see p. 284 for original descriptions of Hypotropis and Hypotropis jukesii]Gray, J.E. 1846 Descriptions of some new Australian reptiles. [Pp. 498-504]. In: Stokes, J.L.(Editor): Discoveries in Australia; With an Account of the Coasts and Rivers Explored andSurveyed During the Voyage of H.M.S. Beagle in the years 1837-38-39-40-41-42-43. Volume1. T. and W. Boone, London [see p. 502, pl. 3 for original description of Hydrus stokesii]Gray, J.E. 1849 Catalogue of the Specimens of Snakes in the Collection of the BritishMuseum. British Museum, London [Pp. i-xv, 1-125; see p. 36, 56 for original description ofChitulia; see for designation of Hydrus gracilis Shaw, 1802 for Type Species of Liopala Gray1842; see p. 46 for original description of Aturia belcheri; see p. 47 for original description ofEnhydrina; see p. 53 for original description of Hydrophis ocellate; see p. 56 for originaldescription of Chitulia inornata and Chitulia fasciata; p. 59 for original description of Hydrusannulatus; see p. 59 for original descriptions of Tomogaster and Tomogaster eydouxii]Gray, M.E. 1930 Note on sea snakes. Australian Naturalist, 8 (4): 88Greene, H.W. 1988 Antipredator mechanisms in reptiles. [Pp. 1-152]. In: Biology of theReptilia. Volume 16. Academic Press, New YorkGreene, H.W. 1997 Snakes: The Evolution of Mystery in Nature, University of CaliforniaPress, BerkeleyGreer, A.E. 1993 Lineage-associated asymmetries in scale overlap patterns in squamates.Herpetologica, 49 (3): 318-326Greer, A.E. 1997 The Biology and Evolution of Australian Snakes. Surrey Beatty, Sydney [Pp.i-xii, 1-358]Greer, A.E. 2006 Encyclopedia of Australian Reptiles. Australian Museum Onlinehttp://www.amonline.net.au/herpetology/research/encyclopedia.pdf Version date: 7 August2006.Grigg, G.C., Shine, R. and Ehmann, H.F.W. (Editors) 1985 The Biology of Australasian Frogsand Reptiles. Surrey Beatty and Sons, Sydney [Pp. i-xvi, 1-527; Note that the publication dateis erroneously given as August, 1985 - actually published not earlier than November 1985]Gritis, P. and Voris, H.K. 1990 Variability and significance of parietal and ventral scales inmarine snakes of the genus Lapemis (Serpentes: Hydrophiidae), with comments on theoccurrence of spiny scales in the genus. Fieldiana. Zoology, (ns) No. 56: 1-13Gudger, E.W. 1938 An unpublished manuscript by the late Bashford Dean; 'Note on the seasnakePelamis platurus (Linnaeus)'. Science, 88 (2276): 144-14580

Australian Biodiversity Record, 2007 (8): 1-124Guillette, L.J. 1987 The evolution of viviparity in fishes, amphibians and reptiles: An endocrineapproach. [Pp. 523-562]. In: Norris, D.O. and Jones, R.E. (Editors): Hormones andReproduction in Fishes, Amphibians and Reptiles. Plenum Press, New YorkGuinea, M.L. 1981 The sea snakes of Fiji. Proceedings of the Fourth International Coral ReefSymposium, Manila, Volume 2: 581-585Guinea, M.L. 1986 Aspects of the Biology of the Common Fijian Sea Snake Laticaudacolubrina (Schneider). MSc Thesis, University South Pacific, Suva, Fiji [Pp. 1-224]Guinea, M.L. 1991 Rainwater drinking by the Sea Krait Laticauda colubrina. Herpetofauna, 21(2): 13-14Guinea, M.L. 1992 The Yellow-bellied Sea Snake Pelamis platurus in the Northern Territory.Northern Territory Naturalist, 13: 37-39Guinea, M.L. 1993 Reptilia, Aves and Mammalia. [Pp. 74-83]. In: Russell, B.C. and Hanley,J.R. (Editors): The Marine Biological Resources and Heritage Values of Cartier and HiberniaReefs, Timor Sea. Report of the Northern Territory Museum Cartier and Hibernia ReefsExpedition, May 1992 Northern Territory Museum of Arts and Sciences, DarwinGuinea, M.L. 1996 Functions of the cephalic scales of the sea snake Emydocephalusannulatus. Journal of Herpetology, 30(1): 126-128Guinea, M.L., McGrath, P. and Love, Bill 1993 Observations of the Port Darwin Sea SnakeHydrelaps darwiniensis. Northern Territory Naturalist, 14: 28-30Guinea, M.L., Tamiya, N. and Cogger, H.G. 1983 The neurotoxins of the sea snake Laticaudaschistorhynchus. Biochemical Journal, 213: 39-41Gunther, A.C.L.G. 1858 Catalogue of Colubrine Snakes in the collection of the BritishMuseum. British Museum (Natural History), London [Pp. i-xvi, 1-281]Gunther, A.C.L.G. 1864 The Reptiles of British India. Royal Society, London [Pp. i-xxvii, 1-444; see p. 367, pl. 25 fig. E for original description of Hydrophis melanosome; see p. 371, pl.25 fig. 1 for original description of Hydrophis atriceps; see p. 375, pl. 25 fig. 0 for use of nameHydrophis longiceps; see p. 377, pl. 25 fig. n for original description of Hydrophis ellioti]Guo, Y.-W., Mao, S.-H. and Yin, F.-Y. 1987 Comparison of cobra plasma albumins with thoseof Banded Krait and sea snakes. Comparative Biochemistry and Physiology, (B) 87 (3): 559-566Gutierrez, J.M. and Bolanos, R. 1980 Karyotype of the Yellow-bellied Sea Snake, Pelamisplaturus. Journal of Herpetology, 14 (2): 161-165Habermehl, G. 1981 Venomous Animals and their Toxins. Springer-Verlag, Berlin and NewYork [Pp. i-ix + 1-195]Halliday, T.R. and Adler, K. (Editors) 1986 The Encyclopaedia of Reptiles and Amphibians.George Allen and Unwin, London [Pp. i-xvi, 1-143]Halstead, B.W. 1959 Dangerous Marine Animals. Cornell Maritime Press, Cambridge,Maryland [Pp. 1-146]Halstead, B.W. 1969 Toxicology of marine animals. In: Firth, F.E. (Editor): The Encyclopediaof Marine Resources. Van Nostrand Reinhold, New YorkHalstead, B.W. 1980 Dangerous Marine Animals that bite, sting, shock, are non-edible.Cornell Maritime Press, Centerville, Maryland [Pp. i-xii + 1-208]81

Australian Biodiversity Record, 2007 (8): 1-124Halstead, B.W. (Editor) 1970 Poisonous and Venomous Marine Animals of the World. UnitedStates Government Printing Press, Washington, DC [Volume 3 - Vertebrates; Pp. 1-1006]Halstead, B.W. (Editor) 1978 Poisonous and Venomous Marine Animals of the World.Princeton, New Jersey [Revised Edition]Halstead, B.W. and Auerbach, P.S. 1992 Dangerous Aquatic Animals of the World: A ColorAtlas of Dangerous Marine Animals. Darwin Press, Princeton, New Jersey [Pp. i-xix + 1-264]Halstead, B.W., Auerbach, P.S. and Campbell, D. 1990 A Colour Atlas of Dangerous MarineAnimals. Wolfe Medical Publications, London [Pp. 1-192]Halstead, B.W., Engen, P.C. and Tu, A.T. 1978 The venom and venom apparatus of the seasnake Lapemis hardwickii Gray. Zoological Journal of the Linnean Society, 63: 371-396Haly, A. 1891 Report on the collections of Reptilia and Batrachia in the Colombo Museum.Government Printing Works, ColomboHan, K.H. 1988 The Reproductive Ecology and Diet of a Sea Snake (Lapemis hardwickiiGray, 1835) from West Coast of Sabah. BSc Thesis, Faculty of Sciences and NaturalResources, University Kebansaan, Malaysia, Sabah Campus [Pp. 1-102]Han, K.H., Stuebing, R.B. and Voris, H.K. 1991 Population structure and reproduction in themarine snake, Lapemis hardwickii Gray, from the west coast of Sabah. Sarawak MuseumJournal, 42 (63): 463-475Harding, J. 1975 We dived into an ocean of sea snakes. Australian Outdoors, (ns) 53 (2): 10-13, 66Harding, K.A. and Welch, K.R.G. 1980 Venomous Snakes of the World: A Checklist.Pergamon Press, Oxford [Pp. 1-188]Hashimoto, Y. 1979 Marine toxins and other Bioactive Marine Metabolites. Japan ScientificSocieties Press, Tokyo [Pp. 1-369]Hattori, Z., Inoue, T. and Okonogi, T. 1974 In front of a pile of sea snake carcasses. Snake, 6:94-98Heatwole, H.F. 1975 Biogeography of reptiles on some of the islands and cays of easternPapua-New Guinea. Atoll Research Bulletin No. 180: 1-32Heatwole, H.F. 1975 Voluntary submergence times of marine snakes. Marine Biology, 32:205-213Heatwole, H.F. 1975 Sea snakes of the Gulf of Carpentaria. [Pp. 145-149]. In: Dunson, W.A.(Editor): The Biology of Sea Snakes. University Park Press, Baltimore [Pp. 1-530]Heatwole, H.F. 1975 Sea snakes found on reefs in the southern Coral Sea (Saumarez,Swains, Cato Island). [Pp. 163-171]. In: Dunson, W.A. (Editor): The Biology of Sea Snakes.University Park Press, Baltimore [Pp. 1-530]Heatwole, H.F. 1975 Predation on sea snakes. [Pp. 233-249]. In: Dunson, W.A. (Editor): TheBiology of Sea Snakes. University Park Press, Baltimore [Pp. 1-530]Heatwole, H.F. 1975 Attacks by sea snakes on divers. [Pp. 503-516]. In: Dunson, W.A.(Editor): The Biology of Sea Snakes. University Park Press, Baltimore [Pp. 1-530]Heatwole, H.F. 1977 Sea snakes, a contrast to other vertebrate divers. Journal of the SouthPacific Underwater Medicine Society, 1977: 35-3882

Australian Biodiversity Record, 2007 (8): 1-124Heatwole, H.F. 1977 Heart rate during breathing and apnea in marine snakes (Reptilia,Serpentes). Journal of Herpetology, 11(1): 67-76Heatwole, H.F. 1977 Sea snakes. Oceans, 1 (3): 126-131Heatwole, H.F. 1977 Adaptations of sea snakes. [Pp. 193-204]. In: Messel, H. and Butler,S.T. (Editors): Australian Animals and their Environment. Shakespeare Head Press, Sydney[Pp. 1-367]Heatwole, H.F. 1977 Serpentes (sea snakes). [Pp. 149-154]. In: Saenger, P. (Editor): TheGreat Barrier Reef. A Diver’s Guide. Poly-graphics, BrisbaneHeatwole, H.F. 1978 Adaptations of marine snakes. American Scientist, 66 (5): 594-604Heatwole, H.F. 1978 Sea snake attacks – Myth or menace. Skin-diving in Australia and NewZealand, 8: 40-45Heatwole, H.F. 1980 Seal predation on a sea snake. Herpetofauna, 11 (2): 24Heatwole, H.F. 1981 Role of saccular lung in the diving of the Sea Krait, Laticauda colubrina(Serpentes: Laticaudidae). Australian Journal of Herpetology, 1 (1): 11-16Heatwole, H.F. 1981 Temperature relations of some sea snakes. Snake, 13 (1): 53-57Heatwole, H.F. 1987 Sea Snakes. University of New South Wales Press, Kensington, Sydney[Pp. 1-148]Heatwole, H.F. 1999 Sea Snakes. University of New South Wales Press, Kensington, Sydney[Revised Edition]Heatwole, H.F. and Burns, G.W. 1987 Final Report for ANPWS Consultancy on Sea SnakePopulations 1984-87. Report for the Australian National Parks and Wildlife Service, Canberra[Pp. 1-64]Heatwole, H. and Cogger, H.G. 1993 Family Hydrophiidae. [Pp. 310-318]. In: Glasby, C.J.,Ross, G.J.B. and Beesley, P.L. (Editors): Fauna of Australia. Volume 2A. Amphibia andReptilia. Australian Government Publishing Service, CanberraHeatwole, H.F. and Dunson, W.A. 1987 Hematological parameters of some marine snakes.Journal of Experimental Marine Biology and Ecology, 113: 289-300Heatwole, H.F. and Finnie, E.P. 1980 Seal predation on a sea snake. Herpetofauna, 11 (2):24Heatwole, H. and Guinea, M.L. 1993 Family Laticaudidae. [Pp. 319-321]. In: Glasby, C.J.,Ross, G.J.B. and Beesley, P.L. (Editors): Fauna of Australia. Volume 2A. Amphibia andReptilia. Australian Government Publishing Service, CanberraHeatwole, H.F. and Poran, N.S. 1995 Resistances of sympatric and allopatric eels to seasnake venoms. Copeia, 1995 (1): 136-147Heatwole, H.F. and Seymour, R.S. 1975 Pulmonary and cutaneous oxygen uptake in seasnakes and a File Snake. Comparative Biochemistry and Physiology, (A) Molecular andIntegrative Physiology, 51: 399-405Heatwole, H.F. and Seymour, R.S. 1975 Diving physiology. [Pp. 289-327]. In: Dunson, W.A.(Editor): The Biology of Sea Snakes. University Park Press, Baltimore [Pp. 1-530]83

Australian Biodiversity Record, 2007 (8): 1-124Heatwole, H.F. and Seymour, R.S. 1976 Respiration of marine snakes. [Pp. 375-389]. In:Hughes, G.M. (Editor): Respiration of Amphibious Vertebrates. Academic Press, London [Pp.402]Heatwole, H.F. and Seymour, R.S. 1978 Cutaneous oxygen uptake in three groups of aquaticsnakes. Australian Journal of Zoology, 26 (3): 481-486 [see data on Laticauda]Heatwole, H.F. and Taylor, J. 1987 Ecology of Reptiles. Surrey Beatty and Sons, ChippingNorton [Pp. 1-325]Heatwole, H.F, Cogger, H.G. and Limpus, C.J. Class Reptilia. [Pp. 200-202]. In: Mather, P.and Bennett, I. (Editors): A Coral Reef Handbook. Third Edition. Surrey Beatty and Sons PtyLtd, Chipping Norton [Sydney]Heatwole, H.F., Heatwole, E. and Johnson, C.R. 1974 Shark predation on sea snakes.Copeia, 1974 (3): 780-781Heatwole, H.F., Minton, S.A.J., Taylor, R. and Taylor, V. 1978 Underwater observations onsea snake behaviour. Records of the Australian Museum, 31 (18): 737-761 [+Figures 1-12]Heatwole, H.F., Seymour, R.S. and Webster, M.E.D. 1979 Heart rates of sea snakes.Comparative Biochemistry and Physiology, A 62(2): 453-456Hecht, M.K., Kropach, C. and Hecht, B.M. 1974 Distribution of the yellow-bellied sea snake,Pelamis platurus, and its significance in relation to the fossil record. Herpetologica, 30(4):387-396Herre, A.W.C.T. 1942 Notes on Philippine sea snakes. Copeia, 1942 (1): 7-9Herre, A.W.C.T. and Rabor, D.S. 1949 Notes on Philippine sea snakes of the genusLaticauda. Copeia, 1949 (4): 282-284Hewer, A. and Mollison, B.C. 1974 Reptiles and Amphibians of Tasmania. [Pp. 51-60]. In:Tasmanian Yearbook No 8. 1974. Tasmanian Government Printer, HobartHibbard, E. 1975 Eyes and other sense organs of sea snakes. [Pp. 355-382]. In: Dunson,W.A. (Editor): The Biology of Sea Snakes. University Park Press, Baltimore [Pp. 1-530]Hibbard, E. and Lavergne, J. 1972 Morphology of the retina of the sea-snake, Pelamisplaturus. Journal of Anatomy, 112 (1): 125-136Hoffstetter, R. and Gasc, J.-P. 1969 Vertebrae and ribs of modern reptiles. [Pp. 201-310]. In:Gans, C., Bellairs, A.d'A. and Parsons, T.S. (Editors): Biology of the Reptilia. Volume 1.Morphology A. Academic Press, London [Pp. 373]Hoge, A.R. and Romano, S.A.R.W.D.L. 1971 Neotropical pit vipers, sea snakes, and coralsnakes. [Pp. 211-293]. In: Bucherl, W. and Buckley, E.E. (Editors): Venomous Animals andTheir Venoms. Volume 2. Venomous Vertebrates. Academic Press, New YorkHolmes, J.C. 1989 A redescription of Simhatrema simhai Chattopadhyaya, 1970 (Trematoda:Exotidendriidae), with comments on its pathogenesis in sea snakes (Serpentes:Hydrophiidae). Proceedings of the Helminthological Society of Washington, 56(2): 156-161Hediger, H. 1934 Beitrag zur Herpetologie und Zoogeographie Neu-Britanniens und einigerumliegender Gebiete.. Zoologische Jahrbucher, Abteilung fur Systematik,. 65(5/6):441-582Hildebrand, M. 1974 Analysis of Vertebrate Structure. John Wiley Sons, New York [Pp. 1-710]84

Australian Biodiversity Record, 2007 (8): 1-124Hilgendorf, F. 1876 Die Japanischen Schlangen. Mitt. Dt. Ges. Nat.-U. Volkerk. Ostasiens 1:29-34 [see p. 31 for original description of Hydrophis pelamoides - possible error forHydrophis pelamidoides Schlegel, 1837].Hin, H.K., Stuebing, R.B. and Voris, H.K. Population structure and reproduction in the marinesnake, Lapemis hardwickii Gray, from the west coast of Sabah. Sarawak Museum Journal,463-475Honders, J. (Editor) 1975 The World of Reptiles and Amphibians. Peebles Press, New YorkHopley, C.C. 1882 Snakes: Curiosities and Wonders of Serpent Life. Griffith and Farran,London [Pp. 1-614]Hoser, R.T. 1989 Australian Reptiles and Frogs. Pierson and Co., Mosman [Pp. 1-238]How, R.A. and Kitchner, D.J. 1997 Biogeography of Indonesian snakes. Journal ofBiogeography, 24: 725-735Hseu, T.H., Jou, E.D., Wang, C. and Yang, C.C. 1977 Molecular evolution of snake venomtoxins. Journal of Molecular Evolution, 10: 167-182Huang, K. and Zhu, Z. 1987 Investigation on the reptiles of the coastal sea of Liaoning. ActaHerpetologica Sinica. 6(1): 78-79 [In Chinese]Huang, W.-S. 1996 Sexual size dimorphism of sea snakes in Taiwan. Bulletin of the NationalMuseum of Natural Science, 7: 113-120Hudson, B.J. and Frohm, T. 1986 Bites due to sea snakes in the Ramu River system.Proceedings of the 22nd Annual Symposium of the Medical Society of Papua New GuineaHudson, B.J. and Pomat, K. 1988 Ten years of snake bite in Madang Province, Papua NewGuinea. Transactions of the Royal Society of Tropical Medicine and Hygiene, 82 (3): 506-508Huey, R.B. 1982 Temperature, physiology and the ecology of reptiles. [Pp. 25-91]. In: Gans,C. and Pough, F.H. (Editors): Biology of the Reptilia. Volume 12. Academic Press, New YorkHutchinson, M.N., Swain, R. and Driessen, M. 2001 Snakes and Lizards of Tasmania. Faunaof Tasmania Handbook No. 9 Fauna of Tasmania Committee, University of Tasmania [Pp. 1-64]Hvass, H. 1958 Reptiles of the World. Methuen, LondonHvass, H. 1964 Reptiles and Amphibians of the World. Methuen, LondonIneich, I. 1986 Histoire naturelle du serpent marin Pelamis platurus (Linne, 1766). BulletinSoc. etudes Oceaniennes, No 236, 20 (1): 1-10Ineich, I. 1988 Le serpent marin Pelamis platurus (Elapidae, Hydrophiinae): Bilan desconnaissances sur sa biologie et sa distribution; situation en Polynesie Orientale. L'AnneeBiologique, (ser. 4) 27 (2): 93-117Ineich, I. and Borsa, P. 2003 Hydrophis coggeri (Cogger's sea snake). Herpetological Review,34(4): 388Ineich, I. and Laboute, P. 2002 Sea Snakes of New Caledonia. IRD Editions, ParisIneich, I. and Rasmussen, A.R. 1997 Sea snakes from New Caledonia and the LoyaltyIslands (Elapidae, Laticaudinae and Hydrophiinae). Zoosystema, 19(2-3): 185-19285

Australian Biodiversity Record, 2007 (8): 1-124Ineich, I., Bonnet, X., Brischoux, F., Kulbicki, M., Séret, B. and Shine, R. 2007 Anguilliformfishes and sea-kraits: Neglected predators in coral-reef ecosystems. Marine Biology, 151:793-802Ingram, G.J. 1990 The works of Charles Walter de Vis, alias "Devis," alias "Thickthorn".Memoirs of the Queensland Museum, 28 (1): 1-34Ingram, G.J. and Covacevich, J.A. 1981 Frog and Reptile Type Specimens in the QueenslandMuseum, with a checklist of frogs and reptiles in Queensland. Memoirs of the QueenslandMuseum, 20 (2): 291-306Ingram, G.J. and Raven, R.J. (Editors) 1991 An Atlas of Queensland's Frogs, Reptiles, Birdsand Mammals. Queensland Museum, Brisbane [Pp. 1-391]Iredale, T. 1971 Serpents of the sea. Proceedings of the Royal Zoological Society of NewSouth Wales for 1969-1970: 19-24Irvine, F.R. 1954 Snakes as food for man. British Journal of Herpetology, 1 (10): 183-189Ishiyama, M., Yoshie, S. and Ogawa, T. 1983 Morphological study of fangs in the pelagic seasnake, Pelamis platurus. Shigaku. Odontology, 70(6): 1220-1229Iskandar, D.T. and Colijn, E. 2001 Checklist of southeast Asian and New Guinean Reptiles.Part I. Serpentes.. Bandung, Indonesia : Biodiversity Conservation Project (IndonesianInstitute of sciences-Japan International Cooperation Agency - The Ministry of Forestry), TheGibbon Foundation and Institut of Technology [Pp. 1-195; see p.139]Jacobs, M. 1985 Sea snake research sheds light on diving biology of marine reptiles.Smithsonian Institution Press, Washington, DC Research Reports No. 45: 5Jan, G. 1859 Prodrome d'une Iconographie descriptive des Ophidiens, et descriptionsommaire de nouvelles espèces de Serpents venimeux.. Reprinted from: Rev. Mag. Zool.(2)10 and (2)11-12, 1858 and 1859. Bouchard-Huzard, Paris [Pp. 1-32; see p. 23 for originaluse of Hydrophis schistotus which was in error for Hydrophis schistosus Daudin, 1803; see p.24 for use of Hydrophis pachycerios; see pl. (d) for use of Hydrophis lacepedei]Jan, G. 1859 Plan d'une Iconographie descriptive des Ophidiens, et description sommaire denouvelles espèces des Serpents. Rev. Mag. Zool. (2)11: 148-157 [see p. 149 for originaldescription of Disteira dumerilii; see p. 149 for original description of Platurus fischeri; see p.150 for original description of Hydrophis problematicus - as H. problematicus; see p. 148 fororiginal description of Hydrophis protervus - note : see also Jan, G. (1859). Prodrome d'uneIconographie descriptive des Ophidiens, et description sommaire de nouvelles espèces deSerpents venimeux.. Reprinted from: Rev. Mag. Zool. (2)10 and (2)11-12, 1858 and 1859.Paris : Bouchard-Huzard 32 pp. (pl. D), as H. protervus].Jan, G. 1863 Elenco Sistematico Degli Ofidi Descritti e Disegnati per l'Iconografia Generale.A. Lombardi, Milan [Pp. 1-143; see p. 109, pl. D for description of Hydrophis abbreviatus asH. abbreviatus, and for Hydrophis brevis as H. brevis; see p. 109 for original use of Hydrophisbicolor maculata - ?nomen nudum; see p. 110, livr. 39 pl. 5 fig. 2 for Hydrophis frontalis - asH. frontalis; see p. 110, livr. 41 pl. 1 for Hydrophis polydonta - H. polydonta] [see also Jan, G.(1859). Prodrome d'une Iconographie descriptive des Ophidiens, et description sommaire denouvelles espèces de Serpents venimeux. Reprinted from: Rev. Mag. Zool. (2)10 and (2)11-12, 1858 and 1859. Bouchard-Huzard, Paris Pp. 1-32]Jan, G. and Sordelli, F. 1872 Iconographie Générale des Ophidiens. Atlas. Milan : Jan andSordelli pp. 39-42 [see original description of Hydrophis propinquus; see livr. 40, pl. 3 fig. 1 foruse of Hydrophis bicolor maculata]Jan, G. and Sordelli, F. 1881 Iconographie Générale des Ophidiens. Atlas 1860-1881. Milan :Jan and Sordelli 50 pp.86

Australian Biodiversity Record, 2007 (8): 1-124Janzen, D.H. (Editor) 1983 Costa Rican Natural History. University of Chicago Press, Chicago[Pp. 1-816]Jayaram, K.C. 1974 Ecology and distribution of freshwater fishes, amphibia and reptiles. In:Ecology and Biogeography in India. Junk, The Hague [Monographiae Biologicae, Volume 23]Jeffries, W.B. and Voris, H.K. 1976 The identification of pedunculate barnicles found onMalaysian sea snakes with notes on their mode and site of attachment. HerpetologicalReview, 7 (2): 87-88 [Abstract only]Jeffries, W.B. and Voris, H.K. 1979 Observations on the relationships between Octolasmusgrayii (Darwin, 1851) (Thoracia, Cirripedia) and certain marine snakes (Hydrophiidae).Crustaceana, 37 (2): 123-132Johansen, K. 1964 Osmoregulering hos marine vertebrater. Fauna [Oslo], 17: 105-120Johnson, M.W. and Brinton, E. 1962 Biological species, water masses and currents. In: Hill,M.N. (Editor): The Sea. Interscience, New York [Volume 2]Johnson, R.G. 1956 The origin and evolution of the venomous snakes. Evolution, 10: 56-65Jong, J.K. de 1927 Reptiles from Dutch New Guinea. Nova Guinea [Leiden], (Zoologie) 15:296-542Jubb, R.A. 1983 A record of the yellow and black sea-snake, Pelamis platurus. Naturalist[Port Elizabeth], 27(3): 38Karlsson, E. 1978 Chemistry of protein toxins in snake venoms. Handbook of ExperimentalPharmacology, 52: 159-212Kasturirangan, L.R. 1952 The alloplacentation of the sea snake Hydrophis cyanocinctus(Daudin). Journal of the Indian Academy of Science, 3: 277-290Keast, J.A. 1959 The reptiles of Australia. [Pp. 115-135]. In: Keast, A., Crocker, R.L. andChristian, C.S. (Editors): Biogeography and Ecology in Australia. W. Junk, The Hague[Monographiae Biologicae, 8; Pp. 1-640]Keast, J.A. 1962 Vertebrate speciation in Australia: Some comparisons between Birds,Marsupials and Reptiles. [Pp. 380-407]. In: Leeper, G.W. (Editor): The Evolution of LivingOrganisms. A symposium to mark the centenary of Darwin's 'Origin of Species' and of theRoyal Society of Victoria held in Melbourne, December 1959. Melbourne University Press,MelbourneKeegan, H.L. and Macfarlane, W.V. (Editors) 1963 Venomous and Poisonous Animals andNoxious Plants of the Pacific Region. Pergamon, Oxford [Pp. 1-468]Keogh, J.S. 1998 Molecular phylogeny of Elapid snakes and a consideration of thebiogeographic history. Biological Journal of the Linnean Society, 63:177-203Keogh, J.S., Shine, R. and Donnellan, S.C. 1998 Phylogenetic relationships of terrestrialAustralo-Papuan Elapid snakes (Subfamily Hydrophiinae) based on cytochrome b and 16SrRNA sequences. Molecular Phylogenetics and Evolution, 10 (1): 67-81 [see alsoHerpetological Review 29 (4), 1998, p. 203]Key, JR., M.M., W.B. Jeffries and H.K. Voris. 1995 Epizoic bryozoans, sea snakes, and othernektonic substrates. Bulletin of Marine Science, 56(2): 462-474Kharin, V.E. 1981 A review of sea snakes of the genus Aipysurus (Serpentes, Hydrophiidae).Zoologicheskii Zhurnal, 60 (2): 257-264 [in Russian]87

Australian Biodiversity Record, 2007 (8): 1-124Kharin, V.E. 1983 A new species of the genus Hydrophis sensu lato (Serpentes:Hydrophiidae) from the north Australia shelf. Zoologicheskii Zhurnal, 62 (11): 1751-1753 [inRussian]; see p. 1751 for original description of Hydrophis macdowelli]Kharin, V.E. 1984 A review of sea snakes of the group Hydrophis sensu lato (Serpentes:Hydrophiidae). 3. The genus Leioselasma. Zoologicheskii Zhurnal, 63 (10): 1535-1546 [see p.1538 for original description of Leioselasma coggeri; see p. 1542 for original description ofLeioselasma czeblukovi]Kharin, V.E. 1984 Sea snakes of the genus Hydrophis sensu lato (Serpentes: Hydrophiidae).On the taxonomic status of the New Guinea H. obscurus. Zoologicheskii Zhurnal, 63 (4): 630-632Kharin, V.E. 1984 [Revision of sea snakes of the subfamily Laticaudinae Cope, 1879 sensulato (Serpentes, Hydrophiidae)]. In: Ecology and Faunistics of Amphibians and Reptiles of theUSSR and Adjacent Countries. Proceedings of the Zoological Institute, Acad. Sci. USSR[Leningrad], 124:128-139 [In Russian; see p. 134 for original description of Pseudolaticauda]Kharin, V.E. 1984 Sea snakes of the genus Hydrophis sensu lato (Serpentes, Hydrophiidae).On taxonomic status of the New Guinea H. obscurus. Zoologicheskii Zhurnal, 63 (4): 630-632Kharin, V.E. 1984 The first record of three species of sea snakes from Vietnam with a note onrare variety of Praescutata viperina. Biologiya Morya [Vladivostok], 1984(2): 26-30Kharin, V.E. 1985 A new species of sea snakes of the genus Enhydrina (Serpentes,Hydrophiidae) from waters of New Guinea. Zoologicheskii Zhurnal, 64 (5): 785-787 [inRussian; see original description of Enhydrina zweifeli]Kharin, V.E. 2006 An annotated checklist of sea snakes of Vietnam, with notes on a newrecord of the Yellow-lipped Sea Krait, Laticauda colubrina (Schneider, 1799) (Laticaudidae,Hydrophiidae). Russian Journal of Marine Biology, 32 (4): 223-228Kharin, V.E. and Cheblukov, V.P. 2006 On a new record of a poorly known and rare seasnake Aipysurus tenuis Lönnberg and Andersson, 1913 (Serpentes: Hydrophiidae) in thewaters of Australia. Russian Journal of Marine Biology, 32 (3): 194-197Kharin, V.E. and Czeblukov, V.P. 2006 A new revision of Sea Kraits of the FamilyLaticaudidae Cope, 1879 (Serpentes: Colubroidea). Russian Journal of Herpetology, 13 (3):227-241Kinghorn, J.R. 1926 A new genus of sea snake and other reptiles collected by H.M.A.S.Geranium during a cruise to north Australia. Proceedings of the Zoological Society of London,1926: 71-74 [see p. 71 for original description of Pseudodistira, and Pseudodistira horrida]Kinghorn, J.R. 1929 The Snakes of Australia. Angus and Robertson, Sydney [Pp. 1-198 +122 plates]Kinghorn, J.R. 1945 Australian poisonous snakes. Australian Museum Magazine, 8 (10): 325-330Kinghorn, J.R. 1951 Reptiles and amphibians. [Pp. 774-788]. In: The Australian JuniorEncyclopaedia. Georgian House, MelbourneKinghorn, J.R. 1956 The Snakes of Australia. Angus and Robertson, Sydney [2nd Edition;first published 1929; note: see also reprinting of 1964]Kinghorn, J.R. 1968 Reptiles. [Pp. 1-125]. In: Animals of the World: Australia. HamlynPublishing, Sydney88

Australian Biodiversity Record, 2007 (8): 1-124Kinghorn, J.R. and Kellaway, C.H. 1943 The Dangerous Snakes of the South West PacificArea. Railway Printing Works, Melbourne [Pp. 1-43][Kinghorn, J.R. and Kellaway, C.H.] 1944 The Dangerous Snakes of the South West Pacific.Australian Military Forces, Melbourne [Note: Authors names were not printed on the work, butadded later by stamp; This is the rare military edition that unintentionally contained classifieddetails of the Australian Military Forces and so had to be withdrawn from distribution; Pp. 1-43]Klauber, L.M. 1935 The feeding habits of a sea snake. Copeia, 1935 (4): 182Klawe, W.L. 1964 Food of the Black and Yellow Sea Snake, Pelamis platurus, fromEquadorian coastal waters. Copeia, 1964 (4): 712-713Klemmer, K. 1963 Liste der rezenten Giftschlangen - Elapidae, Hydrophiidae, Viperidae undCrotalidae. [Pp. 255-464]. In: Die Giftschlangen der Erde Wirkungen und Antigenitat der GifteTherapie von Giftschlangenbissen. Behringwerk-Mitteilungen, N.G. Elwert Universitats,Marburg [Pp. 1-464; see use of Hydrus valakadyn Boie, 1827]Klemmer, K. 1966 Observations on the biology of sea snakes - Hydrophiidae -with remarkson their systematics. Memorias do Instituto Butantan, Simp.Internac.33 (1): 101-103Klemmer, K. 1967 Observations on the sea snake, Laticauda laticaudata in captivity.International Zoo Yearbook, 7: 229-231Klemmer, K. 1968 Classification and distribution of European, North African, and North andWest Asiatic venomous snakes. [Pp. 309-325]. In: Bucherl, W., Buckley, E.E. and Deulofeu,V. (Editors): Venomous Animals and their Venoms. Volume 1. Venomous Vertebrates.Academic Press, New YorkKlemmer, K. 1968 Methods of classification of venomous snakes. [Pp. 275-283]. In: Bucherl,W., Buckley, E.E. and Deulofeu, V. (Editors): Venomous Animals and their Venoms. Volume1. Venomous Vertebrates. Academic Press, New YorkKo, R.C., Valentine, L. and Duggan, R.T. 1960 Note on the biology of Hydrophitremagigantica Sandars, 1960 (Trematoda: Hemiuridae) from the lung of sea snakes (Hydrophiscyanocinctus). Canadian Journal of Zoology, 53: 1181-1184Kochva, E. 1987 The origin of snakes and evolution of the venom apparatus. Toxicon, 25 (1):65-106Kooyman, G.L. 1972 Deep diving behaviour and effects of pressure in reptiles, birds andmammals. Symposia of the Society for Experimental Biology, 26: 295-311Kraus, F. and Allison, A. 2004 New records of reptiles and amphibians from Milne BayProvince, Papua New Guinea. Herpetological Review, 35: 413-418Krefft, J.L.G. 1869 The Snakes of Australia; an Illustrated and Descriptive Catalogue of All theKnown Speices. Thomas Richards, Government Printer, Sydney [Pp. i-xxv, 1-100; see p. 92for original descriptions of Emydocephalus and Emydocephalus annulatus; see p. 93 fororiginal description of Emydocephalus tuberculatus; Note: originally printed with Platescoloured (200 copies) and not coloured (500 copies); note also that a facsimile of thecoloured version was also published by Jeanette Covacevich, Lookout Publications, Brisbane(1984)]Kropach, C. 1971 Sea snake (Pelamis platurus) aggregations on slicks in Panama.Herpetologica, 27(2): 131-135Kropach, C. 1971 Another color variety of the sea snake Pelamis platurus from Panama Bay.Herpetologica, 27 (3): 326-32789

Australian Biodiversity Record, 2007 (8): 1-124Kropach, C. 1972 Pelamis platurus as a potential colonizer of the Caribbean Sea. Bulletin ofthe Biological Society of Washington,2: 267-269Kropach, C. 1975 The Yellow-bellied Sea Snake, Pelamis, in the eastern Pacific. [Pp. 185-213]. In: Dunson, W. (Editor): The Biology of Sea Snakes. University Park Press, Baltimore[Pp. 1-530]Kropach, C. and Soule, J.D. 1973 An unusual association between an ectoproct and a seasnake. Herpetologica, 29 (1): 17-19Kuntz, R.E. 1963 Snakes of Taiwan. Quart. J. Taiwan Mus. 16(1-2): 1-79Kuroda, N., et al 1958 Encyclopaedia Zoologica Illustrated in Colours. Volume 1. Mammals,Birds, Reptiles and Amphibians. Hokuryu-Kan Publishing Co., TokyoLacépède, B.G.E. 1804 Mémoire sur plusieurs animaux de la Nouvelle-Hollande dont ladescription n'a pas encore été publiée. Ann. Mus. Natl Hist. Nat. Paris 4: 184-211 [see p. 198,210 for original description of Leioselasma striata - there is some confusion over the decisionby McDowell (1972) to regard this species as a synonym of Hydrophis cyanocinctus Daudin,1803, a move rejected by Cogger et al (1983) - as cyanocinctus was not known fromAustralian waters. McDowell had suggested that the specimen represented by pl. 18, figs 4-5in Schlegel, H. (1837). (ex Merrem ms) - be made the Neotype of Leioselasma striataLacépède,1804; see p. 199, 210, pl. 57 fig. 2 for original description of Disteira doliata; see p.210 for original description of Disteira; see p. 210 pl. 56 fig 3 for original descriptions ofAipysurus and Aipysurus laevis]Lading, E.A. 1987 A Study of the Sea Snake (Laticauda colubrine) in Kalampunian DamitIsland, Sabah. BSc Thesis, Universiti Kebangsaan Malaysia, Kapus SabahLading, E.A., Stuebing, R.B. and Voris, H.K. 1991 A population size estimate of the yellowlippedsea krait Laticauda colubrina on Kalampunian Damit Island, Sabah, Malaysia. Copeia,1991 (4): 1139-1142Laidlaw, F.F. 1901 List of a collection of snakes, crocodiles and chelonians from the MalayPeninsula, made by members of the “Skeat Expedition”, 1899-1900. Proceedings of theZoological Society of London, 2 (2): 575-586Lamb, F. and Hunter, W.K. 1907 On the action of venoms of different species of poisonoussnakes on the nervous system. VI. Venom of Enhydrina valakadien. Lancet, 2: 1017-1019Lanza, B. 1954 Su un esemplore di Pelamis platurus (L.) catturato in un fiuime della piccolaAndaman. (Serpentes: Hydrophiidae). Monit. Zool. Ital. 62: 67-70Lanza, B. and Boscherini, S. 200 The gender of the genera Podarcis Wagler 1830(Lacertidae), Pelamis Daudin 1803 (Hydrophiidae) and Uropeltis Cuvier 1829 (Uropeltidae).Tropical Zoology, 13(2): 327-329Laurenti, J.N. 1768 Synopsin Reptilium. Vienna: J. Thomae 214 pp. [see p. 109 for originaldescriptions of Laticauda and Laticauda scutate]Lavery, H.J. 1975 Fauna; Reptiles, Birds and Mammals. [Pp. 34-43]. In: May, O.M. (Editor):Queensland Year Book, No 35. Queensland Government Printer, BrisbaneLee, C-Y. (Editor) 1979 Handbook of Experimental Pharmacology. Volume 52. SnakeVenoms. Springer-Verlag, BerlinLee, C.Y. and Chen, Y.M. 1975 Species differences in reversibility of neuromuscular blockadeby Elapid and sea snake neurotoxins. Toxicon, 13 (2): 106-107 [Abstract only]90

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