Progress 10/01/02 to 09/30/07

Outputs
OUTPUTS: Outputs have included:(1) publication of a number of scientific papers in peer-reviewed journals,including one large monograph, (2) participating in an expedition with the Dept. of Customs, Marine Division of the Australian government to investigate the cause of decline of two species of sea snake on Ashmore Reef; this resulted in a report to the government, (3) providing information to BBC(British Broadcasting Company)and to National Geographic in regard films they are producing on sea snakes,(4) field work in New Guinea and Indonesia to collect DNA samples from sea snakes which was contributed to Dr. Ulrich Kuch in Germany for a study he is carrying out in collaboration with myself and several other herpetologists on the phylogeography of the genus Laticauda. PARTICIPANTS: Institutions that were involved in this project were: (1) Australian Government Dept. of Customs, Marine division, (2) James Cook Unversity, Townsville, Queensland, Australia, (3) University of New England, Armidale, NSW Australia, (4) Charles Darwin University, Darwin, NT, Australia and (5) The Australian Museum (Sydney Australia). Institutions that benefited directly from data arising from this project were (1) National Geographic Society and (2) the British Broadcasting Corporation. Individuals involved in collaborative research on this project were (1) Dr. Vimoksalehi Lukoschek (James Cook University), Dr. Helene Marsh (James Cook University), Dr. Michael Guinea (Charles Darwin University),Dr. Ulrich Kuch, (Senckenberg Institute, Germany), Dr. Harold Cogger (The Austgralian Museum) Ms Myriam Preker (Brisbane, Queensland, Australia), Dr. Naomie Poran(NCSU, now gone), Dr. Peter King(Francis Marion University, South Carolina). TARGET AUDIENCES: The target audiences were herpetologists with an interest in snakes, managers of marine parks with an interest in conserving sea snakes, and health officials concerned with snake bite. PROJECT MODIFICATIONS: The first part of the project was concerned mostly with the resistance by prey (frogs, eels) to snake venoms and the effect of venom upon them. These topics were reviewed in the yearly reports and contained in the publications listed in those reports. The later part of the project shifted more to the ecology,biogeograpy and systematics of sea snakes and their role in natural ecosystems.

Impacts
This work has contributed to a better understanding of the interaction of venomous snakes and their prey. Bullfrog tadpoles have little resistance to but develop it at metamorphosis. Cottonmouth venom kills, not by previously believed means, but by collapsing the lungs and causing asphyxiation. Sea snakes were found to vary geograpically in their venom characteristics necessitating that anti-venoms be developed for local populations,not for a widespread species as a whole. Venom also serves in digesting prey. At high emperatures this is not so important as digestion proceeds adequately without the assistance of venom. At low temperataures, however, venom increases both the rate and efficiency of digestion; venomous snakes can thus function at temperatures too low for non-venomous snakes to be able to process food. This project also contributed to an understanding of the taxonomy, distribution and ecology of a major taxa of venomous snakes and has contributed to understanding the dynamics of their ecology. The work on the Great Barrier Reef developed a statistical methods of handling GIS data to assess factors affecting colonization of new reefs by snakes.

Publications

• Cogger, H. G. andH.F. Heatwole 2006. Laticauda frontalis (de VIs, 1905) and Laticauda saintgironsi sn. sp. from Vanuatu and New Caledonia (Serpentes: Elapidae: Laticaudinae)--a New Lineage of Sea Kraits. Records of the Australian Museum 58: 245-256.
• Lukoschek, V., H. Heatwole, A. Grech, G. Burns and H. Marsh 2007. Distribution of two species of sea snakes, Aipysurus laevis and Emydocephalus annulatus, in the southern Great Barrier Reef: metapopulation dynamics, marine protected areas and conservation. Coral Reefs, 17 pages in press. Heatwole,H.,P.King and J.Levine 2007 Laterality in coiling behavior in viperid snakes,new data and a reinterpretation. Laterality,in press.
• Heatwole, H. and V. Lukoschek. 2007 Sea Snakes and Sea Turtles. Chapter in Handbook of the Great Barrier Reef,in press.

Progress 10/01/05 to 09/30/06

Outputs
In the past year research was completed on the geography and taxonomy of venomous sea kraits and a new phase of the study was begun. This involved collaboration with Dr. Ulrich Kuch from Germany in a study of the molecular taxonomy of the genus Laticauda. Previous work was written for publiction and new field work conducted in Australia and western New Guinea on live populations of Laticauda colubrina, one of the key species. The findings were that this species, in fact, contains five cryptic species that differ only slightly, but distinctly, in morphology but which have different venom characteristics. One species is widespread from the Andaman Islands to New Guinea, another also ranges widely but from Sabah northward to Japan and the remaining three have limited distributions, one in Vanuatu, one in New Caledonia and the third in southern Papua. A second project that was completed was a study of the change in distribution of sea snakes on the reefs of the Swain Reef complex on the Great Barrier Reef. This was the culmination of research I have conducted over 30 years and involved comparisons of the local distribution of two species of snakes at various time intervals. Some patch reefs sustained populations throughout this time, others lost their resident populations and still other reefs that were initially barren later developed populations of sea snakes. This project is now being expanded to a greater number of species and throughout the length of the Barrier Reef with the collaboration of Harold Cogger and Vimoksalehi Lukoscheck. We are defining the habitat characteristics of the various species of snakes inhabiting the reefs.

Impacts
The impact of recent work is two-fold: protection from snake bite and zoning the reef for planned use. The discovery that there are cryptic taxa within what was previously considered a single taxon and that the venoms differs among them, means that effective treatment for snake bite in one geographic region may not be so effective elsewhere and more detailed studies of the characteristics of individual venoms should be carried out. The significance of the dynamism of the populations of sea snakes on the Great Barrier reef is that it implies a totally different conservation plan than that true of statically distributed ones. Also, the planned uses of individual reefs (e.g., for resorts) by the Great Barrier Reef Marine Authority must take into account that reefs that are presently snake-free may not remain so. The third phase of the study, assessing habitat characteristics, will aid planning because it will improve the accuracy of the prediction as to which species are likely to inhabit particular reefs over the long term.

Publications

• Stuart, B. L., H. Heatwole and T. F. Lian. 2006. Record of the little-known Rana nigrotympanica Dubois, 1992 (Amphibia: Ranidae) from northern Laos. Hamadryad 30: 108-113.
• Cogger, H. G. and H. F. Heatwole. 2006. Laticauda frontalis (De Vis, 1905) and Laticauda saintgironsi n. sp. from Vanatu and New Caledonia (Serpentes: Elapidae: Laticaudinae)---a new lineage of sea kraits?. Records of the Australian Museum 58: 245-256.
• Lukoschek, V., H. Heatwole, A. Grech, G. Burns and H. Marsh. 2007. Distribution of two species of marine snakes, Aipysurus laevis and Emydocephalus annulatus, in the southern Great Barrier Reef: metapopulation dynamics, marine protected areas and conservation. Marine and Freshwater Research, in press.

Progress 10/01/04 to 09/30/05

Outputs
Progress in the 2004-2005 year consisted of work on venomous sea snakes. One month was spent at James Cook University in Australia collaborating with a colleague, Vimoksalehi Lukoschek, on continuing two projects. One was assembling new locality records of species in eastern Australia to provide a better understanding of habitat selection and local distribution. The other dealt with the changes in distribution of the olive seasnake, Aipysurus laevis, on the Swain Reefs. I had plotted the distribution of that species on the small patch reefs about 20 years ago, and recently we re-surveyed reefs to ascertain which ones had lost this species and to which new ones it had had spread. These projects are nearly finished and we mainly collated the results and have nearly finished writing the papers. The main thrust of my recent research, however, was the completion and publication of a major monograph of 136 pages on the distribution and geographic variation of the venomous sea krait, Laticauda colubrina. This species is widespread, ranging from the Andaman islands through Burma, Thailand, Malaysia, Indonesia, New Guinea and souheastward through the island chains of the Solomon Islands, Vanuatu, New Caledonia, Fiji and Samoa, and northward through the Philippines, Palau, Taiwan to Japan. I carried out an extensive morphological analysis and discovered that this is in fact a complex of closely related taxa, including one ranging from the Philippines to Japan, one from the Andaman Islands to the Solomon Islands, and demes in the remaining eastern islands. The population in southern Papua was quite distinct and was described as a new species. A second new species from New Caledonia is being described in a paper in press. The Palauan population is enigmatic in that it has distinct characteristics on the one hand, but shares others with both the Philippine-Japan one and the Andaman-New Guinean one. Projected work includes a DNA analysis of each of these separate taxa to ascertain the degree of genetic relationship among them. To that end I began assembling tissue samples from critical areas. In September I went on an expedition to Palau to collect fresh tissue for this purpose. I now have fresh tissue from most of the eastern part of the range, Thailand, and Palau. I am now collaborating with Ulrich Kuch from Germany on the DNA analysis of these and other sea snake taxa.

Impacts
Japanese and French venom chemists have found that the characteristics of sea snake venoms vary geographically within widespread species, and consequently antivenoms prepared from one locality may not be maximally effective in other localities. By ascertaining the geographic variation among closely related populations, I have identified taxonomic units that are somewhat different from each other and therefore potentially less cross-receptive to antivenoms. This information will allow selection or development of antivenoms most appropriate for use in specific localities, and will allow assessment of the geographic extent a particular one will be effective.

Publications

• Heatwole, H., S. Busack and H. Cogger 2005 Geographic variation in sea kraits of the Laticauda colubrina complex (Serpentes: Elapidae: Hydrophiinae: Laticaudini). Herpetological Monographs 19: 1-136.
• Cogger, H. G. and H. Heatwole. 2006 Laticauda frontalis De Vis and Laticauda saintgironsi sp. nov. representing a new clade of sea kraits from the Republic of Vanatu and New Caledonia (Serpentes: Laticaudidae). Memoirs of the Queensland Museum, in press.

Progress 10/01/03 to 09/30/04

Outputs
The work on the effect of viper venom on the liver and kidneys of bullfrogs, completed last year and included in last year's report has now been published (see No. 43 below). Interpretation of the damage to tissue required color photographs in the publication and consequently, the Russian Journal of Herpetology was chosen as it routinely publishes in color. It was found that in culture liver is susceptible to venom, and the nuclei of liver cells are destroyed. In the live frog, the serum intervenes and protects liver nuclei. The kidney is more susceptible and in the live animal envenomation destroys the renal capillaries, releasing blood into the tissues of the kidney. Failure of neither the kidneys , nor the liver is the immediate cause of death. Rather, the venom in a live frog destroys the surfactant on the surface of the lung causing it to collapse. The work on lungs was reported last year as complete and ready to write for publication. However, a recent medical case in North Carolina in which a human patient suffered lung collapse following a cottonmouth bite led me to hold up that report. I am exploring this case with a view to summarizing it along with my experimental results in collaboration with the medical doctor in attendance. A major achievement in 2004-2005 was the completion of a monograph on the geographic variation of sea snakes of the Laticauda colubrina complex. This is a project that has been in progress for many years. The manuscript was 186 pages long, single-spaced, and contained 35 tables and 27 figures. It was submitted to Herpetological Monographs and has been received back for revisionary changes. Those will be completed by mid-January 2005. Another project that has been completed is a comparison of the susceptibility of toads and frogs to viper venom. It was found that toads, which are protected from predation by vipers by virtue of toxin glands in the skin, have little resistance to viper venom, whereas frogs, that lack protective cutaneous toxins in the adults, are much more resistant to snake venom. Finally, I continued editing volumes in the series 'Amphibian Biology', one of which appeared in 2004. Another volume, now being edited, deals with the conservation and decline of amphibians, including a chapter on the role of predation, including that by venomous snakes.

Impacts
The studies on viper venom have indicated the target organs that they attack and should provide insight into better ways of dealing with human victims of snake bite.

Publications

• Green, J. M., H. Heatwole, B. Black and N. Poran. 2004. Effect of the venoms of two viperid snakes, the copperhead (Agkistrodon contortrix) and the cottonmouth (Agkistrodon piscivorus piscivorus) on the liver and kidney of the bullfrog, Rana catesbeians. Russian Journal of Herpetology 11: 21-29.
• Stuart, B. and H. Heatwole. 2004. A new Philautus (Anura: Rhacophoridae) from northern Laos. Asiatic Herpetological Research 10: 1-5.
• Heatwole, H. (editor). 2004. Endocrinology, volume 6 in 'Amphibian Biology', Surrey Beatty & Sons, Chipping Norton.

Progress 10/01/02 to 09/30/03

Outputs
In the past year research was completed on the effect of viper venoms on the liver, kidneys, and lungs of bullfrogs. It was found that in culture liver is susceptible to the venom and the nuclei of cells are destroyed. In the live frog, the serum intervenes and protects liver nuclei. The kidney is more suscepible and in the live animal envenomation destroys the renal capillaries, releasing blood into the tissues of the kidney. Failure of neither the kidneys nor the liver is the immediate cause of death. Rather, the venom in a live frog destroys the surfactant on the surface of the lung causing it to collapse, and the animal dies of asphyxiation before the renal or hepatic effects become lethal. The paper on liver and kidney has been submitted, and the one on the lung is in preparation. Since my previous work showed that tadpoles were more susceptible to venom than adult frogs, a project was started to compare the effects of envenomation on these two stages. The experiments have been concluded and preparation of the histological slides are in preparation. Freshwater eels are a potantial prey for cottonmouth moccasins and a study was initiated to ascertain whether these freshwater eels have the unusually high resistance to the venom of their predators as do marine eels to their venomous predators (sea snakes). Again, the envenomations are complete and the histology is in progress. A study of the expansion of sea snakes southward along the Great Barrier Reef, with their concomitant impact on the prey fish in the reef communities has been completed and a paper is in press. Many animals favor one or the other side of their body (handedness) with a concomitant asymmetry in the brain. Snakes lack appendages and might be expected to also lack laterality in the brain. Previous reports suggested laterality, but my work has shown that it does not occur in the direction of coiling of snakes. I have submitted a review re-assessing this topic for copperheads and cottonmouths. I am writing up a paper on the accuracy with which spitting cobras hit the eyes of their victims with venom. They have maximal accuracy at 2 meters distance. Finally, a survey of the reptiles of the New England District in Australia, including the venomous snakes, has been completed and published.

Impacts
The studies on viper venom have indicated the target organs that they attack and should provide insight into better ways of dealing with human victims of snake bite. The study of spitting cobras provides insight into better protection from them. The other projects assess the ecological role of venomous snakes in their environment

Publications

• Heatwole, H., J. de Bavay and P. Webber. 2003. Faunal survey of New England. V. The lizards and snakes. Memoirs of the Queensland Museum 49: 299-325.

Progress 10/01/01 to 09/30/02

Outputs
The following research projects were addressed in 2001-2002. (1) Effect of copperhead and cottonmouth venom on target organs of prey bullfrogs. Bullfrogs were injected with graded doses of venom and later autopsied and microscopic slides made of various organs. Work in previous years investigated effects on muscle and liver and this year emphasis was placed on kidney and lungs. It was found that venom concentrates in the lungs of the frogs, where it destroys the surfactants and the cells that generate surfactants, with the result that the surface tension increases and the lungs collapse. Lungs in envenomated frogs were half the size of those in controls. Lung collapse is probably the cause of death in the prey, a new finding. The kidney is also damaged and shows signs of internal hemorrhage. The results are being written for publication. (2) Feeding of water cobras. This project is continuing. Videos have been made of their feeding on fish underwater. It was found that the snakes search for fish in crevices and identify them by smell. The snakes hold the prey until the venom immobilizes the fish, then they manipulate the prey until the head is reached; the fish is swallowed headfirst. I am now beginning a second phase of this study, an investigation of the adaptations that allow these snakes to hunt underwater. I am measuring their voluntary diving times and will relate that to the ratio of pulmonary and cutaneous respiration, diving bradycardia, and anaerobic metabolism. These studies are just beginning. (3) Digestion of envenomated prey. The part described last year has now been completed for the king snake. For comparative purposes, the project has now been extended to the copperhead. (4) Ecology of the Puerto Rican boa. The field work on movements and home ranges has been completed, and a laboratory study has begun. This involves placing the snakes in thermal gradients to determine preferred temperatures, and will aid in interpreting the results of the field work. The projects on accuracy of spitting cobras and on prey preferences of cottonmouths described last year have been completed and are being written for publication.

Impacts
These studies have enhanced the understanding of the adaptive function of venoms in the feeding of snakes, and has elaborated the way food is captured in snakes of different life styles. It also can lead to better ways of avoiding damage from spitting cobras. A better understanding of the different susceptibilities of different organs can lead to improved means of treating snake bite.

Publications

• Heatwole, H. 2003. Quadrat sampling of reptiles. A Chapter in: "Methods for Measuring and Monitoring Reptile Biodiversity", edited by M. S. Foster; Smithsonian Institution. In press.

Progress 10/01/00 to 09/30/01

Outputs
The following research projects were address in 2000-2001: (1) Treatment of cultured organ slices of prey frogs and of donor snakes with snake venom. The histological slides of venom damage to tissues have been analyzed by a grid analysis of videographed sections. The results have been written and will be submitted shortly. In brief, the results indicate that different species vary in their relative susceptibity to venom and cottonmouth venom is more potent than copperhead venom. The liver in prey species is highly susceptible to venom damage in ccomparison to skeletal or heart muscle. (2) The accuracy of spitting cobras. This project has been finished and being written for publication. The snakes are accurate in hitting a victom's eyes at a distance of 2 m because the venom spreads out as a mist and covers a large area. At closer distances the venom is still in a stream and consequently is less accurate. (3) Prey preferences of cottonmouth moccasins. This work was completed by last year but has been summarized but not yet written for publication. The findings were that fish and frogs are preferred by neonate animals but adults include more mammals. (4) Digestion of envenomated prey. The experiments summarized last year have been extended. Last year it had been discovered that venom aids digestion. The new results reveal that venom assists digestion at low temperatures whereas undigested non-envenomated prey are likely to putrify at low temperatures rather than be digested. (5) Feeding by water cobras. This project is in early stages. Water cobras are placed in large tanks of water and fed fish. Their method of stalking, capturing, and consuming fish are recorded on video and will be analyzed to compare with these activities in sea snakes. (6) Movements and home ranges of the Puerto Rican boa. Boas are implanted with telemeters and the movements of local inhabitants compared with those of snakes introduced from othe areas. The latter move long distances in the direction of their orginal location. Juveniles move greater distances than adults. This project has several more years to go.

Impacts
These studies have enhanced the understanding of the adaptive function of venoms in the feeding of snakes, and has elaborated the way food is captured in snakes of different life styles. It also can lead to better ways of avoiding damage from spitting cobras. A better understanding of the different susceptibilities of different organs can lead to improved means of treating snake bite.

Publications

• Burns, G and H. Heatwole 2000. Growth, sexual dimorphism, and population biology of the olive sea snake, Aipysurus laevis, on the Great Barrier Reef of Australia. Amphibia-Reptilia 21: 289-300.

Progress 10/01/99 to 09/30/00

Outputs
A large number of organ slices of prey frogs and of donor snakes have been treated with various doses of venoms for varying periods of time, and histological slides made for evaluation of direct digestive effect of venom on target tissues. These are now being assessed by quantitative grid analysis of video projections of the slides. Also, studies have been carried out on the accuracy with which spitting cobras can hit the eyes of model humans at different distances. At short distances, the venom is still in a compact stream and accuracy is low. However, at 2 meters, the venom spreads out into a mist and targets a larger area that always encompasses the eyes. Paradoxically the animal is more accurate at the longer distance. At short distances the venom acts as a "rifle" and hits only a small area; at longer distances it functions as a "shotgun" and spreads out over a larger area. Work has also been completed and written up on the prey preferences of cottonmouth moccasins. Fish and frogs are the preferred prey of neonate snakes. Mammals and invertebrates are seldom eaten. By contrast, adults eat significant numbers of mammals. Several experiments have been completed on the digestion of envenomated prey versus nonenvenomated prey. Digestion of envenomated prey is much more efficient although the passage time is not greatly shortened compared to nonenvonomated prey.

Impacts
These studies have enhanced the understanding of the adaptive function of venoms in the feeding of snakes.

Publications

• Heatwole, H., N. Poran, and P. King. 1999. Ontogenetic changes in the resistance of bullfrogs (Rana catesbeiana) to the venom of copperheads (Agkistrodon contortrix contortrix) and cottonmouths (Agkistrodon piscivorus piscivorus). Copeia 1999: 808-814.
• Burns, G and H. Heatwole 2000. Growth, sexual dimorphism, and population biology of the olive sea snake, Aipysurus laevis, on the Great Barrier Reef of Australia. Amphibia-Reptilia 21: 289-300.

Progress 01/01/99 to 12/31/99

Outputs
The work during 1999 consisted of continuing investigations of (1) the histological effects of snake venoms on thin-slices of organs of prey species and of those of the snake species itself, (2) innate prey preferences by cottonmouth neonates, (3) ecology of sea snakes, (4) periodicity and seasonality of venom production and venom characteristics in cottonmouths, (5) accuracy of spitting cobras, and (6) comparative rates of digestion in envenomated and non-envenomated prey. In addition, a new predator-prey system was added for study: a desert mouse and its scorpion prey. Part (1) is a long-term project and more data were accumulated. For (2) the main finding was that neonate snakes that have not previously fed strongly prefer fish and frogs to mammalian (mice) prey, but as they age, they begin to widen their preferences. Project (3) was been nearly completed and all the data from the past few years, as well as a review of all sea snake literature has been published as a book. Project (4) is still in progress. The most important recent finding was that in spring when frogs come to water to breed and are most accessible to the snakes, the snakes' venom characteristics change. I am now investigating whether those changes result in specifically a frog toxin. The most important finding for project (5) was that at 2 meters distance the venom from spitting cobras spreads out and covers a wide area, increasing the likelihood of entering the eyes of molestors. At close range where they can defend themselves by biting, they eject a steam, but with much less accuracy. The results from project (6) and the scorpion work are too premature for summarizing.

Impacts
The impact is that the results of these projects combine to provide a better understanding of predator-prey interactions and the coevolution of predators with their prey. The practical implication is that a better understanding of seasonal changes in characteristics of the venom of cottonmouths may lead to better ways of dealing with snake bite.

Publications

• Heatwole, H. 1999 Seasnakes. Universitiy of New South Wales Press, Sydney
• Heatwole, H., Poran, N.and King, P. 1999 Ontogenetic changes in the resistance of bullfrogs (Rana catesbeiana) to the venom of copperheads (Agkistrodon contortrix contortrix) a dn cottonomouths (Agkistrodon piscivorus piscivorus). Copeia 1999: 808-814.

Progress 01/01/98 to 12/31/98

Outputs
Cottonmouths have been continued to be milked of venom at monthly intervals and HPLC used for separation of the venom into different components. The results show that in the spring some components of the venom increase in proportion relative to others. This may reflect a change in diet primarily to frogs, rather than other vertebrates, at that time of year. Food preferences of neonate cottonmouths have been tested by offering them a choice of foods for their first feeding after they were born. They overwhelmingly prefer fish to mice, and frogs to mice, but there is very little difference in their preferences between fish and frogs. Older cottonmouths are now being tested to see whether those preferences change as they mature. Bullfrog tadpoles are very sensitive to cottonmouth and copperhead venoms, but at metamorphosis (when they are more subject to predation by snakes) they develop an innate resistance to both venoms, which they maintain until they are too large to be eaten by most snakes. Then the resistance declines, but not to the level of sensitivity of tadpoles. By contrast, toads have skin secretions that repel cottonmouths from eating them; accordingly toads do not need a high tolerance to cottonmouth venom and remain sensitive to it throughout life. Bullfrogs in sympatry with cottonmouths are more resistant to cottonmouth venom than are allopatric bullfrogs which have not coevolved with cottonmouths. A study is underway that is comparing the histological effects of cottonmouth and copperhead venoms on the various tissues of prey amphibians at various stages of development. The sensitivity of eels to sea snake venom was tested. Sympatric species of eels had unusually high resistances (52 times the human lethal dose) whereas allopatric ones were highly sensitive, indicating coevolution of prey and predator. Rates of digestion in rat snakes swallowing mice that had not been envenomated compared to those in which the rats had been envenomated revealed that venom serves a digestive function as well as playing a role in subduing prey. This was followed by an analysis of the enzymic action of the salivas of a wide variety of non-venomous species of snakes, and of the salivas and venoms of a wide variety of venomous ones. There were great differences among species. All of the above research contributes to a better understanding of the interaction of venomous predators with their prey, and the counter-adaptations of prey to their predators.

Impacts
(N/A)

Publications

• Duellman, W. E. and H. Heatwole 19989. Habitats and adaptations. Chapter (pp. 30-35) in: Reptiles and Amphibians (Editors, H. G. Cogger ad R. G. Zweifel), Weldon Owen Inc., McMahons Point.
• Stuart, B., W. G. Groom Jr, and H. Heatwole. 1999. Hypersensitivity of some lizards to pilocarpine. Herpetological Review, in press
• Heatwole, H. 1999. Ses Snakes. Universitiy of New South Wales Press, Sydney, about 200 pages, in press.
• Heatwole, H. 1997. Marine snakes: are they a sustainable resource? Wildlife Society Bulletin 116: 175-184
• Heatwole, H. 1998. Diffuse cutaneous and muscular sensory systems: mechanoreception, thermoreception, nociception, chemoreception and kinesthetic sense. Pp. 936-953 in: Amphibian Biology, volulme 3 Sensory Perception (Editors H. Heatwole and E. Dawley), Surrey Beatty & Sons, Chipping Norton
• Burns, G. and H. Heatwole 1998. Home range and habitat use of the olive sea snake, Aipysurus laevis, on the Great Barrier Reef, Australia. Journal of Herpetology 32: 350-358.
• Heatwole, H. and J. Powell 19989. Resistance of eels (Gymnothorax) to the venom of sea kraits (Laticauda colubrina): a test of coevolution. Toxicon 36: 619-625.

Progress 01/01/96 to 12/30/96

Outputs
Field work is continuing on collecting ants across ecological gradients. Transects have been sampled in areas of the Gobi Desert of China, the lowlands of Laos and across an altitudinal cross-section of Madagascar. Voucher specimens of all different species in an area are collected and brought to the lab for identification. Numbers of individuals at each bait site, time of day and microclimatic conditions are tabulated. Investigations are on-going on the predator-prey relationship of an ant and the anoline lizard which preys upon it. Also moss samples are collected in the same geographical regions. Tardigrades, nematodes and rotifers are being extracted and identified. New species are appearing and being described.

Impacts
(N/A)

Publications

• MILLER, W.R.,. MILLER, J. D. and.
• HEATWOLE, H. 1996. Tardigrades of the Australian Antarctic Territories: the Windmill Islands, East Antarctica. Zoological Journal of the Linnean Society 116: 175-184.
• HEATWOLE, H. 1996. Ant assemblages at t.

Progress 01/01/95 to 12/30/95

Outputs
Different groups of snakes were milked of venom at different schedules to ascertain how long it took to replenish the venom supply. On a different schedule, snakes were milked at monthly intervals to ascertain whether there were seasonal changes in either venom yield or venom composition. These experiments are still continuing and not analyzed statistically, but casual inspection reveals that there are marked seasonal changes in the amount of venom produced. Lateralization of the brain of birds and mammals occurs and resides in the neocortex, a part of the brain poorly developed in reptiles. As a test of such central lateralization, snakes were observed for behavioral tendency toward favoring one side or the other in coiling direction; they coil at random and therefore expressed no tendencies toward lateralization. This paper has been submitted for publication. Data previously collected on the sensitivity of a spectrum of prey species to cottonmouth venom was analyzed during the year and a paper on that topic is now in preparation. An expedition to the Ryukyus islands of Japan was carried out and large amounts of sea snake venom collected for use in studies in venom resistance by prey. It has now been lyophlized and is ready for use in experimentation. Papers arising from previous years on the project were published.

Impacts
(N/A)

Publications

• HEATWOLE, H. and COGGER, H. 1994. Sea snakes of Australia. Chapter 5 (pp. 167-205) in: Sea Snake Toxinology , Singapore University Press, Singapore.
• HEATWOLE, H. , CLOUDSLEY-THOMPSON, J. L. (1994). Predation and Defence, amongst Reptiles. Taunton, Somerset: R & A Publishing. ISBN 1-872688-03-9, viii + 138 pp. Frontispiece + 8 illustr. (Pbk) #12.75. Journal of Arid Environments 30.
• HEATWOLE, H. and PORAN, N. 1995. Resistances of sympatric and allopatric eels to sea snake venoms. Copeia 1995: 136-147.
• ZIMMERMAN, K. D., WATTERS, D. J., HAWKINS, C. J., and HEATWOLE, H. 1993. Effects of fractions isolated from venom of the Olive Sea Snake, Aipysurus laevis, on the behavior and ventilation of a prey species of fish. Journal of Natural Toxins 2: 175-186.

Progress 01/01/94 to 12/30/94

Outputs
Previous research on the distribution of sea snakes was written up and published. A new set of experiments involving the action of cottonmouth venom on target organs of bullfrogs was conducted. Techniques of organ culture, developed during the first year of the study, were improved and the effect of venom upon a variety of organs tested and described. An honors student and a PhD student also were trained in these techniques. A book was edited, about half of which dealt with the defensive secretions of amphibian skins and the effect these have on potential predators.

Impacts
(N/A)

Publications

Progress 01/01/93 to 12/30/93

Outputs
Experiments on resistance of eels to sea snake venom were completed, and that work has been extended by a series of experiments on the resistance of a New Caledonian eel (Gymnothorax) to the venom of one of its predators, the sea snake Laticauda colubrinus. This eel has the greatest resistance to sea snake venom of any fish known. Experiments on the resistance of bullfrogs to cottonmouth and copperhead venom were completed. Two aspects arising from that study were pursued. Tadpoles are susceptible, but during metamorphosis develop resistance. The new finding is that metamorphosed frogs lose their resistance again after they grow to a size too large for predation by the snakes. Thus there is an ontogenetic change in innate immunity corresponding to the risk of predation. The second aspect explored the role of sublethal doses of venom on rate of metamorphosis. Tadpoles subjected to low doses of venom resorbed their tails faster during metamorphosis than did controls. Since venom contains proteolytic enzymes, the hypothesis is that these may enhance the normal process of protein catabolism during tail resorption. Experiments quantifying this effect are underway. A technique for testing susceptibility of prey organs to snake venom was developed that allows assessment of relative susceptibility of different organs and identification of agents from serum that reduce the effect of venom. Lung tissue is highly resistant to venom, liver is susceptible and muscle is intermediate.

Impacts
(N/A)

Publications

• HEATWOLE, H. and COGGER, H. 1993. Sea snakes of Australia. pp. 167-197. In: Sea Snakes and Their Venoms. P. Gopalakrishnakoue, ed. National University of Singapore.
• HEATWOLE, H. and COGGER, H. 1993. Family Hydrophiidae, Sea Snakes. Ch. 36 in: Fauna of Australia Vol. 2. C. Glasby, ed. Australian Biological Resource Studies, Canberra. 17p.
• HEATWOLE, H. and GUINEA, M. 1993. Family Laticaudidae, Sea Snakes. Ch. 35 in: Fauna of Australia Vol. 2. C. Glasby, ed. Australian Biological Resource Studies, Canberra. 9p.
• HEATWOLE, H. and PIANKA, E. 1993. Ecology of the Squamates. Ch. 30 in: Fauna of Australia Vol. 2. C. Glasby, ed. Australian Biological Resource Studies, Canberra. 19p.
• DUELLMAN, W.E. and HEATWOLE, H. 1992. Habitat and adaptations. pp. 30-35. In: Reptiles and Amphibians. H. G. Cogger and R. G. Zweifel, eds. Weldon Owen Inc., San Francisco.
• ZIMMERMAN, K.D., HEATWOLE, H., and DAVIES, H.I. 1992. Survival times and resistance to sea snake (Aipysurus laevis) venom by five species of prey fish. Toxicon 30: 259-264.
• ZIMMERMAN, K.D. and HEATWOLE, H. 1992. Ventilation rates in three prey fish species treated with venom of the olive sea snake, Aipysurus laevis. Comp. Bioc.