Australia's Dangerous Snakes: Identification, Biology and Envenoming

Australia’s

Dangerous Snakes

IDENTIFICATION, BIOLOGY AND ENVENOMING

Peter Mirtschin, Arne R Rasmussen and Scott A Weinstein

© Peter J Mirtschin, Arne Redsted Rasmussen and Scott A Weinstein 2017

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National Library of Australia Cataloguing-in-Publication entry

Mirtschin, Peter, 1947– author.

Australia’s dangerous snakes : identification, biology and envenoming / Peter Mirtschin, Arne R Rasmussen and Scott A Weinstein.

9780643106734 (hardback)

9780643106741 (epdf)

9780643106758 (epub)

Includes bibliographical references and index.

Poisonous snakes – Australia – Identification.

Poisonous snakes – Venom – Australia.

Poisonous snakes – Ecology – Australia.

Rasmussen, Arne R., author.

Weinstein, Scott A., author.

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Front cover: The coastal taipan (Oxyuranus scutellatus) is one of Australia’s well-known medically important venomous snakes (photo: Peter Mirtschin).

Back cover (left to right): Lowland copperhead (Austrelaps superbus), extracting venom from a tiger snake (Notechis scutatus), peninsula brown snake (Pseudonaja inframacula) (photos: Peter Mirtschin).

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FOREWORD

Australia has a global reputation for harbouring the majority of the world’s most lethal animals: the ‘most deadly’ spider, octopus, fish (stonefish), etc. The venomous nature of the male platypus is an added toxic enigma. However, the myriad of highly venomous snakes is always the primary collective thought and cornerstone of much fascination, and fear, of Australia’s unique wildlife. Following the common media hype or watching any of the new wave of adventurous wilderness ‘warriors’, one cannot help but be bombarded by Australian serpentine ‘threats’ from land and sea. Too frequently and too simplistically, Australia’s venomous snake fauna is depicted as an unwanted element of our ecosystems that only introduces danger to human populations (especially tourists!). So many voices in the Australian community argue that snakes are to be avoided, eliminated or ignored. However, through studies like this book by Peter Mirtschin, Arne R. Rasmussen and Scott A. Weinstein, we see that snakes are also marvellous. They are biologically engineered masterpieces. Snakes have so many extraordinary adaptations, do so many remarkable things and influence ecosystems in such powerful ways that biologists of all persuasions are focussing on them with renewed enthusiasm. As we delve into the world of the snake, we cannot help but become awestruck by their extraordinary nature and their roles in our lives.

So how much do we really know about the snakes themselves? We are only beginning to unravel the natural history of our terrestrial and marine species and understand their biology, behaviour and evolution. However, understanding how snakes work is just the start of the journey towards truly understanding our most important group of predators. What role do snakes play in our ecosystems? Why do our snakes have such potent venoms? Are they indicator species whose abundance reflects the quality of habitat? Once we have mastered the ecology of snakes, we can then come to understand that they too face serious threats. Climate change, introduced predators, disease, landscape (and seascape) alterations and loss of prey species all can impact the survival of these predators. What are the consequences to ecological systems if snakes disappear?

And what of our relationship with snakes? Some species are actually commonly associated with urban, peri-urban and rural communities and contact is unavoidable. Do snakes play a role in our backyards and if so, is it useful to have them around? Why do some people love snakes, while others have phobias? If the worst does happen and someone is bitten, what then is the current best treatment (first aid and hospital), to ensure survival?

With the help of many researchers, experts, and photographers from a diversity of interests, this book unearths the secret lives of one of Australia’s most famous and iconic group of animals. The large and dangerous snakes of our terrestrial landscapes and tropical marine waters are not only described and identified in detail, but also with clarity and genuine affection. Peter Mirtschin, Arne R. Rasmussen and Scott A. Weinstein have devoted their lives to understanding snakes, snake venom and the medical management of snakebite envenoming. The book delivers the most recent information on the ecology of the dangerous snake species and the role they play, and the threats they face in natural and human-altered environments, as well as the latest in snakebite management. That snakes deliver such a vital role in balancing the delicate forces that shape ecosystems has only recently become clearer and leads to only one clear conclusion – the survival of snakes is unexpectedly and intimately interwoven with our own.

Chris Daniels

Professor of Biology University of South Australia Presiding Member, Adelaide and Mount Lofty Ranges, Natural Resources Management Board December 2015

CONTENTS

Foreword

Acknowledgements

1Introduction

2The relative danger of snakes

Snakebite and envenoming: a global perspective

Snakebite envenoming morbidity and mortality: public health problems of poverty and economic disadvantage

Risks from snakes: some further considerations

Venom lethal potency and risk: incorrect perceptions and valid concerns

3Morphology and general features of snakes

Basic properties and features of snakes

Front-fanged snakes, non-front-fanged (‘rear-fanged’) snakes

4Identification of dangerous snakes

Mistaken identity 1

Mistaken identity 2 (or not?)

Snake scalation

Key to identification of medically important elapid snakes in Australia

Key for terrestrial elapids identification

Key for sea snake identification

Key to Australian species of Aipysurus

Key to Australian species of Hydrophis

Key to Australian species of Laticauda

5The dangerous snakes described

Terrestrial snakes

Sea snakes and sea kraits

6Conservation of Australia’s dangerous snakes

Conservation terms

The threats on venomous snakes including medically important species

The road to recovery: solutions, viability and pragmatism

Biological control revisited: a delicate balance

Raising funds for habitat restoration and conservation

Laws and limits: statutory regulations and the purported ‘toll’ of collecting

Native animals as pets

7Production, actions and uses of venoms

The adaptive significance of snake venoms

Venom functions: an evolved armamentarium in the pursuit of prey

Venom production

Venom components

Procurement of snakes for venom collection

Effective venom production and captive husbandry

Venom extraction and safety

Antivenom

Snake venom for research: venom fractionation and characterisation

Some representative uses for venoms: from antivenoms to potential pharmacotherapeutics

Snake venom research: realities and the future

8Snakebite and envenoming: risks, realities and clinical management

Snakebite envenoming in Australia

Snakebite prevention

First-aid and first-response actions for snakebite envenoming

Management of envenoming in the hospital

Medico-legal issues/pitfalls

Discharge and follow-up

Long-term considerations/prognosis

Australia’s venomous snakes and their medical importance

Other medically important or potentially medically important species less frequently involved in human envenoming

Snakes of lesser medical importance

The future of snakebite management: possibilities and probabilities

9Snakes and people

Risks

Do we need to remove snakes?

Evaluation and impacts of relocation

How to avoid snakebites

Sea snakes: risks and realities

Benefits of having snakes around us

Appendix 1. The pioneers of venom production for Australian antivenoms

Early ‘snake men’

Twentieth century venom producers and snake handlers

Other snake handlers

Glossary

References

Index

ACKNOWLEDGEMENTS

Dr Bill Nimorakiotakis provided some valuable input during the early preparation and writing. Rosemary Thomson spent countless hours sourcing material on her father David Fleay. Her efforts helped attain an accurate portrait of her father’s lifelong contributions to the understanding of Australian venomous snakes. Meg Slater and Grayson Slater assisted with the procurement of material about their father, Ken, and also provided several photographs. John Weigel provided material about Eric Worrell, and also checked parts of the text. John Cann was most helpful with material and contacts, and gave permission to use material freely from his book Snakes Alive. Snake Experts & Antidote Sellers of Australia. He also provided some valuable historical images of the upper reaches of the Murray River. Dr Ken Winkel provided some valuable historical notes and letters from the Commonwealth Serum Laboratories (Seqirus). Albert Ramsamy checked the information about Ram Chandra. The late Jeanette Covacevich and Frank Madaras provided valuable information about Charles Tanner. Frank also provided diagrams and information about venoms and antivenoms. Neil Charles provided snippets and photos about David Fleay, and Paul Masci helped with some clotting diagrams. Neville Goddard and Roy MacKay assisted with photographs and information about Kevin Budden and Bill Hosmer. Dorothy Dale helped verify the accuracy of the story about her brother, Kevin Budden. Vince Meara helped add information about Bill Hosmer. Dr Bart Currie gave some assistance with the Ken Slater report, and provided some information for Chapter 5. Joan Crothers helped compile information about Frank Little. The authors thank Scott Eipper from Nature For You for his many contributions of photographs and assistance with some select edits of Chapter 5.

Special thanks to Prof Chris Daniels for providing some editorial assistance, and for encouraging Peter Mirtschin (PJM) to see the project through to completion, as well as for writing the Foreword.

Mark Hutchinson reviewed the terrestrial snake key and helped with selected diagrams, especially the computed tomography (CT) scans. Mark was also always willing to allocate time for PJM to examine and photograph South Australian Museum specimens. He also engaged in useful discussion with Scott Weinstein (SAW) regarding current hypotheses of squamate evolution.

During the early inception of the book, Ruchira Somaweera helped with some of the information about sea snakes, provided some useful editing of that material and contributed several fine images.

Kate Sanders for her generous help obtaining much needed images of several sea snake species and some technical advice about sea snakes.

Hal Cogger for inspiration and invaluable information; we also thank Nathan Dunstan, Luke Allen, Geoff Abraham and Brendan Schembri, who provided many opportunities for photographing specimens at Venom Supplies long after PJM retired, and especially Nathan who also on numerous occasions, unhesitatingly obtained information from Venom Supplies files.

David Williams provided useful information on several elapid species including some SVDK images of tests against D. vestigiata venom, and Bryan Fry helped with early drafts and provided advice to PJM.

Professor John Harris generously provided additional background regarding his important earlier research on the venom action on muscles and some outstanding images.

Professor Julian White provided some uncommon references, resources and accounts of personal experiences with several important snakebite cases as well as some of his invaluable archival snakebite photos and diagrams. Thanks to the Medical Library staff at the Women’s and Children’s Hospital, Adelaide, South Australia, for their time and efforts needed to procure several rare references for SAW. Dr David Bates provided SAW with help with figure preparation and their layout. We gratefully acknowledge the photographic contributions of Dr Judy Ou, as well as the Royal Flying Doctors, Central Operations.

Professor Rick Shine and his clerical assistant were always responsive to requests for papers. Rick also supplied information about himself for Chapter 6. Thanks also to: Brian Bush for information about P. atriceps and various Pseudonaja spp.; Harry Ehmann for information regarding P. atriceps; Tony Woods for information regarding the anti-tumour activities of several venoms. Mogens Andersen for allowing us to use his immaculate sea snake scale drawings; and Marucia Chacur for providing information about unpublished work regarding the analgesic effects of Naja kaouthia venom.

There were many photographers who were most generous in supplying valuable images without which the book would have been notably deficient. Although some have already been acknowledged, their contributions merit another mention along with others who unselfishly provided high quality photos: Mogens Andersen, Henry Cook, Scott Eipper, Brian Bush, Grant Husband, Brad Maryan, Lyall Naylor, Gary Stephenson, Brian Bush, Chris Malam, Kanishka Ukuwela, Ray Lloyd, Steve Swanson, Mahree-dee White, Lauren Vonnahme, Hal Cogger, Julian White, Rob Porter. Tony Courtney, Vinay Udyawer, John Read, Sally South, David Peacock, Jason Wishart, Glenn Edwards, Peter Tremain, Ray Chatto, Brent Johnson, Terri Shine, Alex Bloom, Aaron Fenner, Frozan Jamali, Tony Woods, Frank Madaras, Marucia Chacur, Emma Bateman, Brendan Schembri, Jeff Abraham, Judy Ou, Hamish Tristram, National Library of Australia, Steve Hoff, Roy Mackay, Neville Goddard, John Harris, Albert Ramsamy, John Cann, Neil Charles, Jane Melville, John Weigel, John Cann, Meg Slater and Grayson Slater, Eric Zillmann and the Royal Flying Doctors, Central Operations.

For their help and interest, we also thank: Professor Inn-Ho Tsai (Taiwan); Professor R Manjunatha Kini (Singapore); Byron Manning, Cleland Wildlife Park (assistance with dingo photo); Amanada Parker, Cleland Wildlife Park (assistance with python photos); Bradley Oliver (assistance with python photos).

The authors are grateful to Elsevier Press for allowing use of material substantially resourced from Mirtschin (2006) that is included in Appendix 1. It has been re-written and expanded here with the kind permission from Elsevier Press, as has some material published in Weinstein et al. (2013) that is included in Chapter 3.

PJM would like to thank his wife Mia for caring, nursing his brain and advising him through the many ups and downs of the journey preparing this manuscript, which would never have been possible without such patience and unselfish allotment of her time.

SAW also gratefully thanks his parents, the late Herbert Weinstein and Lenore Weinstein, who have tolerated numerous legless and multi-legged visitors for decades, and from his earliest years encouraged his consuming interests. For needed moral support, he also thanks: Dr Alison Dundon, Mark DiGirolamo, Dr Indraneil Das, Csaba and Monica Egri, Virginia Elder, Dr Khal Ismail, Dr Dan E Keyler, Brian Marian, Dr Pankaj Malik, Mark J Nerenberg, Gjorgjino Precaj, Dr Elizabeth Reynolds, Gerie Shearrow, Jerry Stachor, Jack Streitman, Dr Taksa Vasaruchapong, Stephen V Zivkovic and the late Norman Pachtman.

ARR would also like to thank Allan Greer, Mic Guinea, Ross Sadlier, Laura Sanders, Knud Botfeldt, Lasse Rasmussen, Rasmus Rasmussen and Annette Syberg.

1

Introduction

During the early days of the European settlement of Australia, snakes were abundant, and many snakebites occurred. The first antivenom for a medically important Australian elapid snake – tiger snake antivenom – was developed in 1930. Subsequently, in response to public pressure, other antivenoms were produced: taipan antivenom in 1955; brown snake antivenom in 1956; and death adder antivenom in 1958. The need for additional antivenoms was then recognised, and further products followed: Papuan black snake antivenom in 1959; sea snake antivenom in 1961; and the polyvalent in 1962 (Brogan 1990). The decline in numbers of snakebite deaths (Table 1.1) has been due to the introduction of antivenoms, ever improving knowledge of and use of better first aid, improved medical awareness and care (especially intensive care procedures such as mechanical ventilation), a public better educated about snakes, and also probably due to the decline in snake numbers, especially tiger snakes (Mirtschin et al. 1999). Snakebite has remained at about the same frequency over recent years, despite increases in the human population (Sutherland 1992; Sutherland and Tibballs 2001).

In 21st century Australia, many snake species have declined, and it appears as though the average size of individuals within many species has declined as well. In Australia, as in most countries, over the last century there have been innumerable changes to most habitats placing great pressures on many snake species. A few species may have benefited from these changes (e.g. because of increased availability of rodents attracted to human dwellings), but most have declined. The major environmental pressures that have most affected various snake populations include: direct habitat loss through land clearing; habitat modification caused by grazing domestic animals; coral bleaching and the concomitant loss of large populations of marine animals; introduction and presence of feral animals; loss of prey species; application of herbicides and pesticides; the impact of pest plant species; and probably global warming as well. The net effect of these pressures has caused a downward spiral in populations of most native animals that has not yet been halted, or even detectably slowed. Increasingly, global warming threatens to pose an evergrowing threat. If recent geophysical analyses prove correct, global warming threatens to further alter habitats in drastic ways, thus probably forcing the extinctions of many species.

Table 1.1. Decline in snakebite deaths in Australia.

Currently, medically important (‘dangerous’) snakes occupy all habitats of the whole of the mainland, Tasmania and many offshore islands, with sea snakes occurring in coastal waters around the northern half of Australia. In most habitats, there are more than one dangerous snake species. Even urban environments in our major cities have snake populations, and it is surprising to see how some species manage to fit into highly modified environments. Sea snakes occur in waters off two of our capital cities and off a number of the larger coastal provincial cities.

Interaction between humans, domestic animals and snakes inevitably occurs, and some people are gradually becoming more tolerant of co-existence with snakes, and even enjoying their presence. Australia manufactures some of the best antivenom in the world, and also has a medical fraternity who are quite well versed in snakebite management. There are now a number of recurring and regularly updated courses and symposia, which focus on the detailed topic of medical treatment of snakebite. This has contributed to the development of an emergency medical system and retrieval services that have improved capabilities to handle life-threatening envenoming from highly venomous Australian elapid snakes.

Our knowledge of some snake species has grown over the last 25 years, and cumulatively now represents a solid foundation for future management of some species threatened with possible extinction (e.g. Hoplocephalus spp., broad-headed snakes). In contrast, for many other species, vital knowledge about even their basic natural history is still lacking and represents a challenge to wildlife conservation agencies. For instance, studying the ecology of a species such as the inland taipan, Oxyuranus microlepidotus, presents real hurdles due to its isolation. Tiger snakes, once a very common species, are now in serious decline, and yet there is much to learn about their natural history. Their insular populations offer a tantalising opportunity to study a single snake species adapted to a diverse set of habitats. However, sometimes interest in expanding our knowledge of venomous snakes may arise from an unexpected source. With the growing demand for various seafoods, and the associated large harvests from the sea, some funding from seafood industries has been allocated to investigate the impact on sea snakes. Whether viewed as self-serving, or not, this has facilitated further study, and added to the basic knowledge about the ecology and ethology of several sea snake species. Nevertheless, the basic biology and natural history of most species of these enigmatic snakes has hardly been studied.

The conservation of Australian venomous snakes is of real concern, and in most cases it cannot be separated from general environmental losses and considerations. The challenge for future generations will be to wisely allocate funds for objectives that will yield the greatest benefits to most species. In this book, we have tried to generally describe the greatest threats to the irreplaceable populations of many of these snakes, and encourage efforts to protect their long-term survival. It is imperative that governments recognise the inherent value in protecting all of Australia’s unique wildlife – even the irrationally maligned venomous species. Government agencies must also ensure that their efforts are clearly focussed on major threat reductions.

In Chapter 7 and Appendix 1, we also outline the history of venom collection associated with antivenom development and consider venoms, and some representative examples of the modern applications of their components. It is important to appreciate that the journey of some of Australia’s pioneering amateurs and professionals who interwove their lives with snakes in order to learn about their behaviour, biology and venoms. Many of these early workers took great risks by working with some of the world’s most venomous species, and several paid the ultimate price with their lives. Although most of us have an inkling of the potent toxicity of some venoms, there is relatively limited recognition of the positive uses of these venoms. In cancer research, haematology, neurology, cardiovascular medicine and inflammatory diseases, venoms are used either for research, diagnostics or occasionally as therapeutic agents. A small handful of snake venom-derived components have been developed into important medications, and have been added to the pharmacotherapeutic armamentarium against human diseases.

When we are confronted by a snake, given a photograph of one, or find a dead one on the road, determining its identity will no doubt present a challenge for some of us. Many will use the coloured photographs in the book and combine them with the range maps of the snakes to determine the most likely identification. This can be sufficient, and this book contains many colour and pattern variations to compare with a specimen or photo. Some of the more studious will approach the identification more systematically, and use scale counts by following the diagrams provided. If this is insufficient, we have provided identification keys, which may help those willing to attempt more formal taxonomic identification. This can only be done if the snake is available, and it is safe to do so. For those not proficient in snake handling, counting scales on live snakes should not be attempted.

Treating snakebite is largely a specialised area of medical practice, and ideally best reserved for those skilled in this specialty (clinical toxinology). Fortunately, in Australia, the aforementioned highly specialised courses and workshops conducted by experts in this field have contributed to the distribution of a handful of knowledgeable people among most Australian states. Even so, it is best that seriously envenomed victims be transferred to a large emergency unit or intensive care ward as soon as possible. The first-response to snakebite is included in this book, as is discussion of the latest methods and criteria needed to effectively and compassionately handle an envenoming emergency.

Reptile houses are often the most visited exhibits in any zoo, and snakes are the primary attraction. This is not because they are ‘loved’, but often because people are drawn and fascinated by animals they interpret as ‘deadly’. For interested readers who don’t like snakes, we hope this book helps change that view to some extent. If this book manages to help some readers appreciate the value of having snakes in the environment, whether it be simply in the scrub or even in suburbia, then a major intention of this work will have been achieved.

2

The relative danger of snakes

Snakebite and envenoming: a global perspective

Just how ‘dangerous’ are snakes? In reality, only a relatively small number of potentially medically important snakes are ‘dangerous’ when harassed, inappropriately handled or accidentally contacted by an unsuspecting human. However, a number of medically important venomous species persist around agriculturally worked locations in several tropical regions, and thus unfortunately constitute a significant public health problem in several countries. In order to compare Australia’s potentially medically important snakes proportionally with those in other geographic regions, it is useful to briefly consider medically important snakebite from a worldwide perspective.

Global estimates/speculations of morbidity and mortality from snakebite envenoming consist of mostly crude approximations because of limited documentation. Aside from recent speculations of worldwide mortality between 100 000 and 200 000+, the true numbers of persons affected by permanent disabilities as a consequence of snakebite envenoming, remain largely unknown. Below are some estimates, and comments about their limitations, of the incidence of snakebite envenoming in several representative regions in which snakebite is a significant public health problem. Table 2.1 summarises some of the figures included in several published studies that have assessed snakebite incidence.

Africa and the Middle East

In a meta-analysis of documented reports, Chippaux (2011) estimated 60 snakebites/100 000 persons with an approximate average of 314 000 envenoming and 7300 deaths (1.5 deaths/100 000 persons), while Kasturiratne et al. (2008) reported a higher estimate of approximately 500 000 persons envenomed annually (depending on region, 3.33–93.34 snakebites/100 000 persons), with a high estimate of 32 195 fatalities (0.018–5.909 deaths/100 000 persons); the highest burden was in West Africa. Chippaux’s analysis also suggested that 95% of snakebites occurred in rural areas, as did 97% of fatal outcomes. The average envenomed victim was a male agricultural worker/farmer around 18–22 years old (Chippaux 2011).

As these figures are derived from published information and/or general estimates, it is clear that they are essentially ‘educated guesses’, and may over- or underestimate the actual incidence. Cautiously, with all factors considered, these approximations are more likely underestimates because of the likelihood of underreporting from rural locales, which constitute the main regions with substantial snakebite burden (de Silva et al. 2013). Also, it is likely that a significant number of envenomed victims do not present to medical facilities because of fear, superstition or local dependence on ‘traditional healers’.

Latin America and the Caribbean

The recent speculations about global incidence included regional assessments of snakebites and related mortality with subsequent high and low estimates. These speculative estimates pinpoint other regions where snakebite envenoming figures as a serious public health problem. For example, Kasturiratne et al. (2008) also reported high estimates of 135 123 snakebites for all of Latin America and the Caribbean, with 3459 deaths (depending on region, 0.008–1.05/100 000 persons). The highest snakebite incidence/100 000 people, was in the Andean region of Latin America, and Panama had a particularly notable annual snakebite incidence (79.8/100 000) (Kasturiratne et al. 2008). The highest snakebite mortality/100 000 people, was in the Caribbean region. As noted in the overview for other regions, these speculations are very likely gross underestimates.

Table 2.1. Estimates and speculations about the annual global incidence of snakebite, and comparison of geographic regions.

Asia

The estimates of snakebite in this very large geographic area containing marked venomous snake diversity also support the perception of snakebite envenoming as a serious, and underreported, public health problem. Comparison of two studies can provide a sense of the difficulties in assessing the true scope of this problem, the limitations that plague acquisition of accurate global snakebite data, and the judicious care that must be exercised when considering published estimates. Kasturiratne et al. (2008) performed a literature-based analysis, and thereby derived speculations about the regional incidence and mortality from snakebite envenoming. Their high estimates for Asia in total (including the Asian Pacific region) suggested >1.1 million annual snakebite envenoming (depending on region, 1.55–84.65/100 000 persons) with some 57 600 deaths (depending on specific region, 0.010–3.2 deaths/100 000 persons, Kasturiratne et al. 2008), and with the greatest burden noted in South Asia.

However, as observed earlier, these estimates must be viewed with great caution because a field-conducted survey analysing 123 000 deaths in randomly selected areas in India alone reported 562 deaths attributed to snakebites (Mohopatra et al. 2011). Ninety-seven per cent of deaths from snake-bite envenoming occurred in rural areas and were more common in males (59%) than females (41%), The incidence peaked at ages 15–29 years (25%) and during the monsoon months of June to September (Mohopatra et al. 2011). Therefore, this study suggested that there were some 45 900 annual snake-bite deaths in India alone with an annual age-standardised rate of 4.1/100 000 persons (Mohopatra et al. 2011), while the literature-based analysis of Kasturaratne et al. (2008) suggested a high estimate of approximately 57 600 deaths for all of Asia. Thus, it is likely, even with some expected error in the projected numbers, that available approximations are notable underestimates, and the actual mortality and morbidity from snakebites in the regions most affected by this problem remain largely and regrettably speculative.

Papua New Guinea (PNG)

Some recently collected figures from several provinces in PNG (2015 population approximately 7.7 million) have suggested an estimated 1519 snakebite envenomings, with 207 fatalities occurring annually in medical facilities, and another 780 deaths as a result of ‘non-care’ (Williams 2016), meaning a lack of modern medical treatment. A substantial number of tribal peoples of PNG are located distant from and/or avoid population centres such as Central Province (Dr Alison Dundon pers. comm.), and specifically its capital, Port Moresby, where most of the available, and thus limited, information has historically been collected. Therefore, many available figures (e.g. Table 2.1) are also likely to be significantly underestimated, and available information suggests that snakebite is a serious public health problem in many parts of PNG.

Snakebite envenoming morbidity and mortality: public health problems of poverty and economic disadvantage

In 2013, the World Health Organization (WHO) regrettably removed snakebite from its list of ‘Neglected Tropical Diseases’, then re-classified it as a ‘Neglected Condition,’ but then dropped it altogether in 2015. This can be interpreted as an apparent ‘lack of will’ to firmly tackle this serious problem, possibly because of the complicated nature and economic challenges of factors associated with the incidence, prevention, management and politics of many snakebites in economically disadvantaged countries, which have differences from those associated with other even larger public health challenges such as, for example, infectious diseases and malnutrition. However, it is encouraging that during the 2017 WHO 10th meeting of the Strategic and Technical Advisory Group for Neglected Tropical Diseases (NTD), snakebite envenoming was again added to the NTD portfolio.

Comparison of the approximated incidence and mortality from snakebites in envenoming ‘hotspots’ such as South-East Asia, the Indian subcontinent, Africa and regional Latin America with that in Europe, the United States, and Australia factually supports the perception that residing in an economically disadvantaged ‘Third World’ community carries a far greater risk of snakebite-related mortality and long-term morbidity. For example, in recent years, conservatively 2800–5500 persons have been reported to be envenomed by venomous snakes annually in the United States (population approximately 323.6 million) with an estimated 53% admission rate noted between 2001 and 2005, and a mortality rate less than 1% (Gold et al. 2004; Seifert et al. 2009; Langley et al. 2014), although some afflicted patients do suffer post-envenoming morbidity that may have lifelong consequences. Likewise, with the inclusion of East Thrace (‘European Turkey’) and West-Central Russia (‘European Russia’), an estimated 7500 viperid bites occur annually in Europe, which include some 1000 serious envenoming, but fewer than five fatalities (Chippaux 2011), a mortality rate less than 0.07%. This is especially noteworthy because the population encompassed within the whole of Europe amounts to some 739 million people.

Just a gross comparison of the very likely significantly underestimated broad range of snakebite mortality, >7000 to >30 000 deaths, annually among the population of all of Africa, approximately 1.21 billion people, with that of all of Europe, highlights the far greater risk of death from snakebite envenoming in rural areas of economically disadvantaged envenoming ‘hotspots’.

Risks from snakes: some further considerations

Which snakes do we consider dangerous? Clearly most large venomous snakes are an obvious consideration. Some people, however, develop allergies to all snakes and, in some instances, these allergies can be life threatening. Large pythons also present certain risks.

In certain circumstances, even snakes without a specialised venom delivery system can present a danger to humans. This especially applies to large pythons (e.g. Fig. 2.1). Provoked and unprovoked attacks by such species are occasionally recorded, sometimes causing significant damage or even death. For example, many keepers of pythons like to freely handle even very large specimens by hanging them around their necks or around the necks of other people in a show of the snake’s harmlessness. Most of the time these pythons tolerate this type of handling quite well and present little risk, but there have been instances when people have been overcome by them and had to be rescued by onlookers or, rarely, have even been killed. The risk here is asphyxiation, and not venom. Some boas and pythons have proteins in their oral secretions that cross-react with antivenoms raised against venoms of some venomous snakes (Minton and Weinstein 1987), and likely represent conserved products from shared common origins of squamate reptiles. However, these proteins are probably an example of ‘pre-adaptation’ (Gould and Vrba 1982), and do not constitute ‘venom’, nor do they have any medical significance (Weinstein et al. 2011). However, this phenomenon does have some relevance to the use of the Seqirus Snake Venom Detection Kit® (SVDK), which is sometimes used to confirm or support identification of the correct antivenom to use for a medically significant envenoming (see Chapter 8). This is because these proteins are present in venomous and non-venomous species such as pythons and thus a person bitten by a python may have a positive wound swab if tested with the SVDK (Jelinek et al. 2004).

However, this book is primarily focussed on medically important Australian members of the front-fanged snake family Elapidae. Elapids have a clearly defined high pressure venom gland and an effective venom delivery system, including fangs containing an enclosed canal, or lumen, that facilitate venom injection.

Venom lethal potency and risk: incorrect perceptions and valid concerns

Ranking venomous Australian snakes in order of danger to humans is a relatively academic exercise. This is because venom toxicity is most often determined with limited animal experiments, inevitably with small mammals (mainly mice). The most common means to compare experimental lethal potency is with the LD50 (lethal dose 50%). In order to calculate the LD50, different dosages of venom are administered to a group of mice in order to determine the lethal potency of the venom via a given route (e.g. intravenous, intramuscular, subcutaneous). The dose that causes a 50% death rate in a defined period of time (most commonly 24 hours) is calculated as the LD50. Despite its common utility, usefulness and effectiveness in understanding acute toxicity in a mammal model, the test has serious drawbacks with regard to animal ethics considerations. Also, because mouse physiology (and volume of distribution) has differences with human physiology and that of most domestic animals, the relativity between mice and humans or domestic animals is a rather tenuous comparison. In addition, rodents and other small mammals do not constitute the major type of natural prey animal for many species of snakes. Therefore, prey-specific toxins may affect one group of animals (e.g. birds or lizards) and have notably less effect on others such as small mammals. Thus, the experimentally derived lethal potency indices may not accurately reflect the lethal potency of some venoms for the natural prey of the snake whose venom is assayed with this method.

Fig. 2.1. A woma python, Aspidites ramsayi . Pythons do not possess fangs or a dedicated venom delivery system. Their oral secretions may contain proteins that exhibit immunological cross-reaction with several venom proteins from front-fanged elapid snakes, but there is no evidence that these function as toxins, and they have no apparent role in prey acquisition, nor do they have any medical significance.

The prey specificity of some venoms can be reflected by the effects observed in the bitten victim (humans versus other animals). Procoagulants (venom fractions that stimulate unregulated blood clotting) are the most prominent contributing cause of death and serious morbidity after human envenoming from brown snakes (Pseudonaja spp.). However, bites in domestic animals (mostly dogs and cats) from Pseudonaja spp. more often cause serious symptoms of neurotoxicity manifested by paralysis of the limbs and other parts of the body (see Chapter 8). Pseudonaja venoms have potent postsynaptic neurotoxins and extremely potent presynaptic neurotoxins but they are present in small quantities, although other neurotoxic species (e.g. tiger snakes, Notechis scutatus) have similarly proportioned neurotoxins in their venoms, and more commonly cause paralytic effects in humans (see Chapter 8). Thus, this marked difference in venom action in the envenomed subject may be related to proportional body size, innate susceptibility to a given toxin and/or other multi-factorial influences.

Bites from tiger snakes, many of the sea snakes and taipans can be quite different and the neurotoxins are far more important for both humans and animals possibly because of some of factors outlined above regarding the clinical effects of neurotoxins in the venoms of these species in comparison with those found in brown snake venoms.

Table 2.2. Factors affecting snakebite.

As alluded above, using mice as a sole indicator of relative danger via an experimental lethal potency assay (LD50) can be fraught with error and lead to false conclusions. Importantly, many antivenom producers solely rely on small animal testing to rate their antivenom potency, and this may yield clinically misleading conclusions about the efficiency of antivenom for a given venomous snake species.

The net pathophysiological effects from any snakebite will depend on many factors. These factors can be divided up into three categories: factors related to the snake, victim and management. Examples of some of the components involved with each factor are listed in Table 2.2.

Therefore, the health of a victim, as well as their tolerance for physiological insult, and immunity, are key factors that may influence the medical effects of snakebites. Bites inflicted on elderly, physically frail or very young victims are often more serious than those on heathy adults. Exposure to venom can sometimes cause allergic hypersensitivity (e.g. anaphylaxis) that can manifest in rash, wheezing, or a rapid, life-threatening fall in blood pressure and/or bronchoconstriction. This serious effect is more likely to occur in envenomed victims who have had a previous exposure to venoms, or other snake products. Exposure to powdered venoms is a particularly sensitive trigger of these reactions in some individuals, especially those who regularly associate with venoms (researchers, commercial antivenom producers, keepers of venomous snakes). Worldwide, many individuals who work with dangerous snakes and their venoms have developed hypersensitivity to the dried venoms. Tom (Pambo) Eades, the first person to collect venom in a systematic way for the Commonwealth Serum Laboratories (now Seqirus) in the 1930s, experienced many snakebites and frequently inhaled powdered venom, which led to the development of a serious hyperreactive airway condition and eventually caused his death in 1942 (Carter 1956; Cann 1986; Appendix 1). One of the authors of this book (PJM) has also developed an allergy to powdered snake venom, after a 20-year exposure while working with snake venom, and another author (SAW) experienced progressively expanding rashes when working with several specific rattlesnake venoms. Interestingly, these reactions were likely individual venom/species protein-specific, as they did not occur with other viper species or elapids. Subsequent to these reactions, special precautions had to be taken when working with venoms. Some precautions that are used under these circumstances are shown in Figs 2.2 and 2.3.

In general, bites from large snakes that normally yield more venom are more medically serious. The toxicity of the venom depends mostly on the snake species, but there can be intra-specific, geographical and ontogenetic (age-related) variation as well. Some species such as the taipan, Oxyuranus scutellatus, are well known to deliver repeated bites, thus potentially delivering greater amounts of venom.

Victims who receive correct first aid quickly and are transported to hospital, then given proper management are more likely to recover without serious complications. Most fatal outcomes after snakebites in Australia occur because of heart attacks or strokes that occur before the envenomed victim reaches the hospital.

Fig. 2.2. Nathan Dunstan from Venom Supplies (Tanunda, South Australia) handling venom. Note the respiratory mask and rubber gloves. The tiniest trace of venom can cause an allergic reaction to the handler and the end user of the venom. Ideally, this work should be performed with the operator prepared as shown, while the samples are contained in a laminar flow hood.

Fig. 2.3. Lyophilised venom, which is sealed under partial vacuum with nitrogen gas added. The nitrogen enables better storage of venom against degradation. Sealed vials ensure a safer way to handle and transport venoms protecting against aerosol (airborne venom particles) leakage.

Fig. 2.4. This West African Gaboon viper Bitis rhinoceros has hooked its fangs over the venom collecting vessel instead of inside of it. The huge fangs are inside the fleshy sheath – vagina dentis .

The biting mechanism of snakes can be a factor. Australian venomous snake fangs are typical elapid fangs and are relatively small compared with some snake species from other parts of the world, especially the viperid snakes (Fig. 2.4). Of the Australian snakes, taipans and death adders have relatively large fangs, tiger snake fangs are intermediate and brown snake fangs are small in relation to the bearer snake’s size. Fig. 7.7 compares fangs from three Australian species with that of the West African Gaboon viper (Bitis rhinoceros). Fig. 2.5 compares a range of non-Australian venomous snake fangs to a range of Australian venomous snake fangs.

In 1979, the Commonwealth Serum Laboratories published a summary of the lethal potency of most of Australia’s dangerous snakes (Broad et al. 1979a). Table 2.3 combines this data with several other sources to show relative toxicities of some of the tested species. These comparative figures are based on the acute toxicity in mice. However, comparing toxicities determined by different experiments is inherently complicated because of the many variables that could differ between laboratories, testing methods and operator interpretation. Therefore, this information is purely provided as a relative and superficial comparison. Table 2.4 compares the toxicity of a few snake venoms that, when assayed in mice, causes the death of all of the test animals (LD100) and also compares the venom toxicities for different animal species.

Fig. 2.5. Fangs of various medically important Australian terrestrial venomous snakes compared with a number of non-Australian venomous snake fangs. The fangs are: Non-Australian: B. r West African gaboon viper, Bitis rhinoceros ; C. a Eastern diamondback rattlesnake, Crotalus adamanteus ; B. a puff adder , Bitis arietans ; C. v Uracoan rattlesnake , Crotalus vegrandis ; B. n rhinoceros viper, Bitis nasicornis ; A. b cantil, Agkistrodon bilineatus ; B. s eyelash viper , Bothriechis schlegeli ; C. u Aruba Island rattlesnake, Crotalus unicolor ; O. h king cobra, Ophiphagus hannah ; N. k Thai cobra, Naja kaouthia. Australian: O. s coastal taipan, Oxyuranus scutellatus ; O. m inland taipan, Oxyuranus microlepidotus ; A. a common death adder, Acanthophis antarcticus ; P. g spotted black snake , Pseudechis guttatus ; P.c Collett’s snake, Pseudechis colletti ; P. a king brown snake, Pseudechis australis N. s tiger snake, Notechis scutatus ; A. s lowland copperhead, Austrelaps superbus ; T. c rough-scaled snake, Tropidechis carinatus ; H. s Stephen’s banded snake , Hoplocephalus stephensi ; P. t common brown snake, Pseudonaja textilis ; P. guttata speckled brown snake, Pseudonaja guttata.

Table 2.3. The relative toxicity of Australian terrestrial and other snakes.

Sources: Mouse toxicity figures are taken from Broad et al. (1979a) unless otherwise stated; yield figures were taken from Mirtschin et al. (2005) unless otherwise stated. The two different values categorised as ‘saline’ or ‘BSA’ (bovine serum albumin) represent two different diluents used to reconstitute the dried venom for injection. See pp. 96–97 for comments about the contentious subspecies of N. scutatus (not included in this table).

Abbreviations: s.c. subcutaneous, BSA bovine serum albumin, LD50 lethal dose 50%, intravenous, N/A not available

Given the dubious value of extrapolation of results and ethical considerations about invasive methods in experimental animals such as the LD50, in-vitro methods such as nerve muscle preparations, clotting studies or phospholipase A2 assays are becoming essential methods of venom bioassay. Although these methods provide important information about specific toxicities, they are usually restricted to detecting the effect of a single class of toxins. For example, the commonly used nerve muscle preparation – the chick biventer cervicis muscle assay – measures neurotoxicity only in avian (bird) tissue. In this method, biventer cervicis muscle and associated nerve fibres are dissected from the tissue surrounding the chicken’s cervical spine and subjected to electrical stimuli after being exposed to venoms or toxins (Ginsborg and Warriner 1960). This assay can be used to distinguish between neuromuscular blocking agents that cause depolarisation (essentially, cause a sudden change in the electrical charge of a cell), and those that do not. Both reduce the contractions caused by nerve stimulation, but depolarising drugs also cause a contracture of the muscle (Ginsborg and Warriner 1960). Table 2.5 depicts a comparison using this method between the action of venom from some sea snakes and the tiger snake (Chetty et al. 2004).

Table 2.4. Certainly lethal (LD 100 ) doses for various animals against various snake venoms.

Source: Kellaway (1929).

N/A not available

Table 2.5. A comparison of venom toxicity using the ‘isolated chick biventer cervicis nerve–muscle preparation assay’.

The time needed to observe a 90% reduction in twitches is measured. The in-vitro neuromuscular blockade of nerve transmission in this avian tissue sample is indicated by the shorter time needed to achieve reduction in the twitch response. This does not always reflect medically relevant neurotoxic potency (source: Chetty et al. 2004).

Blood clotting tests are another method commonly used to evaluate the pro- or anticoagulant activities of venoms (see Table 2.6). These tests can be relatively simple, and are carried out using the plasma from animal or human blood, or can be more complex assays using isolated clotting factors or artificial substrates.

All these methods of testing toxicity of venom have selectivity for certain groups of toxins. Therefore, the selection of a suitable method for testing venom actions would depend on the scope of biological activities desired in the study. More importantly, when testing the efficacy of antivenom, it is important to use a collection of tests rather than relying on a single method, because snake venom is a cocktail of many different components (often hundreds), and many cause significantly different effects, or are synergistic.

Most of the few studies (and most are limited sampling and/or anecdotal) that have considered this question have not shown any gender-related marked difference in experimental lethal potency of venoms. Also, no formal studies have been performed comparing gender-specific venoms from Australian snake species.

There also haven’t been any comparisons of venoms from snakes newly emerged from hibernation in comparison with samples taken later in the season.

Table 2.6. Clotting time in seconds for various terrestrial snakes using venom concentrations of 25 μg/mL in normal plasma in the presence of co-factors phospholipid and calcium chloride.

In this assay, the shorter clotting times indicate the stronger coagulant effect of the venom, because normal clotting in a control test tube (no venom added) occurs in approximately 60 seconds (source: Chester and Crawford 1982).

It’s often been said that juvenile snakes are more ‘deadly’ than adult snakes. It has been shown in studies with some non-Australian snakes, that venoms from juvenile snakes can be more experimentally toxic than venom from adult snake venoms (Gutierrez et al. 1980; Minton and Weinstein 1986; Lopez-Lozano et al. 2002; Saravia et al. 2002, 2015; Wray et al. 2015). Work carried out on three Australian species, Notechis scutatus, Oxyuranus scutellatus and Oxyuranus microlepidotus has shown that in these taxa there is little difference in toxicity and activity between adult and juvenile venoms in the activities tested (Tan et al. 1992, 1993a, 1993b). This may be partly due to similar prey in both juvenile and adult snake of these three species. Also, even in the species in which juvenile venoms have greater experimental toxicity, the venom yield is notably smaller than that of an adult. Therefore, receiving a bite from an adult would obviously be more medically concerning.

The seasonal variation of Australian snake venoms has been inadequately studied. One study of various American rattlesnake venoms, using isoelectric focussing, showed that no variation in patterns was evident over a 20 month period for any individual snake (Gregory-Dwyer et al. 1986). One Australian study examined the venom from a single brown snake, Pseudonja textilis, over a year and some changes were noted in some of the venom components, and the efficacy of brown snake antivenom for this venom (Williams and White 1992).

The relative danger from medically important venomous snakes is indeed relative to many factors because highly venomous snakes such as Pseudonaja spp. may refrain from injecting venom, while some species with venom less toxic to humans may on occasion inflict a life-threatening envenoming. The commonly held credo often holds true, that ‘the most dangerous venomous snake is the one that just bit you’.

Perhaps in the near future, the growing popular awareness of threatened and extirpated animal species might overpower the common human unwarranted fear of snakes including venomous species, and supplant it with an appreciation for their unique and vivid beauty. If this optimistic wish proves true, then future generations might be able to admire all snakes, not as potentially ‘dangerous’, but rather as a highlight of a walk through a natural setting. As the late P.H. William Bachmann closed his eloquent poem about snakes,

‘That day is pleasant, if some pathway leads,

To your bright beauty, flowing through the reeds.’ (Bachmann 10 January 1949; in Kauffeld 1969).

3

Morphology and general features ofsnakes

Snakes are secretive animals and many aspects of their natural history remain poorly understood, or even unknown. Their evolutionary history is fragmented and contains large gaps because of a poor fossil record. Thus, although there is so much to learn about snakes, an appreciation of their distinctive physical adaptations and physiology is essential in order to grasp their important features and functions. Such an understanding is also important to critically consider hypotheses about their evolution, and comprehending their diverse roles in the natural world.

Basic properties and features of snakes

Reptiles belong to the Class Reptilia, which contains all reptiles including turtles, tortoises, crocodilians, lizards, snakes and several other groups. The next level of classification hierarchy, the order Squamata, contains the reptiles that have similar physical characteristics, as well as other features such as ecological preferences and even behaviours.

Two separate suborders of the order Squamata exist: the lizards, sub-order Sauria (or, Lacertilia) and snakes, suborder Serpentes. In fact, the grouping of lizards and snakes in the Squamata appears to be a natural classification, meaning, that although it is ‘man-made’ grouping, these animals undoubtedly share their evolutionary history. Essentially, snakes are derived legless lizards. The distinction between the two suborders relies on the possession of a number of features and is summarised in Table 3.1, although in some species these distinctions can be quite subtle.

Although there are some notable exceptions to these ‘rules’ listed in Table 3.1, most lizard and snake species possess the features in their respective categories. The absence of limbs in snakes provides them stealth and access to a wider variety of microhabitats in search of prey while creating minimal sound or substrate disturbance. The possession of legs would hinder less detectable movement through grass, cracks, burrows and leaf litter. Although most lizards have legs, there are some that lack rear appendages (Fig. 3.1), anterior limbs or these extremities are substantially reduced. Sea snakes use their paddle-like tail to propel through the water, which also facilitates their egress into rock and coral crevices.

The serpent brain is completely enclosed in the skull, thus affording optimum protection during struggles, and protection against biting and clawing of prey. With a flexible jaw connection that distends the ligament connecting the upper and lower jaw, snakes are able to swallow prey much larger than their head size, offsetting the disadvantage of being unable to chew their food into small pieces. The snakes found in Australia have been assigned into six families (see Table 3.2).

The Elapidae (terrestrial snakes and sea snakes) contains all of the medically significant, or ‘dangerous’, Australian snakes.

Table 3.1. The major differences between snakes and lizards.

Front-fanged snakes, non-front-fanged (‘rear-fanged’) snakes

The World’s living snake fauna consists of approximately 3580 species, and the majority belongs to the superfamily Colubroidea, the ‘advanced snakes’ (e.g. those with derived anatomical traits), that evolved during the Oligocene-Miocene periods (approximately 34 to 23 million years ago) of the Cenozoic Era (Table 3.3). It should be noted that some authors use a more segregated, but complex phylogeny in which the advanced snakes are all included in the Infraorder Caenophidia with the families Elapidae and Lamprophiidae as part of the Superfamily Elapoidea. The recognition of up to 11 inclusive families in this phylogeny is dependent on a given author’s cladistics interpretation. Therefore, although the Colubroidea is used here for convenience, it should be noted that the dynamic nature of squamate reptile taxonomy might alter the usage/taxonomic accuracy of this particular phylogeny.

Fig. 3.1. Appearing superficially like a snake, this legless lizard ( Delma inornata ) has all the typically defining lizard features except visible legs.

Table 3.2. The families of snakes found in Australia.

Approximately 700 colubroid snake species (roughly, some 20%) are front-fanged (front-fanged colubroids, FFC), and possess either relatively ‘fixed’, or distensible front fangs in the maxilla that have variable morphology, but all have an enclosed lumen (canal) and orifice, which superficially resemble a hypodermic needle (see Fig. 7.8). This modified dentition is associated with a venom gland that commonly contains a stored bolus of venom that is ejected under pressure exerted by contraction of the skeletal muscle fibres inserted in the gland.

The ‘fixed’ fangs are found in members of the family Elapidae and a single genus, Homoroselaps (African dwarf garter or harlequin snakes, two species), of the otherwise non-front-fanged family Lamprophiidae, while the distensible fangs occur in members of the family Viperidae, and a single genus, Atractaspis (mole vipers, burrowing asps or stiletto snakes, arguably about 21 species), of the Lamprophiidae. In general, elapid fangs may be considered ‘fixed’ (meaning relatively immobile and in a permanently erected position in the maxilla), but there are a number of species (e.g. death adders, Acanthophis spp.) that have significant fang mobility. The term ‘fixed’ is primarily used in order to grossly distinguish elapid fangs, which are frequently called, ‘proteroglyphous’, from the generally more mobile fangs, often termed, ‘sole-noglyphous’, of viperids. The variability and embryological origins of ophidian dentition in the rear of the upper jaw, the posterior maxillae, renders the firm, broad categorisation of ‘fangs’ as somewhat artificial, and the terminology is largely used for general convenience.

For several centuries, many species of FFC have been formally recognised for their medical importance. The characteristics and risks of envenoming inflicted by numerous clinically important taxa have been documented with increasing medical accuracy since the late 19th century. Of the living FFC belonging to the families Viperidae and Elapidae, probably about 50% (some 300–350 species) are known to inflict medically significant bites on humans. The Viperidae consists of the Old World vipers, subfamily Viperinae, and the New and Old World pit vipers, subfamily Crotalinae, while the Elapidae consists of Australian species such as tiger snakes, brown snakes, death adders, taipans and black snakes, as well as other well-known groups from other geographic regions such as cobras, mambas, coral snakes, sea snakes and their allies.

In reality, a relatively small proportion of FFC species are responsible for most of the global annual snakebite-related human mortality and morbidity, which has been roughly approximated to include 45 000–100 000 deaths (see Chapter 2). Several examples of these most medically important FFC species include: Echis spp. (the saw-scaled, or carpet, vipers, Viperidae, Viperinae, ~11 species, Africa, Middle East, Asia), Naja naja and N. kaouthia (respectively, the common Asian cobra and the monocled, or monocellate cobra, Elapidae, Asia), Oxyuranus scutellatus canni (Papuan taipan, Elapidae, Papua New Guinea), Bitis arietans (the puff adder, Viperidae, Viperinae, sub-Saharan Africa), Russell’s viper (Daboia russelii, D. siamensis, Viperidae, Viperinae, Asia), Bungarus spp. (kraits, Elapidae, approximately 14 species, Asia), Bothrops asper and B. atrox (respectively, the terciopelo, and barba amarilla, or common lancehead, Viperidae, Crotalinae, Latin America; both are occasionally called ‘fer-de-lance’, a term that might be more accurately used for the Martinique lancehead, Bothrops lanceolatus). These species collectively account for a large proportion of the global human mortality and/or morbidity from snakebite. Their prominent medical importance is a result of: their abundance and persistence in the vicinity of human agrarian (e.g. farming) populations; the marked toxicity of their venoms for humans; the unfortunate higher frequency of poorer health of many of the people subjected to this greater risk of serious snakebite in the relevant geographic regions (e.g. poor nutrition and high parasite burdens in themselves can contribute to a poor outcome); the likewise regrettable limited availability of rapid, medically supported retrieval, as well as an affordable, adequate and safe supply of effective, regionally specific antivenom, and additional factors associated with the snakes and the co-habiting human populations (see Chapter 8).

In addition to the FFC species that are known to have prominent medical importance, there are also a number of lesser known front-fanged species (including a number of small fossorial Australian elapids and the aforementioned Homoroselaps spp.) of uncertain medical