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Copyrighted Material Some pages are omitted from this book preview. Giant Giant Snakes A Natural History John C. Murphy & Tom Crutchfield

Snakes, particularly venomous snakes and exceptionally large constricting snakes, have haunted the brain for a millennium. They appear to be responsible for our excellent vision, as well as the John C. Murphy & Tom Crutchfield anxiety we feel. Despite the dangers we faced in prehistory, snakes now hold clues to solving some of humankind’s most debilitating diseases. Pythons and boas are capable of eating prey that is equal to more than their body weight, and their for this are providing insight into diabetes. Fascination with snakes has also drawn many to keep them as , including the largest . Their popularity in the trade has led to these large constrictors inhabiting southern . This book explores what we know about the largest snakes, how they are kept in captivity, and how they have managed to traverse ocean barriers with our help. Copyrighted Material Some pages are omitted from this book preview. Copyrighted Material Some pages are omitted from this book preview.

Giant Snakes A Natural History

John C. Murphy & Tom Crutchfield Copyrighted Material Some pages are omitted from this book preview. Giant Snakes

Copyright © 2019 by John C. Murphy & Tom Cructhfield All rights reserved. No part of this book may be reproduced in any form or by any electronic or mechanical means including information storage and retrieval systems, without permission in writing from the publisher. Printed in the of America First Printing March 2019

ISBN 978-1-64516-232-2 Paperback ISBN 978-1-64516-233-9 Hardcover

Published by: Book Services www.BookServices.us

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This book is dedicated to the survival of Giant Snakes around the world.

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Front Cover: Nathalie Aall Back Cover: Reticulated (Michael J. Jowers), Yellow (JCM), (JCM), (Bill Lamar) Inside Front Cover: Reticulated Python (JCM) Inside Back Cover: Reticulated Python, Morph (JCM) Inside Dedication page: Ceylonese Python, pimbrura. JCM

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Acknowledgments

The authors offer their sincerest thanks to the following individuals for their photographs, stories, and helpful comments. For comments on the manuscript and personal stories we thank: Steve Barten, Harry Green, Tom Lorenz, Paul Moler, Gordon Schuett, Russ Gurley, Ron St. Pierre, Nick Leszai, Ken McCloud, Sara Murphy, Joe Sambono, Vic Morgan, Janet Weems, and Andrew Wyatt. Heinz Wilhelm helped us with translations; Ricky Duffield provided info on the warehouse of pythons that was supposedly blown away. Monique Crutchfield and Penny Crutchfield helped TC (Tom Crutchfield) in recalling events. The authors greatly appreciate the use of photos from the following: Gavin Bedford, Myke Clarkson, Maheta Jaydeep, Arno Naude, Bill Lamar- Green Tracks, Bill and Kathy Love, Bill Stiffler, Breck Bartholomew, Chris Gillette, Clarence Aber- crombie, Cor Viljoen, Joe Fauci, Joe Sambono, Joe Wasilewski, Jose Luis Ulgade Trejo, Kevin McCurley- New England , Max Jackson, Michael Powell, Ron Ramsey, Mike Rochford, Ray Van Nostrand Sr., Rhett Stranberry, Renato Yabiku - Huachipa Zoo, Michelle Sutton, Sebastian Holch, Tal Feinberg and Tom Lorenz -Georgia Southwestern State University. The cover art was done by Nathalie Aall at aallformsoflife.com. The images that start each chapter are as follows: “Anxiety, Vision and Snakes” is a painting titled “The Constrictor” (1867) by Aloys Zötl, an Austrian artist known for his paintings of natural history subjects. “Size and Shape” is a North African Python (JCM). “ Origins & Biology” illustrates a pelvic girdle in a cleared and stained specimen (JCM.) “Pythonids, an Overview” is a juvenile (JCM). “Giant Constrictors of Australasia” is a large Scrub Python (Joe Sambono). “Giant Pythons of the Afro-Asian ” is a North African Python (Michelle Sutton). “The Reticulated Python Clade” is a Reticu- lated Python (JCM). “Booid Snakes, an Overview” is a Ruschenberger’s Treeboa (JCM). “The Anaconda Clade, Giant Aquatic Boas” illustrates a Green Anaconda (Bill Lamar). The Clade is a Boa imperator (JCM). “Giant Snakes in Captivity” shows two exceptionally large Burmese Pythons, one a color morph, the other a wild morph (Jose Luis Ulgade Trejo). “Giant Snakes in Florida” is a Burmese Python in a cypress swamp in southern Florida (Chris Gillette). “Extinct Giant Snakes” is a highly modified Green Anaconda (Bill Lamar).

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Preface

Since the publication of Tales of Giant Snakes (Murphy and Henderson, 1997), much of what we know about the largest snakes in the world has changed. The record has revealed remains of a truly colossal snake, , that was living in northeastern 58 to 60 Ma. Evidence and hypotheses about how snakes evolved their giant size have become more detailed, and ecological field studies of giant snakes have become much more common. Giant and near-giant snakes are more popular than ever in the pet trade; these “designer snakes” are bred in captivity for their unique colors and patterns. The novelty skin industry is consuming giant snakes in higher numbers than ever before. Giant snakes have become invasive in southern Florida. Of lesser importance, the record Murphy and Henderson (1997) considered as representing the largest known extant snake has proved to be false. This has happened to many previous records believed to be valid. Extant giant snakes do not get as long or as massive as most people think based on the liberal acceptance of old and unreliable records that have been published and republished for many years. Murphy and Henderson (1997) defined giant snakes as those species that exceed 20 feet (6.1 m). In this book, we will continue with that definition for giant species and describe the near-giant species as those that approach 6.1 m, but may not reach it. At this point, the extant giant snakes appear to be the Green Anaconda ( murinus) of , the Reticulated Python ( reticulatus) and Burmese Python (Python bivittatus) of Southeast , the South African Python (Python natalensis), North African Python (Python sebae) from sub-Saharan , and Kinghorn’s Scrub Python ( kinghorni) of northern . Near-giants are the Indian Python (Python molurus), and the Western ( olivaceus barroni) from . Other species that obtain significant size are discussed: the Oenpelli Python Simalia( oenpelliensis) from Australia, the Papuan Python ( papuana) from New , and the Cuban Boa, ( angulifer). A discussion of the Boa Constrictor (Boa constrictor) is included, not because it’s a giant snake—but because so many people think it is! On March 6, 2015, the U.S. Fish and Wildlife Service (USFW) finalized the Constrictor Rule, a controversial policy designed to reduce the probability that more invasive constricting snakes will find their way into ecosystems in the U.S. The Constrictor Rule declared the Reticulated Python, DeSchauensee’s Anaconda, Green Anaconda and Beni Anaconda as “injurious” under the Lacey Act. A fifth species, the Boa Constrictor, was removed from consideration for listing as an inju- rious wildlife species after an outcry from keepers of pet snakes and their lobbying

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organization, the United States Association of Reptile Keepers (USARK), applied political pressure for the Boa Constrictor removal from the listing. The rule sparked considerable controversy because these species are in the pet trade and the designer snake industry, and some are already established as in southern Florida. This book is not intended to be a platform for political activism. It is about science, natural history, and how and exceptionally large serpents relate to each other. Both authors have kept giant and near-giant snakes in captivity for decades, and we can sympathize with those who want to maintain and breed giant snakes in captivity. They are amazing , and there is much to be learned from their lives in captivity as well as in the wild. Keeping them in captivity is challenging because of their size and food requirements, and we discuss this at length in one of the chapters. However, in a foreign landscape, they are capable of devastating the local fauna. The argument for keeping giant snakes in captivity can be reduced to individual rights versus the common environmental good. This book is not a reworked Tales of Giant Snakes. It is an updated look at the scientific and popular literature, as well as personal experiences and those of people the authors know. It is intended to provide a context for giant snakes and their many relationships with humans. If you are a reader unfamiliar with the term clade, it refers to an ancestor and all of its descendants. by definition are monophyletic; that is, they have one ancestor. Thus, a group of species that shared a common ancestor forms a clade. In traditional classification, a is a clade. In traditional classification, all the members of a , in theory, should share a single ancestor, making it also a clade. Although this book was written for the general public, we have included a section targeted to people who keep giant snakes as pets but may not be as familiar with them as we are. Generating an appreciation of giant snakes, their , and the science that builds knowledge of the natural world are also on our list of goals for this book. Common English and scientific names are provided. The scientific name is followed by the name of the person(s) who originally described the species. If the person’s name is in parentheses, it means the species is now placed in a different genus than it was when it was first described. The maps were made using ArcView, with localities obtained from VertNet and the literature. Climatic map data were obtained from GLDAS (Global Land Data Assimilation System).

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Abbreviations and Acronyms

m: meters JCM: John C. Murphy kg: kilograms Ma: Million years ago NRM: Naturhistoriska riksmuseet (Swedish Museum of Natural History) SDNHM: San Diego Natural History Museum SSD: Sexual Size Dimorphism SVL: Snout to vent length TC: Tom Crutchfield TL: Total length UINHS: University of Illinois Natural History Survey YBP: Years before present YPM: Yale Peabody Museum Note: We use metric units for all measurements in this book. For the metric- challenged we have added a conversion table.

Metric conversions made simple. Convert Into Multiply by centimeters inches 0.3937 feet meters 0.3048 hectares acres 2.471 inches centimeters 2.54 kilograms pounds 2.205 kilometers miles 0.6214 meters feet 3.281 miles kilometers 1.609 pounds kilograms 0.4536 Fahrenheit Celsius subtract 32 0.5556 Celsius Fahrenheit multiply by 1.8 Add 32

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Fronticepiece. The struggle between man and snake is illustrated in this plate from ’s Malayan Archipelago. The snake involved is a Reticulated Python, which the man is trying to extract from the house. His posture suggests both his anxiety and determination in removing the snake. Anxiety over snakes is likely based upon hominins’ evolutionary history of living alongside venomous snakes, giant constrictors, and hypercarnivores. The plate is titled “Ejecting an Intruder.” The house was used by Wallace while he was in Amboyna collecting insects. Wallace observed the snake in the thatched roof while he was reading on the couch. Realizing it was a python and that it had been in the roof for at least a day, he called two of his workers and asked them to remove the snake; however, they refused and ran out of the house. He called in more men from a nearby plantation, but only one was willing to actually attempt to remove the snake. He made a noose of rattan, mounted it on a pole, secured the snake in the noose, and pulled it out of the ceiling. There was a long struggle to get the snake outside, where it was killed. Wallace noted the that snake was about four meters long, large enough to swallow a child.

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xi Copyrighted Material Some pages are omitted from this book preview. Giant Snakes Contents Acknowledgments ...... v Preface ...... vii Abbreviations and Acronyms ...... ix Chapter 1. Anxiety, Vision, and Snakes ...... 1 Chapter 2. Size and Shape ...... 9 Sexual Size Dimorphism ...... 12 Becoming A Giant Species ...... 13 Allometry, Heterochrony, and Growth ...... 15 Ontogeny and Snakes ...... 17 Pritchard’s Rule ...... 18 Acceptable Evidence for Giant Size ...... 20 A Critique of the Three Approaches ...... 21 The Giants and Near-giants ...... 24 Skin Length vs. Total Length ...... 24 Popular and Often-Repeated Stories That Are Misleading ...... 26 Our Perspectives on Maximum Size ...... 31 Scrub Python, Simalia kinghorni ...... 32 Burmese Python, Python bivittatus ...... 32 Indian Python, Python molurus molurus ...... 34 Ceylonese Python, Python m. pimbura ...... 34 South African Python, Python natalensis ...... 34 North , Python sebae ...... 35 Reticulated Python, Malayopython reticulatus ...... 36 Green Anaconda, Eunectes murinus ...... 37 Chapter 3. Snake Origins & Biology ...... 39 Body and Tail Lengths ...... 46 Temperature Control ...... 48 Chemosensory Systems and Thermal Imaging ...... 49 Active Foraging and Ambush ...... 51 Food and Feeding ...... 51 Ecdysis ...... 55 Diel Color Change ...... 56 Modes of Reproduction ...... 57 Sex Determination & Facultative ...... 58 Multiple Paternity and Mating Systems ...... 63 Relative Clutch Mass (RCM) ...... 64 Geography, Climate, and Snake Body Size ...... 65 Some Ecology ...... 66 The Genome ...... 70 xii Copyrighted Material Some pages are omitted from this book preview. Contents

Chapter 4. Pythonids – An Overview ...... 73 The Afro-Asian Python Clade, Genus Python ...... 77 The Indo-Malaya Python Clade, Genus Malayopython ...... 79 The Carpet & Tree Python Clade, Genus Morelia ...... 80 The Dwarf Python Clade, Genus ...... 82 The Clade, Liasis + Apodora ...... 84 The Scrub Python Clade, Genus Simalia ...... 84 The Black-headed Python Clade, ...... 86 The Ringed Python Clade, Bothrochilus + ...... 87 Chapter 5. Giant Constrictors of Australasia ...... 89 Scrub Python, Simalia kinghorni (Stull, 1933) ...... 91 Olive Python, L. Smith, 1981 ...... 96 Oenpelli Python, Simalia oenpelliensis Gow, 1997 ...... 100 Papuan Olive Python, Apodora papuana (Peters and Doria, 1878) . . . . 103 Chapter 6. Giant Pythons in the Afro-Asian Clade ...... 105 TheMolurus Clade ...... 108 Burmese Python, Python bivittatus bivittatus Kuhl 1820 ...... 111 Python, Python bivittatus hainannus (Zang 2015) ...... 122 Python, Python bivittatus progschai Jacob, Auliya & Böhme 2009 123 Indian Python, Python molurus Linnaeus, 1758 ...... 123 Ceylonese Python, Python molurus pimbura Deraniyagala, 1945 . . . . 137 African Members of the Molurus Clade ...... 138 South African Python, Python natalensis Smith 1840 ...... 142 North African Python, Python sebae Gmelin, 1788 ...... 144 Chapter 7. The Reticulated Python Clade ...... 149 The Jampea Reticulated Python, Malayopython reticulatus jampeanus (Auliya, Mausfeld, Schmitz, and Böhme 2002) ...... 152 Selayar Reticulated Python, Malayopython reticulatus saputrai (Auliya, Mausfeld, Schmitz and Böhme 2002) ...... 153 Reticulated Python, Malayopython reticulatus reticulatus (Schneider 1801) . 153 Chapter 8. Booid Snakes, an Overview ...... 169 West African Burrowing Boa – Calabariidae ...... 172 The Dwarf Boa Clade of the New World – Charinidae ...... 174 The Sand Boa Clade – Family Erycidae ...... 175 The acificP Boa Clade - Candoiidae ...... 177 The eotropicalN Boa Clade - ...... 177 The Boa Clade ...... 177 The reeboaT Clade, ...... 178 The Anaconda Clade, Eunectes ...... 178 The Rainbow Boa Clade, ...... 180 The slandI Boa Clade, Chilabothrus ...... 180 The alagasyM Boa Clade, Sanziniidae ...... 181 xiii Copyrighted Material Some pages are omitted from this book preview. Giant Snakes

Chapter 9. The Boa Constrictor Clade ...... 185 Short-Tailed Boa, Boa constrictor amarali (Stull, 1932) ...... 188 Red-Tailed Boa Constrictor, Boa constrictor constrictor Linnaeus 1758 . . 190 Long-Tailed Boa Constrictor, Boa constrictor longicauda Price and Russo 1991 ...... 191 Argentine Boa Constrictor, Boa constrictor occidentalis Ihering 1911 . . 191 Orton’s Boa Constrictor, Boa constrictor ortonii Cope 1878 ...... 192 Central American Boa Constrictor, Boa imperator imperator Daudin 1803 193 Pearl Island Boa, Boa imperator sabogage (Barbour 1906) ...... 194 Dominican Boa Constrictor, Boa nebulosa (Lazell 1964) ...... 194 Tet’chien or St. Lucian Boa, Boa orophias Linnaeus 1758 ...... 195 Mexican West Coast Boa Constrictor, Boa sigma (Smith 1943) ...... 195 Chapter 10. The Anaconda Clade, Giant Aquatic Boas ...... 197 Beni Anaconda, Dirksen, 2002 ...... 200 DeSchauensee’s Anaconda, Dunn and Conant, 1936 . 200 The reenG Anaconda, Eunectes murinus (Linnaeus, 1758) ...... 201 , Eunectes notaeus Cope 1863 ...... 211 Chapter 11. Giant Snakes in Captivity ...... 215 A Brief History of Herpetoculture ...... 219 Giant Snake Color Morphs ...... 219 Monsters of God as Pets ...... 224 Housing and Maintaining Giant Snakes ...... 231 Medical Care for Giant Snakes ...... 235 Giant Snakes as Educational Ambassadors ...... 236 Infectious Diseases in Snakes ...... 238 Breeders Became Exporters ...... 241 Chapter 12. Invasive Giant Snakes in Florida ...... 243 The efariousN Alteration of the South Florida Landscape ...... 247 Imports ...... 252 How did pythons become established in Florida? ...... 252 Distribution of Burmese Pythons in Florida ...... 258 Estimating Population Size ...... 258 The ietD of Florida Pythons ...... 259 Python Ecology in Florida ...... 261 They Adapt ...... 263 Reproduction in Florida ...... 265 The Impact on Native Wildlife and Disease ...... 267 Controlling Populations ...... 269 The iscoveryD of North African Pythons in Miami ...... 271 The ybridH Python Issue ...... 272 The Boa Constrictor in Florida ...... 274 Other Points of Possible Dispersal ...... 275 xiv Copyrighted Material Some pages are omitted from this book preview. Contents

Chapter 13. Extinct Giant Snakes ...... 277 Family Boidae† ...... 280 Chubut Boa, Chubutophis grandis† Albino ...... 280 Talisma Anaconda, cf Eunectes sp.† ...... 280 Titanic Boa, Titanoboa cerrejonensis† Head, Bloch, Hastings, Bourque, Cadena, Herrera, Polly, and Jaramillo ...... 280 The Family Madtsoiidae ...... 280 Garstin’s Giant Snake, garstini† Andrews ...... 281 Patagonian Giant Snake, bai† Simpson ...... 281 Malagasy Giant Snake, Madtsoia madagascariensis† Hoffstetter . . . . 282 Naracoorte Giant Snake, naracoortensis† Smith ...... 282 Camfield Giant Snake, camfieldensis† Scanlon ...... 283 Dinosaur-eating Snake, Sanajeh indicus† Wilson et al...... 283 The Family Paleophidae ...... 284 Eocene Snake, colossaeus† Rage ...... 285 Shark River Eocene Snake, Palaeophis grandis† (Marsh) ...... 285 Potomac Eocene Snake, Palaeophis virginianus† Lynn ...... 285 Choctaw Giant Aquatic Snake, schucherti† Lucas . . . . . 285 Family ...... 286 Bluff Downs Pliocene Python, Liasis dubudingala† Scanlon and Mackness, 2001 ...... 286 Riversleigh Python, Morelia riversleighensis† (Smith and Plane) . . . . . 287 Appendix ...... 288 Appendix 1. Some DNA Basics ...... 288 Appendix 2. How the Anaconda Got its Name ...... 289 The Second Story: How the Green Anaconda got its Scientific Name. . . . . 293 Appendix 3. Giant Snakes in Oral Traditions ...... 294 The Ancestral Written Snake Story ...... 296 Names and Ancestry ...... 302 Appendix 4. Attacks and Deaths from Constrictors ...... 303 Glossary ...... 307 References ...... 312 Index ...... 340 Author Information ...... 346

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Chapter One Anxiety, Vision, and Snakes

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On June 15, 2018, fifty-four-year-old Wa Tiba left her small village, Persiapan Lawela, in Sulawesi, to tend to her garden located about half a mile from her house. Tiba’s family reported that she wanted to check on the garden because wild boars often destroyed her crops. When Tiba didn’t return home by sunrise, her sister went to look for her near the garden. She could only find some of her sister’s belongings, including a flashlight, her machete, and her sandals. About 100 local people, including Tiba’s relatives, launched a search party. They encountered a 23-foot Reticulated Python with a bloated midsection about 30 meters away from where Tiba’s belongings were found. Suspicious that the snake had swallowed the woman, they killed it and carried it out of the garden. Back at the village, the villagers sliced open the snake and found the body of Wa Tiba, who had been swallowed head first. The woman was just a different kind of prey. Humans have always been on the menu of giant snakes. You have experienced it; everyone has at some time. You may not understand what it is, but you have sensed the uneasy feeling and the worry that comes with being human. It does not seem to have a specific trigger. It is a feeling that something is wrong, danger is near. For some, the anxiety can be debilitating. It takes many forms, and often it goes unacknowledged by the conscious mind. Panic, phobias, tense muscles, sleep problems, self-consciousness, and compulsive behaviors are all symptoms. Anxiety is a physiological response set in the subcortical and deep cortical structures of the brain. Fear and anxiety are not exclusively associated with , although preda- tion is the most ancient selection pressure shaping the fear response system. Fear not only organizes escape and avoidance responses to dangers, but also activates defen- sive responses and deactivates some cognitive processes, such as searching for a mate. The source of anxiety lies in our inheritance: we’ve spent millions of years trying to escape predation. The fear response is more influenced by the ancient species that struggled to survive than any present-day challenges. As Ann Druyan and Carl Sagan (1995) succinctly put it in the title of their book, we live in a Demon-Haunted World. Today, those demons are mostly extinct, but the fear response survives within us, and sometimes it is aroused at inappropriate moments. Appropriate moments would be times when we are trying to avert a car crash, avoid a falling object, or escape a violent person. Our hominin ancestors spent millions of years exposed and relatively defenseless. For many of those years, humans faced hypercarnivores. In the Eastern Hemisphere, 3 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes cave bears, cave lions, cave hyenas, and even the extant African lion, hunted our ancestors. In the Western Hemisphere, wolves (though probably not the gray wolf of today), saber-toothed , and the dire wolf may have hunted humans across the landscape. Hypercarnivores were sometimes solitary hunters, but on occasion, they hunted in social groups (Van Valkenburgh et al. 2016). Some hypercarnivores even took the form of giant and , including , giant , and, of course, giant snakes. Writer and biologist, David Quammen (2003) was quite blunt about the situation in the opening of his book, Monster of God. Great and terrible flesh-eating beasts have always shared the landscape with humans. They were part of the ecological matrix in which Homo sapiens evolved. They were part of the psychological context in which our of identity as a species arose. They were part of the spiritual systems that we invented for coping. The teeth of big predators, their claws, their ferocity and their hunger, were grim realities that could be eluded but not forgotten. Every once in a while, a monstrous carnivore emerged like doom from a forest or a river to kill someone and feed on their body. It was a familiar sort of disaster—like auto-fatalities today—that must have seemed freshly, shockingly gruesome each time, despite the familiarity. And it conveyed a certain message. Among the earliest forms of human self-awareness was the awareness of being meat. As technology evolved, humans did their best to eradicate large predators to make the world safe for humankind. We speared, poisoned, trapped, and shot as many large predators as we could. Today, few survive. Relatively few individuals of the few species of big cats are still around: the , the , the tiger, the African and Asian lions, the American mountain lion. There are few recorded deaths from despite their living in close prox- imity to humans. African lions kill about 250 people each year. kill about 85 people per year in and probably take a few more humans over the remainder of their distribution. In the United States, there were two deaths from mountain lions between 1986 and 1998 and a few more recent ones. This large is expanding its range and is expected to be hunting along the eastern seaboard very soon, if it is not doing so already. undoubtedly kill a few people each year. However, the total number of humans taken by big cats is minute compared to the deaths caused by motor vehicles or firearms. Bears will kill and eat humans; however, more humans are killed or mauled because the bear is defending its young, a kill, or its territory than are killed from predation. Bears are not hypercarnivores in the true sense. They are omnivorous, so they will eat fruit and seeds as well as meat. There are about three deaths annually from bears in North America. In recent times, black, grizzly, and polar bears have hunted, killed, and eaten humans on rare occasions (Schmidt and Clark 2018, Smith and Herrero 2018). 4 Copyrighted Material Some pages are omitted from this book preview. Chapter 1 - Anxiety, Vision, and Snakes

Of the large canids, only the gray wolf remains. Documented attacks on humans by gray wolves are few and far between. There is one documented human death from a wolf in 2005, none before, and only one attack on a human after that date (MacNay 2007, Butler et al. 2011). Some ranchers in the western United States still insist on killing them and strongly oppose efforts to re-establish wolf populations near their ranches. They cite the economic loss of stock animals—yet we can’t help feeling that it is those ancient demons that are responsible for the ranchers’ attitudes. Wolves have a bad and quite undeserved reputation. After all, many of us live with the domesticated form of the wolf and consider them our best friends. The reintroduction of the gray wolf in Yellowstone National Park has also provided an opportunity to study human behavior toward large carnivores. Of course, the wolf did not stay within the boundaries of the park, but expanded its distribution into areas where people were raising cattle and other livestock. Solomon (2018) was told by ranchers, “Wolves are Democrats.” He then explains, “Ranchers view wolves as a symbol of big government and regulations and all the ways that distant bureau- crats and coastal elites want to destroy the cherished rural ranching culture of the West.” This attitude betrays the ancient anxiety, exposes the deeply divided politics of America, and reveals an attitude that some humans believe they can do what they please with the ecosystem. Humanity’s war on hypercarnivores has been a resounding success. Large carni- vores are damn hard to find; humans are all over the planet. There are now more than 7.5 billion humans living in the biosphere. Medical technologies have increased human lifespans as well as overall health, and the population has exploded. We have developed the ability to control our population size through medical technology— something that almost all other animal species do through evolution—but ancient belief systems and ignorance have prevented some of us from taking advantage of this important technology. Humans have become the most abundant large animal and the most dangerous animal on the planet. The danger of human population growth is not just to the planet’s wildlife; we are a danger to ourselves. In 1908 Teddy Roosevelt wrote, We are coming to recognize as never before the right of the Nation to guard its own future in the essential matter of natural resources. In the past we have admitted the right of the individual to injure the future of the Republic for his own present profit. In fact, there has been a good deal of a demand for unre- stricted individualism, for the right of the individual to injure the future of all of us for his own temporary and immediate profit. The time has come for a change. As a people, we have the right and the duty, second to none other but the right and duty of obeying the moral law, of requiring and doing justice, to protect ourselves and our children against the wasteful development of our natural resources, whether that waste is caused by the actual destruction of such resources or by making them impossible of development hereafter.

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Since Roosevelt wrote those words, the human population has more than doubled. Most of our ancestors gave up hunting, scavenging, and gathering as a way to obtain food about 12,000 years ago (plus or minus a few thousand years). They settled into agricultural communities as the food supply became reliable. This change of lifestyle allowed humans to specialize; not everyone needed to be a farmer. Technological advances allowed many of our ancestors to move into industrialized cities. Today, buildings protect us from temperature extremes and severe weather events. Vaccines protect us from communicable diseases that killed many of our ancestors, and we lead healthier lives because of our understanding of the germ theory of disease and simple methods of sanitation. There are still places in the world where people live alongside large carnivores. Big cats, bears, and wolves are relatively rare because of humans. Of course, there are sharks, but humans are not an aquatic species, and most of us spend relatively little time in the water where we are exposed to a shark attack. However, big snakes are still relatively abundant in South America, Africa, and Asia, and many of them live in close contact with people. Humans have an incredible skill for seeing snakes. Experimental evidence suggests that the danger that snakes posed to our ancestors may be responsible for the evolution of our excellent vision. This has been called the snake detection theory. Lynne Isbell, a behavioral ecologist at the University of California, Davis developed the theory after an encounter with a cobra. The encounter would have been at the closer range if her eyes hadn’t quickly detected the snake and alerted her brain to the snake’s presence. Since that encounter, Isbell and colleagues have performed a variety of experiments to find support (or denial) for the original hypothesis. Isbell first published her snake detection theory in 2006. She hypothesized that lemurs have poorer vision than African monkeys and apes because lemurs are known only from , a continental island with no venomous snakes and where constrictors tend to be relatively small. Neuroscientists at the University of Toyama in Japan and the University of Brasilia in worked with Isbell to find evidence for her theory in brain-based research. In a paper published in 2013, the team described how images of snakes affect the pulvinar—a cluster of neurons in the thalamus, an evolutionarily ancient part of the brain. Pulvinar neurons are believed to help direct our attention to a potential threat. have larger pulvinar clusters than other animals, and certain parts of the cluster are unique to primates. Primates evolved a snake detection-avoidance system for avoiding envenomation. Other relied on the development of different systems, such as chemical resistance to snake . The researchers inserted electrodes into the brains of two captive-born Macaque monkeys that had never encountered snakes. They measured the electrical spikes from individual neurons in two regions of the pulvinar cluster while showing the

6 Copyrighted Material Some pages are omitted from this book preview. Chapter 1 - Anxiety, Vision, and Snakes monkeys images of snakes both coiled and elongated. They also observed and recorded responses from images of Macaque faces with both angry and neutral expressions, Macaque hands in various positions, and geometric shapes such as circles and stars. The snake images stimulated the pulvinar neurons more than any other images. Of the 91 neurons that became active during the experiment, 40% were more active when the Macaques were viewing snake photos than other photos. The neurons fired more frequently than when the monkeys were responding to faces, hands, or shapes. The photos that came in second for causing the neurons to fire were angry monkey faces. Also, the snake-responsive neurons became activated about 15 milliseconds faster than those that responded to angry faces and about 25 milliseconds faster than the neutral shape-detecting neurons. Isbell and her colleagues concluded that the experimental results support the idea that primates have built-in neural mechanisms for recognizing snakes as a specific threat. During our long evolutionary history in Africa, our ancestors had to adapt to the local hypercarnivores and venomous snakes. The view of humans and their ancestors living alongside giant snakes is supported by the fossil record. One of our ancient relatives, Australopithecus anamensis inhabited Kanapoi, a site in the Rift Valley of southwest of Lake Turkana. The hominin are dated at 3.9 and 4.2 Ma. Australopithecus anamensis was bipedal and living in a savanna environment with a diverse fauna. Grazing mammals were more numerous than browsing mammals. The fossil beds included remains of fish, turtles, crocodiles, elephants, giraffes, rhinoceroses, , , hippos, and (Harris and Leakey 2003). The most common squamate fossils at Kanapoi were from a large python similar to Python sebae (Head and Muller 2018). Australopithecus anamensis males reached 50 kg and 1.52 m; females were slightly smaller at 32 kg and 1.34 m. Their teeth were similar to modern humans and show wear from eating coarse vegetation. Australo- pithecus anamensis was potential prey for the giant python of Kanapoi. A more recent fossil site in the Buia area of Eritrea also held remains of an early hominin and a python (Delfino et al. 2004). It seems likely that the Buia python exceeded five meters. The hominin present was Homo erectus, a species that attained a maximum weight of 68 kg and a maximum height of 85 cm in maximum height. (Anton, 2003). While this size and weight may have been reached by some individuals, they would be maximums, and most H. erectus would have been smaller as children, adolescents or even adults. Weights reported for the African Python are in the 55–65 kg range and may be greater (Pitman, 1974). Thus, hominins and giant pythons were inhabiting the same biotic communities since the appearance of the hominins in the fossil record. There is also evidence that behavior has been shaped by snakes in multiple lineages of living primates. Gardner et al. (2015) observed a female Coquerel’s Sifka, a lemur, captured by a Madagascar Ground Boa. Members of the female lemur’s troop attacked the snake and injured it so badly that it later died from its injuries. In Costa 7 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes

Rica, Boinski (1988) observed an adult male White-faced Capuchin Monkey hitting a pit viper with a branch. Chimpanzees communicate danger to other chimps using alarm calls. Crockford et al. (2012) demonstrated that chimpanzees were more likely to use an alarm call when a group of chimps was unaware of the presence of a snake compared to when the group was aware of the snake’s presence. There is also a video of a captive chimp flipping a harmless snake with its hand towards the fence of its enclosure in an attempt to get rid of it. One of the authors (TC) was recently in the Peruvian Amazon. As an experiment, he and Bill Lamar showed a Boa Constrictor to a group of habituated Squirrel Monkeys, and the monkeys literally went into fear mode, screaming warn- ings. On the same expedition, a wild bushmaster (Lachesis muta) was discovered in the daytime, causing a troop of Black Saddle Tamarins to seemingly miraculously appear above them screaming warnings to them and to each other. It seems wild primates are as afraid of snakes as most of the human population. We don’t want to imagine a world without giant snakes. This line is in fact how we feel about giant snakes, but it was also part of the title of a 2017 article about Shariar Caesar Rahman (Hance, 2017). Rahman is the CEO of the Creative Conservation Alliance, an organization that started a python project in 2013 at Lawachara National Park in northeast . The 1,250-hectare park is completely surrounded by villages and tea plantations. The park contains Burmese Pythons, the same species that has invaded southern Florida. Conflict between humans and the Burmese Python would seem preordained. The Creative Conservation Alliance team hired and trained some villagers to capture and handle the pythons. Villagers called the trained “parabiologists” when they had a problem with a python. Ten pythons were rescued and surgically implanted with radio transmitters. The team discovered the snakes preferred life in the village. One female snake was captured and returned to the forest eight times in a year. While the species is invasive in the United States, it is considered Vulnerable by the IUCN within its natural distribution. Snakes had a direct impact on primate evolution, and they helped make us the success we are today. Anxiety, our excellent vision, and our communication skills seem to be the result of living with a dangerous fauna, one that included snakes. Snakes are symbols for cults, religions, and medicine around the world. This book looks at the natural history of the largest snakes and their relationships with us in the past and the present. We know relatively little about the lives of giant snakes. This book is an attempt to summarize what we do know about the largest living serpents and to increase appreciation for them.

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Chapter Two Size and Shape

9 Copyrighted Material Some pages are omitted from this book preview. Chapter 2 - Size and Shape

Limbless, cylindrical bodies provide snakes with opportunities not available to species with more conventional tetrapod body forms. Looking at a snake, an informed observer can usually tell if the snake lives in trees, in the water, below the surface of the ground, or on the surface. Snake body forms reflect their macro- , hunting methods, prey size, prey shape, and ancestry. Arboreal snakes have low body-mass/length ratios (a slender, lightweight body), laterally compressed bodies, long tails, large eyes, a large vertebral scale row, and a center of gravity that is more posterior than non-tree-dwelling species. Many arbo- real snakes have ventral scales with a keel: a bent edge on each side. Aquatic snakes have high body-mass/length ratios, cylindrical bodies that can be laterally compressed when swimming, eyes and nostrils dorsally oriented on the head, valvular nostrils, and no enlarged vertebral scale row. They frequently have short tails, but some have quite long tails, depending upon their ancestry. Many aquatic species have ventral scales reduced in width and overall size. Fossorial snakes—snakes that live underground—have fused head scales, small eyes, narrow, pointed heads, often with a reinforced rostral scale, and short tails. Like aquatic snakes, they may have valvular nostrils. The dorsal scales are often iridescent because of their ultra-smooth surface, which prevents particles from adhering to them. Terrestrial snakes have laterally oriented eyes and nostrils. They vary in body size, tail length, and eye size. They usually have broad ventral scales without keels. Of course, not all snakes are perfectly adapted for each of these lifestyles. Many use a combination of , so they may spend time in trees or bushes, bask and hunt in the water or live in a burrow in the mud at the edge of a pond or stream and actively forage in the water or on the ground. Snakes with long heads tend to specialize in feeding on mammals, while snakes with shorter heads are usually dietary generalists (Pearson et al. 2002; Vincent and Herrel, 2007). Head-length/body-length ratios may change during an individual snake’s lifetime as they shift from a generalist juvenile diet to a more specialized adult diet. Head size may also be related to in species where the sexes have different diets.

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Size is fundamental to a snake’s biology. Body size determines the size of prey an animal can kill and consume, the number of or young a female can hold, how quickly the animal can warm or cool its body, and how rapidly water is lost through its skin. Size may also influence success in social endeavors like male combat or mating. As ectotherms, snakes depend upon their environment for heat, and they have the advantage of requiring fewer calories from food than endothermic birds or mammals of similar weights. The disadvantage of being an ectotherm is sluggish- ness at low temperatures. Long, cylindrical, bodies can quickly collect heat from the environment by stretching out to increase the body’s surface area. The same behavior at low temperatures causes rapid heat loss. Snakes can conserve heat by coiling their body to reduce the surface area exposed to the environment. Larger snakes will warm and cool slower than smaller snakes. Thus, being large can lower the cooling rate and help the snake retain heat. The ability to hold heat is thermal inertia. Color and reflectance also play a role in heating. Snakes with darker bodies collect heat more quickly than snakes with lighter colors. Snakes with darker coloration tend to live at higher altitudes, higher latitudes, or use cooler microenvironments like deep shade forests, than their lighter colored relatives. Large body size can also accommodate larger prey. Thus, a bigger snake can take in more calories per meal. A correlation between the body size and the size of its prey is well known (Shine 1991). Large snakes usually take large prey and often delete small prey from their diet, giving them a competitive edge because they can use prey that smaller snakes cannot. Natural selection is continually monitoring individuals for being larger or smaller in all environments. On islands or mountaintops, available prey may be small or in short supply and snakes downsize for energy efficiency. Or larger prey may be seasonally available, like newborn , and snakes are selected for increased size to be able to consume the larger prey and store the energy for lean times. Size may also be a way to avoid predators. Giant snakes are going to have fewer predators than small snakes. Small size means more potential predators, but smaller animals also have more places for concealment. Increased body size is likely expensive in terms of calories, thus benefits from it are probably substantial. These could include increased survival of offspring and resistance to environmental extremes. Sexual Size Dimorphism Male and female animals that differ in size, shape, coloration, behavior, and other characteristics are said to be sexually dimorphic. For quantifying differences in body sizes of male and female snakes, the mean adult body length of the bigger sex is divided by the mean adult body length of the smaller sex and subtracted from 1.0. The number is arbitrarily given a minus sign if the male is larger and no sign if

12 Copyrighted Material Some pages are omitted from this book preview. Chapter 2 - Size and Shape the female is larger. No difference between the sexes is scored as a zero. The greater the difference in the sexes, the larger the number. Studies on sexual size dimorphism (SSD) in snakes suggest species with larger males have males competing for females through combat or ritualized behavior, where a large male body size increases the probability that larger males will sire more offspring (Shine 2003, Dubey et al. 2009). Of the giant and near-giant snakes, the only one known to have males larger than females is the Scrub Python (Simalia kinghorni). This may also be true for some of the lesser known, poorly studied near- giant species. One of the most exhaustive studies done on wild Reticulated Pythons exam- ined more than 1000 specimens from southern . (Shine et al. 1998a, b) The researchers found that females attained much greater body sizes than males, even though they had similar head lengths at the same body size. Females achieved a maximum mass of 75 kg, whereas the maximum mass of males was only 20 kg. The largest females reached 6 meters, while the largest males reached only 3.5 meters. Rivas and Burghardt (2001) found adult male Green (Eunectes murinus) were in the 2–14 kg range, whereas adult females ranged from 8 to 80 kg. Dirksen (2002) found males reached a maximum length of about 3.4 meters, whereas females grew to 6.5 m. Dorcas and Willson (2011) reported measure- ments of Burmese Pythons (Python bivittatus) in the Florida . Their data included males measuring 0.914–3.96 m and females measuring 0.914–5.18 m, with fewer males in the larger size classes. Burmese Pythons appear to become sexually mature at about 2.7 m, and most of their animals were below this size. However, the largest individuals were females. Thus, females of most giant snake species are significantly larger than males. Larger body size in one sex reduces competition for food between the sexes. Females can feed on larger prey than males, which reduces competition with males. Larger females can hold more eggs and offspring. In nature, males of these species typically trail the females and court them, often in groups, a mating strategy termed scramble competition. Species using this mating strategy often form mating balls, where the female is surrounded by dozens of males in a writhing mass of snakes that looks like a ball. Species that use scramble competition are thought not to use combat or ritualized competition bouts between males to determine which male will mate with the female. Becoming A Giant Species Why do some species become giants while others do not? Factors influencing body size include food availability, sexual selection, competition from other species, and predators.

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In a review of gigantism, Vermeij (2016) defines a giant as one species that is the largest member of its clade, trophic level, or ecosystem either at a particular time or for all time. He defines a local giant as the largest-bodied species in a specific loca- tion or period. Species that are the largest members of their ecological (trophic and habitat) category at the global spatial scale and throughout the Phanerozoic eon (541 Ma to the present) are referred to as global giants. Thus, gigantism appears tens to hundreds of millions of years after mass extinc- tions and long after the origins of clades in which the giants evolved. Giant marine species correlate with high planktic or seafloor productivity. However, on land the correspondence between photosynthesis productivity and gigantism is weak. Global giants are aerobically active animals, not gentle giants with low metabolic demands. He found oxygen concentration in the atmosphere correlates with gigantism in the Paleozoic but not afterward. The evolution of efficient gas-exchange systems was present in clades containing giants. Although temperature and habitat size are essential in the evolution of immense scale in some cases, the most critical enabling circumstance is a highly developed ecological infrastructure in which vital resources are abundant and efficiently recycled and reused, permitting activity levels to increase and setting the stage for large animals to evolve. Following Vermeij (2016), the snakes discussed here are best considered local giants. Head and Polly (2007) found snake body-size significantly correlated with the number of vertebrae (pleomerism) in many lineages. Therefore, increasing the number of body segments (=vertebrae) produced during embryonic development can be a factor in the evolution of body size. They examined the vertebral count in snakes that exhibit gigantism: booids, pythonids and the typhlopids. Although members of the blind snake family () are small and could not be considered giants, there are a few huge species (relative giants) within this family of usually small snakes. The relatively giant blind snakes were included in the Head and Polly (2007) study, and they found that Typhlops and Rhinotyphlops possess a positive relationship between body size and the number of vertebrae, confirming that pleomerism is essential in these snakes. However, the giant boas and pythons have fewer than expected vertebrae, suggesting that a separate process is responsible for gigantism in boas and pythons. The lack of correlation between body size and the vertebral count in giant snakes demonstrates the dissociation of segment production in early development from body growth during maturation. Thus, gigantism in boas and pythons is achieved by modifying development at a different stage in the snakes’ life history. Climate also has a role in giant snake evolution. In a survey of terrestrial ecto- therms, Makarieva et al. (2005) analyzed body-length data for the largest representa- tives of 24 taxa of terrestrial ectotherms from tropical, temperate, and polar envi- ronments. They found ectothermic terrestrial giants become shorter for each 10°C

14 Copyrighted Material Some pages are omitted from this book preview. Chapter 2 - Size and Shape decrease in ambient temperature. They quantified the reduction in maximum body size to compensate for the drop in metabolic rate required by the lower temperature. The results suggested that the upper limit to body size within each of the ectotherms is set by a temperature-independent critical minimum value of mass-specific meta- bolic rate. A fall below this rate would impact the organism’s biological perfor- mance and decrease its chances of survival. Therefore, larger ectotherms evolved at warmer temperatures. The discovery of Titanoboa in the Cerrejón Formation of Colombia, South America by Head et al. (2009) added evidence to the hypothesis that giant ectotherms increase or decrease their size with temperature fluctuations. The fossil vertebrae from Titanoboa were dated at 58–60 Ma, and the snake may have reached 13 m in length and about 1,135 kg in weight. Head et al. used the relationship between temperature and size derived by comparison to modern organisms. The authors calculated a snake of this size would require a minimum mean annual temperature of 30–34°C to survive, an estimate consistent with the hypothesis of a hot in the Neotropics. A high concentration of atmospheric carbon dioxide was evident at the time. Comparison of paleotemperature estimates from the equator to those from South America’s mid-latitudes suggests a steep temperature gradient during the early Paleocene greenhouse conditions, similar to that of today. The paleoenvironment and faunal composition of the Cerrejón Formation indicate Titanoboa was leading an anaconda-like existence. Head et al. estimated that Titanoboa required minimum temperatures of 32–33°C, temperatures 6–8°C warmer than temperatures reconstructed from plants within the same formation, and much warmer than modern values. If giant fossil snakes can in fact act as paleothermometers, and more giant snakes can be discov- ered, it may provide insights into paleoclimates and a better understanding of giant snake evolution. Allometry, Heterochrony, and Growth Clifford Pope opened his classic book, The Giant Snakes, with a chapter on Sylvia, a Python bivittatus who was found on October 3, 1945 in a sugar barrel behind an army mess hall. She was brought to the United States by a colleague of Pope’s and delivered to him at the Chicago Natural History Museum (now the Field Museum) in December of the same year. When, Sylvia arrived in Chicago, she was just over a meter in length. Pope raised her and took monthly measurements. Sylvia’s size at birth was unknown, but from February 1946 it took just 15 months for her to double her length. A 13-year study by Madsen and Shine (2000) on free-ranging Water Pythons (Liasis fuscus) and their prey, the Dusky (Rattus colletti), at Fogg Dam in the 15 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes

Northern Territory of Australia revealed an impressive result on growth. Water Pythons are about 450 mm (17.7 inches) in body size at birth, and they can reach an adult length of 3 m. The study included body-size data on 471 snakes of known age that were marked after they had hatched in captivity. Madsen and Shine (2000) found a high correlation between snake growth rates and the availability of prey. Growth rates between siblings were not correlated, suggesting that environmental factors were involved in growth. Young snakes grew faster in years of abundant prey than those produced in years when prey was scarce. Individual snakes that grew quickly in their first year of life continued to grow during the study. The authors termed this the “silver spoon” effect, a phenomenon created by an abundance of food in the first year of life, setting the stage for long-term growth and adult body size. This was most obvious in the faster-growing females. The was more abundant in years with higher rainfall, and Madsen and Shine suggested that the current distribution of body sizes in the population is the result of the annual variation in precipitation during the preceding decades. Expanding on this work, Ujvari et al. (2011) examined the effects of climate on the annual food supply and found that a flood-induced famine reduced the body size at which females became mature. This suggests that body size and sexual maturity in pythons is plastic and influenced by food availability when the pythons are young. Postnatal growth in usually maintains geometric proportions. A common exception is head size, which may show allometric growth. That is, the head may be proportionally larger at birth, and the body grows slightly faster than the head after birth or hatching. However, linear dimensions increase at about the same proportional rate. A measure of length (L), therefore, is a suitable estimate of size. At the same size and time, the surface area of the skin, or a cross-section of the body, or a cross- section of an organ increases in proportion to the square of the length (L2), whereas volumes increase proportionally to the cube of the length (L3). Density remains almost constant, so body mass shows the same relation as volume (L3). Therefore, if one animal is twice the length of another, its body mass will be approximately eight times as much (Scanlon 2014). Applying this to the Green Anaconda, a 2.088 m specimen weighs 5.9 kg. A 4.16 m individual would weigh 47.2 kg, and an 8.32 m individual would weigh 377.6 kg. This would be a massive snake, and it is easily understood why snakes with thick, long bodies like the anaconda are aquatic: they require water to help support all that weight. By contrast, consider an 8 m python captured in that weighed a mere 150 kg. Reticulated Pythons (Malayopython reticulatus) have a much more gracile build and are less aquatic and more arboreal than the anaconda. This is not to say that they do not spend time in the water. Reticulated Pythons do enter the water and will hunt from the water, but they also climb trees.

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Evolution can change the body form of organisms by altering the rate at which different parts of the body grow (allometry) or by modifying the timing or rate of events (heterochrony) that occur during growth. The term allometry was coined by Huxley and Tessier (1936) while studying the growth of the massive claw of the male Fiddler Crab. They found the claw grew at a faster rate than the rest of the body. Heterochrony is a change in developmental timing. It can refer to the mecha- nisms that embryos use to measure time during development. Recently, studies have examined the evolution of increased segment number in snakes, which is a heterochronic modification of the somite clock in the snake embryo that has been an important driver of evolutionary change. If the somite clock speeds up, the snake will have more vertebrae, more ventral scales, and a longer body. If the clock slows, the snake will have fewer vertebrae, fewer ventral scales, and a shorter body. Esquerré et al. (2017) examined how allometry (growth of different body parts at different rates) and heterochrony (changes in timing or rates that result in changes in size or shape) drive morphological diversification of pythons. They found heteroch- rony to be the primary driver of initial divergence within python clades and shifts in allometric trajectories make it possible for pythons to have novel head shapes and body sizes. The broad pattern they found is that pythons tend to get broader heads and proportionally smaller and more dorsally situated eyes as they grow. Falk et al. (2017) noted that body condition is frequently estimated as a body condi- tion index (BCI) using length and mass measurements. Female Burmese Pythons exhibited more positive allometry than males in both total mass and fat mass, and fat mass was more positively allometric than total mass in both sexes. Males and females exhibited different allometric relationships between length and mass. Boas and pythons are often very similar in appearance. Similarities in head shapes, body sizes, color patterns, behavior, and diet in both clades are the result of these developmental processes that are selected for by evolution and produce snakes that look similar to each other but have entirely different ancestors. Ontogeny and Snakes Ontogeny refers to the development of an individual from the time the is fertilized to death. Here we use the term to refer to changes that an organism under- goes post-birth (or hatching). Some snakes undergo a color change from when they hatch until they mature because they also change their habitat. Venomous species change their chemistry from birth until they increase their body size because they switch to a different prey, often eating frogs or lizards as juveniles and mammals as an adult. Schuett et al. (2005) manipulated the diet of juvenile Boa Constrictors to test the hypothesis that prey size induces phenotypic plasticity of the skull and jaw, as well

17 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes as to see if the onset of sexual size dimorphism was determined under a feeding schedule where total prey mass consumed by the snakes was held constant. They used 23 neonate Boa Constrictors from a litter sired by one male. The neonates were placed into two treatment groups, but all were maintained under identical environ- mental conditions. Group 1 was fed weanling mice throughout the entire study; group 2 was fed weanling mice, followed by of increasing size as the snakes grew larger. At the end of the study, group 1 consumed more meals, but both groups consumed an equivalent mass of rodents. The snakes were measured at 5 weeks and 58 weeks. Between-group differences of the skull and jaw sizes were not significant; thus, prey size did not exert an influence on growth of the head bones. In contrast, sex differences accounted for significant differences in body length, total length, and body mass. They found sexual dimorphism emerged early during ontogeny under conditions where prey mass consumed was held constant, suggesting that a genetic influence is likely. Pritchard’s Rule Pritchard’s Rule (Pritchard 1994) predicts that the maximum size of a snake species will be 1.5 to 2.5 times the minimum adult female length. Rivas (2000) found the smallest mature female Green Anaconda to be 2.10 m, so the maximum expected size would be 5.25 m. Willson et al. (2014) reported a 2.10 m female Python bivittatus that contained 11 eggs. So, Pritchard’s Rule would again predict the maximum-sized individual to be about 5.25 m. The size of the North African Python Python( sebae) and the South African Python (P. natalensis) at sexual maturity has been confused in the literature. Branch (1988) estimated sexual maturity at 200-300 cm. Therefore. the maximum estimated size would be 7.5 m (using 2.5 m x 3) for one or both of these species. At the time of his writing, all African Pythons were P. sebae. Minimum size at sexual maturity for P. sebae was considered 1.9 m by Luiselli et al. (2007). Following this rule, its maximum size would be about 6.25 m. Shine et al. (1999) found the smallest mature female Reticulated Python out of 1000 snakes examined from Sumatra was 2.5 m, which would suggest the maximum size for P. reticulatus is about 6.25 m. The problem with this system is the phenoplasticity of snakes in the wild and in captivity. Well-fed captive pythons may become sexually mature at smaller sizes than free-ranging pythons that have to hunt for a living. Also, most of the snakes Pritchard used to develop the rule were colubrids and booids, and pythonids may not follow the same patterns. Thus, these predictions are not hard numbers. Table 2-1 shows the size data for neonates, mature animals, and maximum lengths and weights, as well as maximum size estimates, using Pritchard’s Rule calculations (in parentheses, using 2.5 as a multiplier). 18 Copyrighted Material Some pages are omitted from this book preview. Chapter 2 - Size and Shape

Table 2-1 Size & Weight Data for Neonates, Mature Animals Hatchling/ Size at sexual Est. Maximum Common Name Maximum Neonates maturity Weights Size Using Scientific Name Lengths TL MM TL MM Pritchard’s Rule Boa Constrictor1 430–575 M 0.9–1.6 ~30 kg 4.9 m 5.75 m Boa constrictor F 1.2–2.3 Green Anaconda2 508–956 M 1.8–2.8 ~200 kg 7.79 m M 7.0m Eunectes murinus F 2.4–3.5 F 8.7m Yellow Anaconda3 409–593 M 1.5–2.5 50 kg 4.25 m M 6.25m Eunectes notaeus F 2.3–4.25 F 10.6m Burmese Python4 ? 8.22 m 480–790 M 2.0 182 kg M 5.0m Python bivittatus F 2.6 5.75 m confirmed F 6.5 m Indian Python5 ? 5.5 m 480–790 M 2.0 52.2 kg M 5.0m Python molurus F 2.6 4.6 m confirmed F 6.5m South African Python6 500–700 M 1.7–2.5 5.8 m M 5.0m Python natalensis F 1.7–3.0 F 6.5m North African Python7 500–700 M 1.7–2.0 113 kg ~6.5 m M 5.0m Python sebae F 1.7–2.0 F 5.0m Reticulated Python4 ? 10 m 550–850 M 1.6–3.5 158 kg M 8.75m Malayopython reticulatus F 2.4–4.5 6.35 m verified F 11.25m Scrub Python8 ~8 m 633–729 M 2.43 91 kg M 6.1m Simalia kinghorni F 2.33 5.6 m verified F 5.8m Western Olive Python9 550-880 ? 2.5-3m 9.3 kg ? 6.5 m 7.5 m Liasis olivaceus barroni Papuan Python10 609-698 ? 22.5 kg 5.13 m ? Apodora papuana Oenpelli Python11 830– ? 6 kg 4.57 m ? Morellia oenpelliensis ?1,000 References: 1. Snow RW, Krysko KL, Enge KM, Oberhofer L, Warren–Bradley A, Wilkins L. 2007. Introduced populations of Boa constrictor (Boidae) and Python molurus bivittatus (Pythonidae) in southern Florida. Biology of Boas and Pythons. Edited by RW Henderson and R. Powell. Eagle Mountain Publishing, Eagle Mountain, Utah, USA. 2007:365–86. 2. Murphy JC, Henderson RW, Henderson RW. 1997. Tales of giant snakes: a historical natural history of anacondas and pythons. Krieger Publishing Company, Malabar. 3. Waller TO, Micucci PA, Alvarenga ER. 2007Conservation biology of the yellow anaconda (Eunectes notaeus) in northeastern . . Pp. 340–362 in: Biology of the boas and pythons. Eagle Mountain Publishing, Eagle Mountain. 4. Barker DG, Barten SL, Ehrsam JP, Daddono L. 2012. The corrected lengths of two well–known giant pythons and the establishment of a new maximum length record for Burmese pythons, Python bivittatus. Bulletin of the Chicago Herpetological Society. 47(1):1–6. 5. Minton SA. A contribution to the of West . Bulletin of the AMNH; v. 134, article 2. And Barker et al. 2012. 6. Branch B, Erasmus H. 1984. Captive Breeding of pythons in , including details of an interspecific hybrid (Python sebae natalensis X Python molurus bivittatus). The Journal of the Herpetological Association of Africa. 30(1):1–0. 7. Broadley DG. 1983. FitzSimons' snakes of . Delta Books. 8. Freeman A. 2016. A study in power and grace: The . Wildlife Australia, 53(3):27. 9. Whitlock, F. 1923. A trip to the and Hamersley Ranges, Northwest Australia. Emu 22: 259–273. 10. Tryon BW, Whitehead J. 1988. Reproduction in a little‐known Python, Liasis papuanus (Peters and Doria). Zoo Biology 7(4):371–9. 11 . Gow, C. 1989. A complete guide to Australian snakes. Angus and Robertson, Sydney. 19 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes

Acceptable Evidence for Giant Size What is acceptable evidence for the size of a giant snake? Snakes, like fish, stim- ulate the human brain to embellish and exaggerate. Anyone who has spent time looking for snakes is familiar with the local telling a story about the one they saw yesterday, last month, or last year. Or, the one killed by their neighbor or grandfather. Herpetologist James Oliver (1958) was well aware of the tendency for humans to exaggerate in his book Snakes in Fact and Fiction. He noted the existence of human interest in extremes, the difficulty of catching a huge snake, the challenge of accurately measuring a giant snake, and the fact that scientists are usually more interested in average sizes than the extreme size because the largest specimen of any animal is likely to be an “abnormal giant.” Clifford Pope also discussed the problem in his 1961 book, The Giant Snakes, and noted three approaches to determining the valid size of snakes. The first approach is strictly scientific and demands concrete proof. The snake is measured alive or dead. The specimen is in a museum collection or zoo, so that the measurements can be verified. Therefore, measurements taken in the field without the animal for a voucher would be unacceptable. The second approach is to accept field measurements made by experienced explorers. Both Oliver and Pope took this approach. It assumes that people are intellectually honest, have good memories, will not make exaggerated claims, and can control their emotional reaction when they find an enormous snake. The third approach is to accept stories at face value. Included in this category would be second- or third-hand stories. Using this approach assumes that people don’t exaggerate or misinterpret observations they make about snakes, and if multiple people are reporting 12, 15, or 20 m snakes, there must be some truth to the stories. Pope and Oliver considered the second method acceptable. Unfortunately, this approach has produced record-sized snakes that do not stand up to scrutiny over time. Investigation of some long-established records has revealed once-held maximum sizes reported for the Boa Constrictor, the Anaconda, and the Reticu- lated Python have serious flaws. The maximum-size records of snakes are published in somewhat obscure places where they can be overlooked. Often reports of giant snakes may be taken at face value because of a person’s reputation and are picked up by writers so that they become part of popular culture. Therefore, encyclopedia articles and record books may contain misleading information passed from one generation to the next. Murphy and Henderson (1997) attempted to draw attention to these problems, but were minimally successful based upon the fact that many references to giant snakes still contain the same erroneous measurements reported in the older literature. While each approach has its problems and advantages, there is only one that is acceptable. 20 Copyrighted Material Some pages are omitted from this book preview. Chapter 2 - Size and Shape

A Critique of the Three Approaches The third approach proposed by Pope would accept size estimates and hearsay. If the reporter was credible and not prone to exaggeration, the possibility exists that some people encountered enormous snakes. An example is anthropologist Mary Kingsley’s (1897) hearsay description of a giant African Python. She wrote, I am assured by the missionaries in Calabar that there was a python brought into Creek Town in the Rev. Mr. Goldie’s time that extended the whole length of the mission-house verandah and to spare. This python must have been over 40 feet [12.1 m]. I have not a shadow of doubt it was. Kingsley traveled extensively in , she was a trained scientist, and in her judgment the report was accurate. The second approach proposed by Pope and Oliver would accept Colonel Percy Fawcett’s 1907 encounter with an 18.9 m Anaconda. This oft-cited story is used to support the existence of super-sized snakes. Fawcett was a Royal Artillery officer who had served the British government in Ceylon, did secret service work in North Africa, and led a life of adventure. Bored with army life, he retired to do survey work when he received an offer from the Royal Geographic Society early in 1906. The Society asked Fawcett to survey parts of that had no accurate maps. While near the Peruvian-Bolivian border, Fawcett and his team happened upon a huge snake. After encountering the snake, and killing it, he wrote, We stepped ashore and approached the reptile with caution. It was out of action, but shivers ran up and down the body like puffs of wind on a mountain tarn. As far as it was possible to measure, a length of forty-five feet [13.7 m] lay out of the water, and seventeen feet [5.1 m] in it, making a total length of sixty-two feet [18.8 m]. Its body was not thick for such a colossal length—not more than twelve inches in diameter—but it had probably been long without food. I tried to cut a piece out of the skin, but the beast was by no means dead, and the sudden upheavals rather scared us. A penetrating foetid odour emanated from the snake, probably its breath, which is believed to have a stupefying effect, first attracting and later paralyzing its prey. Everything about this snake was repulsive. In May of 1925 Fawcett entered an uncharted area of Brazil with his son Jack and one of Jack’s friends. The three men were never seen again. Fawcett’s son Brian compiled his father’s memoirs. Discussing the above story, Brian wrote: “When this serpent was reported in , my father was pronounced an utter liar!” Fawcett was probably not a liar, but most certainly a shameful exaggerator. It seems likely that Fawcett happened upon an enormous Anaconda, but in all proba- bility it wasn’t 18.8 m long. And if it smelled as bad as he suggests, it might have been dead when he found it. The size, the odor, and the suggestion that it was only 0.3 m in diameter make this story unbelievable. Anacondas are well-known snakes, and 21 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes

Calabaria reinhardtii volontany Sanzinia madagascariensis cf. dumerili Madagascar Acrantophis madagascariensis Acrantophis dumerili } continentalis placata } Lichanura trivirgata West United States bottae Charina unbratica } & muelleri Africa Eryx jayakari Eryx conicus & Eryx johnii } Middle East Eryx colubrinus Africa Eryx jaculus } Eryx elegans Eryx tataricus vittatus Eryx miliaris Eryx tataricus Asia Eryx miliaris nogaiorum} Melanesia, Micronesia bibroni Candoia aspera Candoia carinata Candoia superciliosa Candoia superciliosa crombiei } Mexico & Boa imperator Boa imperator sabogae } Central America Boa constrictor occidentalis Boa constrictor amarali Boa constrictor constrictor Corallus annulatus Corallus caninus Central America, Corallus hortulanus South America, Corallus cookii Eunectes murinus Eunectes notaeus Epicrates alvarezi Epicrates cenchria Epicrates maurus Epicrates assisi Epicrates crassus Chilabothrus inornatus Chilabothrus monensis granti Chilabothrus subflavus Greater Antilles Chilabothrus fordii Chilabothrus chrysogaster Chilabothrus exsul Chilabothrus strigilatus fosteri Chilabothrus strigilatus Chilabothrus strigilatus mccraniei BS>50%, PP>.095 BS<50%, PP>.095 BS<50%, PP<.095 Python regius Africa Southeast Asia Figure 2-1. Molecular technologyPython curtus has provided us with a direct view of the genetic code and allows us to see relationships betweenPython organisms bivittatus at a level we could only guess at two decades ago. Reynolds et al. (2014) produced a phylogeneticPython molurus tree of} pythons and booid snakes based upon molecular evidence. We have modified the treePython to show anchietae the pythonid phylogenyAfrica with the giant species highlighted in blue and the near-giant speciesPython highlighted sabae } in green. Thus, enormous body size has evolved multiple times Maylayopython timoriensis in each of these clades. Maylayopython(See Glossary reticulatus if you are unfamiliar with diagrams of phylogenetic trees.) Maylayopython reticulatusContinued jampeanus on next page. Maylayopython reticulatus saputrai } Morelia carinata 22 & Morelia spilota variegata Morelia viridis Antaresia maculosa Antaresia perthensis Antaresia stimsoni orientalis Australia & New Guinea Antaresia childreni Liasis papuana Liasis olivaceus Liasis fuscus sauvensis Liasis mackloti Similia oenpelliensis Similia boeleni Melanesia Similia tracyae Similia amethistina Similia clastolepis Similia nauta } Similia kinghorni Australia & New Guinea Aspidites ramsayi Aspidites melanocephalus} Bothrochilus hoserae Bothrochilus boa Indonesia & Bothrochilus albertisii} Calabaria reinhardtii Sanzinia volontany Sanzinia madagascariensis Acrantophis cf. dumerili Madagascar Acrantophis madagascariensis Acrantophis dumerili } Central America Ungaliophis continentalis Exiliboa placata } Lichanura trivirgata West United States Charina bottae Charina unbratica } & Mexico Eryx muelleri Africa Eryx jayakari Eryx conicus South Asia & Eryx johnii } Middle East Eryx colubrinus Africa Eryx jaculus } Eryx elegans Eryx tataricus vittatus Eryx miliaris Eryx tataricus Asia Eryx miliaris nogaiorum} Melanesia, Micronesia Candoia bibroni Candoia aspera Candoia carinata Candoia superciliosa Candoia superciliosa crombiei } Mexico & Boa imperator Boa imperator sabogae } Central America Boa constrictor occidentalis Boa constrictor amarali Boa constrictor constrictor Corallus annulatus Corallus batesii Corallus caninus Corallus cropanii Central America, Corallus ruschenbergerii Corallus hortulanus South America, Corallus grenadensis Lesser Antilles Corallus cookii Eunectes murinus Eunectes notaeus Epicrates alvarezi Epicrates cenchria Epicrates maurus Epicrates assisi Epicrates crassus Chilabothrus angulifer Chilabothrus inornatus Chilabothrus monensis granti Chilabothrus subflavus Greater Antilles Chilabothrus fordii Chilabothrus chrysogaster Chilabothrus striatusCopyrighted Material SomeChilabothrus pages exsul are omitted from this book preview. Chilabothrus strigilatus fosteri ChilabothrusChapter strigilatus 2 - Size and Shape Chilabothrus strigilatus mccraniei

Python regius Africa Southeast Asia Python brongersmai Python bivittatus Python molurus } Python anchietae Africa Python sabae } Maylayopython timoriensis Maylayopython reticulatus Maylayopython reticulatus jampeanus Maylayopython reticulatus saputrai } Morelia carinata Morelia bredli Indonesia & Southeast Asia Morelia spilota Morelia spilota variegata Morelia viridis Antaresia maculosa Antaresia perthensis Antaresia stimsoni orientalis Australia & New Guinea Antaresia childreni Liasis papuana Liasis olivaceus Liasis fuscus Liasis mackloti sauvensis Liasis mackloti Similia oenpelliensis Similia boeleni Melanesia Similia tracyae Similia amethistina Similia clastolepis Similia nauta } Similia kinghorni Australia & New Guinea Aspidites ramsayi Aspidites melanocephalus} Bothrochilus hoserae Bothrochilus boa Indonesia & Papua New Guinea Bothrochilus albertisii}

BS>50%, PP>.095 BS<50%, PP>.095 BS<50%, PP<.095

Figure 2-1 continued from page 22. there is no evidence to suggest that they could reach the size stated. Also, healthy, live snakes do not have a bad odor. The Fawcett story is included here because, in theory, it meets the criteria of the second category proposed by Pope. Fawcett was indeed an experienced explorer; however, experience in exploration does not ensure that a person will avoid exag- gerations and fantastic stories. Murphy and Henderson (1997) defined supersnakes as those reported to be in the 15–45.7-meter range, they frequently have huge eyes (boas and pythons have relatively small eyes) that glow in the dark, and they have supernatural powers over humans and other animals. The Fawcett story is a borderline supersnake story because his comments about the animal’s psychic power are restricted to the commonly accepted idea at the time that snakes could mesmerize prey. He merely exaggerates the size and adds the comment about the odor from a story he had prob- ably previously read or heard. Expectations about giant snakes in Fawcett’s mind were undoubtedly predetermined by his previous knowledge of travelers’ tales. 23 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes

The Giants and Near-giants There are six species of extant giant snakes—species that in all likelihood exceed 6.1 meters. They are the Green Anaconda Eunectes( murinus), the Reticulated Python (Malayopython reticulatus), the North African Python, (Python sebae), the South African Python (Python natalensis), the Burmese Python, (Python bivittatus), and the Scrub Python (Simalia kinghorni). Another six species get quite large but may not exceed the 6.1 meters: the Indian Python (Python molurus), the Western Olive Python (Liasis olivaceus barroni), the Oenpelli Python (Morelia oenpelliensis), the Papuan Python (Liasis papuana), the Yellow Anaconda (Eunectes notaeus), and the Cuban Boa (Chilabothrus angulifer). Then there is the common Boa Constrictor Boa( constrictor), a species that rarely exceeds 4 m (13 ft.) in nature or captivity, but is a giant in the mind of the public. Given the plasticity of snakes to grow and the difficulty of measuring live snakes, even under the best conditions, the exact sizes of giant snakes are elusive and are unlikely to be readily verified. Therefore, this book focuses on natural history and giant snake and human relationships instead of body sizes that may or may not be verifiable. Interest in the size of the largest snakes is likely to be a reason the reader picked up this book. Below, we summarize the largest specimens of each species based on the available data. Skin Length vs. Total Length The cover photo of the September 1998 issue of Reptile magazine illustrated a large snakeskin with 16 children and an adult sitting along the dried skin on a lawn. The 8.8 m skin was said to be from a snake that was attacking cattle in Brazil in 1935. The short article reported that the skin was not stretched, and with the head and part of the tail missing, the snake measured close to 9.14 m and weighed more than 227 kg (Moser and Robertson 1998). This record was also commented on by Lutz Dirksen (2002), who stated the skin might have been a 9 m snake in life. In fact, we think it was substantially smaller. Stories about giant snakes based upon skins are ubiquitous. Reading them, you get a sense of certainty that the skin documents the existence of an enormous serpent. However, this is not true. It has been well-known for a very long time that skins removed from the snake stretch. In fact, it is virtually impossible to remove a snakeskin without stretching it about 20% of its length. In fact, the skins may stretch much more than the 20% quite by accident, but on occasion, it is on purpose. Snakes have evolved skin that stretches as an to feeding on large prey. Growing up, I (JCM) remember feeding a small gartersnake multiple frogs to see how many it would eat. After 20 or 25 frogs (they were small) the snake’s skin was

24 Copyrighted Material Some pages are omitted from this book preview. Chapter 2 - Size and Shape stretched to the point where the bones in the frogs’ feet were visible through the skin. Snakeskin is remarkable tissue. Because large constrictors tend to eat large meals, their skin needs to be able to stretch and retain enough strength to keep the body from bursting open. Science likes experiments and data. It is the only way to know the secrets of the universe. newspaperman and herpetologist R. R. Mole (in Oliver 1958) measured a 3.11 m Boa Constrictor just after its death. He then skinned the snake and measured the skin. It was 3.71 m, an increase of 19.3%. Harvard herpetologist Arthur Loveridge (1931) repeated the experiment with an African Python. Loveridge wrote: With a view to obtaining data which might prove of assistance in esti- mating the actual length of a snake whose dried skin only is available, I measured one of these snakes in the flesh and found it to be 2180 mm, while its dried, and not unduly stretched, skin measured no less than 2650 mm. That is to say an increase of at least .21 of the total length should be allowed for, or in other words, a dried skin is nearly a quarter as long again as was the living reptile from which it was taken. The skull of this same snake measured 84 mm in its greatest length so that it may be assumed that a python is about twenty-six times longer than its skull, though this proportion varies with age for the larger snake measured 4330 mm in the flesh with a skull length of only 128 mm or a thirty-third of its total length. Herpetologist William Lamar killed a large Green Anaconda in Colombia in 1978 (in Murphy and Henderson, 1997). He carefully measured the snake’s dead body and found it to be 7.4 m. Aware that snake skins stretch, Bill skinned the snake with care to keep stretching to a minimum. After hours of work, the skin was removed and measured. It was now 13.5 m, a gain of 3.9 m or more than 40% of its original length. The maximum size of the Boa Constrictor Boa( constrictor) has been considered to be the Watkins-Colwell and Leenders (2003) record. William Duellman cited the 4.45 m Boa constrictor (2005), a record thought to be the maximum size for the boa. However, there is a problem. The measurement is based upon a skin, albeit a dried skin that was rehydrated. Skinning a snake always results in a stretched skin, and it seems unlikely that rehydrating causes it to shrink back to its original size, although to the best of our knowledge, there is no experimental data to support or deny this. It seems improbable that rehydrated skin shrinks back to the size of the snake. Instead, it is highly likely that it takes up water, becomes more flexible, and stretches even more. Our take on the 9.144 m-long skin reported by Moser and Robertson in 1998 and discussed at the start of this section is that it likely came from a snake that was about 7 m—possibly much less.

25 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes

Popular and Often-Repeated Stories That Are Misleading The Green Anaconda is considered to be the largest snake on the planet, while the Reticulated Python is usually considered to be the longest snake. However, the exact numbers are elusive. Here are some frequently repeated numbers and the stories behind them. During most of the last half of the 20th century, the longest snake was consid- ered to be the Anaconda, because of the Dunn-Lamon record. In 1944 Emmett Reid Dunn, a well-known and respected herpetologist published an article on the reptiles of Colombia in the journal Caldasia. It included a statement that his friend, Robert Lamon, a geologist working for the Richmond Oil Company had killed and measured an 11.5-meter Anaconda in eastern Colombia. Raymond Gilmore, of the U.S. Fish and Wildlife Service, investigated this record. In 1954 he found Robert Lamon working for the Northern Natural Gas Producing Company in Calgary, Alberta. In a letter dated May 19th, 1954 Lamon wrote to Gilmore and stated that he did kill and measure an anaconda on the Meta River. In that letter, Lamon wrote, I remember measuring the beast with a four-meter stadia rod, and if my memory serves me right, it required almost three lengths of the rod to obtain the dimensions, but I could not swear to this in that it may have been almost two lengths of the rod. However, this occurred sometime in about 1939 or 1940 just before I met Dix Dunn in Colombia. Therefore, the measurements must have been fresh in my mind, and if I so reported it to Dunn, I feel confident that the 11.5 meters is correct. In his unfinished manuscript, Gilmore wrote, The verdict on the anaconda of 11.5 m (37.5 ft.) is now up to the herpe- tologists. I think that I have shown that it is questionable and that the snake could have been 7.5 m or 24 feet (almost 2 lengths, 7.5 m, of a 4-meter stadia rod instead of almost 3 lengths, 11.5 m). The 11.5 m Anaconda was widely cited as proof that the longest snake was, in fact, the Green Anaconda. The record was quoted by Oliver (1958), Pope (1961), Minton and Minton (1973), and in numerous references, including several encyclo- pedias. Unfortunately, Gilmore’s letter went unpublished and unrecognized. After his death, the letter and unfinished manuscript were deposited in the archives of the SDNHM, where Gilmore had been Curator of Mammals. Van Wallach, then at Harvard University, obtained copies of the files and in the early 1990s sent them to JCM. In 1993, they were published in the Bulletin of the Chicago Herpetolog- ical Society.

26 Copyrighted Material Some pages are omitted from this book preview. Chapter 2 - Size and Shape

Rivas (2000) measured 780 specimens of the Green Anaconda in over seven years. The largest specimen he measured was slightly more than five meters. The largest weighed 97.5 kg, and the average of 45 specimens was 30.8 kg. Field measurements of snakes have also turned out to be less than reliable. Oliver (1958) and Pope (1961) both accepted them but consider the following. Clif- ford Pope’s (1961) classic text, The Giant Snakes, reports what Pope considered to be the record size for the Boa Constrictor (Boa constrictor). He wrote, The accepted record length for the boa constrictor is unusual in that it is based on a field measurement (by a scientist) that appreciably raises the formerly accepted maximum. This field measurement was made by Colin F. Pittendrigh, who encountered the big snake one morning in a swampy area of the Central Range of Trinidad. It was coiled up in the hollow end of a tree trunk, from which it had to be extracted by means of poles. After it was shot, Pittendrigh determined its length in the flesh as 18½ feet. This is the kind of field observation which cannot be lightly discredited. An 18.5-foot boa would be 5.64 meters. Oliver (1958) cited the same record as being the maximum sized Boa Constrictor. Sherman and Madge Minton (1973) also accepted this record in their book Giant Reptiles, as did John Mehrtens (1987) and Scott Weidensaul (1991) in their books on snakes. In the 1986 edition of Snakes of the World, Chris Mattison reported the Boa Constrictor to reach 6 m, a length not supported by any specimens. But in his Encyclopedia of Snakes (1996), he reported a maximum size of 4 meters. Hans E. A. Boos (1992), curator of the Emperor Valley Zoo in Trinidad, contacted Pittendrigh. Boos received a letter from Pittendrigh dated March 12, 1980. Pittendrigh was then Director of the Hopkins Marine Station operated by Stanford University. Pittendrigh described the 1944 encounter with the enormous snake in the Guico-Tamana area of Trinidad. The snake was basking on a fallen tree when it was discovered, killed, and skinned. The skin was said to be 30 feet long (9.1 m), but the animal was estimated at 18 feet long (5.5 m). That night the skin was destroyed by stray . Boos pressed Pittendrigh for more information and the possible confusion between the Boa Constrictor (Boa constrictor) and the Green Anaconda (Eunectes murinus). In a second letter, Pittendrigh admitted that he did not know the difference between the Anaconda and the Boa Constrictor. Boos later contacted Yussuf Khan, a local man who had worked with Pittendrigh and asked him about the big snake. Khan said it was large, green, and that it was a “Huille,” the local name for the Anaconda. Thus, the record-sized, 18-foot (5.49 m) Boa Constrictor was almost certainly a misidentified Green Anaconda. The Reticulated Python named Colossus, which was maintained in captivity at Pittsburg’s Highland Park Zoo, was considered to be the longest snake ever kept in a 27 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes zoo by the Guinness World Records Limited (Wood 1972). Murphy and Henderson (1997) agreed with this assessment, and it appeared the snake measured 11.2 m on 15 November 1956 as reported in Barton and Allen (1961). However, Barker et al. (2012) inquired at the Carnegie Museum, where Colos- sus’s remains are housed, the snake was skinned, and the hide and bones were kept. The collection manager found the data collected on Colossus at the time he was prepared. Barker et al. wrote, “Colossus truly was an immense, big-bodied, giant snake, but he was 6.35 m (20 ft. 10 in) in total length, exceptionally long for a male python, but nowhere near as long as was generally reported.” Measuring a giant snake, even when it is in captivity and the workforce and tools available are excellent, is difficult. An accurate measurement is at best ambiguous. There is another issue with the Colossus record that was not directly addressed by Barker et al. (2012). It has nagged us for some time: Colossus was a male, a male that reached 6.35 m in captivity. Also, at one point Colossus weighed “more than 200 pounds.” This is more than 90.7 kg. This same snake was attributed as weighing 133.7 kg. The length and weight could be the result of it being a well-fed captive. Or, more likely, Colossus may have been a female, and misidentified as a male. Colossus does not fit the profile of the well-studied Sumatran Reticulated Python population regarding length or weight. However, its origin was thought to be . Colossus simply should not have gotten this big unless he was a she, or unless it represents a different species of Reticulated Python with different sexual dimorphism. TheGuinness Book of World Records website describes a 7.67 m, 158.8 kg Retic- ulated Python named Medusa that was owned by Full Moon Productions, Kansas City, Missouri in 2011. The Reticulated Python named Twinkie at the Reptile Zoo in Fountain Valley, California was reported to weigh 158 kg and was more than 6.1 m (Virata, 2014). The Reticulated Python named Fluffy at the Columbus Zoo and Aquarium was said to be 7.3 m and weigh 136 kg. It died in October 2010. The longest recently-documented snake seems to be the Reticulated Python named Samantha, a resident of the for almost a decade. She was hatched in in about 1970. The snake was captured and held in a boxcar in Samarinda, Borneo. The large python came to the attention of the Bronx Zoo via Paul Raddatz, a leather dealer in Wisconsin. When the snake arrived at the zoo in 1993, it was 6.4 m long and 80 kg and eventually grew to 8 m and 125 kg). Bill Holmstrom, the supervisor of the Bronx Zoo’s World of Reptiles, considered Samantha a mellow and easy-going creature (Santroranov, 2012). Fragrant Flower was a twenty-first century giant that shrinks. Just because a snake is in captivity does not mean its size is safe from exaggeration. John Aglionby 28 Copyrighted Material Some pages are omitted from this book preview. Chapter 2 - Size and Shape

(2004), a reporter for The Guardian newspaper investigated the story of a 14.85 m, 447 kg python that was said to be 150 years old. The specimen was living in a tourist park in the village of Curug Sewu, about 40 miles south of Semarang in central . People paid 2,500 rupiahs to view the giant snake. Darmanto owned and kept Fragrant Flower. He initially claimed the snake came from Sumatra, while a tourism officer said it came from a locality in Java. The Reticulated Python lived in a 9.5 x 4.5 m cage of corrugated iron and was fed several dogs each month. Aglionby placed a tape measure on the giant snake and found that it fell far short of the 14.85 m length reported by the park. He measured the animal at 6.5 to 7 m and estimated its weight to be a maximum of 100 kg. When Darmanto was queried about the actual size of Fragrant Flower, he said, …you must understand that a python’s length is not constant… Depending on the weather, on how recently he has eaten, and when he last shed his skin, Fragrant can stretch and contract a great deal. A few days ago he stretched himself out halfway round the cage. The non-scientific mind can always supply an answer. A report of a 10-10.2 m Reticulated Python from Sulawesi (formerly known as the Celebes) is frequently cited as the record-sized snake. The origin of this story can be traced to an article written by Harry C. Raven, a museum collector who traveled the between 1912 and 1923. The published report comes from a 1946 article in Natural History Magazine about an incident that happened in 1912. Raven wrote: I left the schooner and went inland a short distance to camp on the mountains, which were covered with virgin jungle. The white men at the mine told me of a huge python one of their relatives had killed a few days before my arrival and showed me a very poor photograph of it taken after it had been killed and dragged to camp. Though the print was dull, you could see a man standing on the huge body, which was about a foot thick. The civil engineer told me it was just ten meters long. I asked him if he had paced off its length, but he said no, he had measured it with a surveying tape. There are reasons to seriously doubt Raven’s story. It has all the classic signs of fabrication or a seriously exaggerated snake story. The snake was killed just before he arrived, and the photo was poor in quality. Also, exactly how do you get a photograph developed in a few days in 1912 Sulawesi? Polaroid or digital camera technology was not available. A 10 m snake is likely to be much more than 0.3 m in diameter. Additionally, Raven goes on to tell about another big snake that was captured by tribal people while he was there, but they ate it! Alas, he had no proof. The Raven story is not acceptable documentation for a 10 m Reticulated Python. There is a similar discussion of this story in Murphy and Henderson (1997) and on 29 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes the Internet. It is a record that is still cited as being the largest documented snake. Old fables die hard. While working on this book, we came across a photo published in a British newspaper in 1975. The photo was reportedly taken in the Celebes in 1898 (Figure 2-2).

Figure 2-2. An old newspaper photo of three large python skins from Sulawesi. We suspect the 10-meter Sulawesi python story is based on this photograph. It was published in an undetermined British newspaper in 1975. However, the photo was reportedly taken in 1898 in the Celebes (now Sulawesi). A second copy of the same photo noted that these skins were 28.5, 26.5, and 24.5 feet respectively. The photograph was submitted by Patrick J. Brett, Heather Close, Cedar Rd., Hook Heath, , Surrey. Keeping in mind that snake skins always stretch and assuming that the lengths of the skins are correct, reducing the size of each by a conservative 20% will provide a liberal estimate of the actual size of the animal they came from. Courtesy of Breck Bartholomew. Rewards for a Giant Snake

Carl Hagenbeck (1910) may have been the first person to offer a reward for a super-sized snake. He became involved when one traveler asked if anyone would pay £500 for a snake that was 12.2 meters long. Hagenbeck had been in the animal business for some time, and the largest snake he had seen was 7.92 m. Therefore, he thought he was justified in engaging in the reward offer. He and his friends offered £1000 for a 9.1 m snake. To claim the reward, the snake had to be delivered to Hamburg, Germany alive and healthy. From about 1910 until 2002 the New York Zoological Society (The Wildlife Conservation Society) continued to offer a steadily increasing reward for any healthy, live snake 9.1 m or more in length. The only potential candidate species would likely be the Reticulated Python and the Green Anaconda. Individuals have tried to grow captive snakes to meet the size needed to claim the reward, but that approach has also been unsuccessful. The reward started at $1,000. It was subsequently raised to $5,000. In January 1978 it rose to $15,000; then at the 1980 Members Meeting of the NYZS, William G. Conway announced that the reward had been raised to $50,000. The reward was canceled in 2002, most likely to discourage people from disturbing large snakes in the wild. 30 Copyrighted Material Some pages are omitted from this book preview. Chapter 2 - Size and Shape

Our Perspectives on Maximum Size There is one squamate that shrinks when food is not available—the Marine , Amblyrhynchus cristatus (Wikelski and Thom 2000). However, there is no evidence that snakes shrink. Luiselli (2005) tested data he had on the Australian Water Python and about 16 species of African snakes that he had captured and recaptured. With each capture and recapture, he had collected data on length. He found no evidence that snakes do what the Marine Iguana does. However, of 6,431 recaptures, 2.8% showed shrinkage, and 14 out of 16 species examined had at least a single case of shrinkage. (The percentage of shrinkage cases was not significantly different among terrestrial, semi-aquatic, and arboreal species.) However, the percentage of shrinkage cases varied considerably in relation to the “measurability” score of single species. Luiselli found an exponential growth curve for the relationship between the two variables: the mean body length and the ease of measurability. Large, vigorous snakes were more difficult to measure correctly than small non-vigorous species. Measurability was influenced by temperament and physical danger, i.e., defensive species were more difficult to measure accurately than placid species, and highly venomous snakes were more difficult to measure than non-venomous species. Loss of body size at the second capture event compared to the first (in terms of percentage) was trivial compared to shrinkage data for marine . Both the frequency of occurrence of shrinkage events and the percent of the decrease in the body length did not differ interseasonally. Luiselli concluded that shrinkage events are caused by measurement errors and are more frequent among large and dangerous species (i.e., large boas and pythons, cobras, mambas, and large vipers). I (TC) have had a very large Reticulated Python that has been kept outside in Homestead, Florida. She was caught wild near Miami in 2012 as a three-meter escaped or released pet. At one point I had friends present who were capable of dealing with a giant snake, so I attempted to get an accurate measurement. Going to great lengths to ensure accuracy, we measured her at 5.56 m long in a tight coil. She had been measured before, and the measurements were always near 6.1 m. Measuring a coiled snake is not the same as measuring a snake stretched out. It is almost impossible to get a 100% reliable measurement on giant snakes unless they are dead. The length can change dramatically depending on compression or expansion of the vertebra. The vertebral column has a degree of flexibility in all vertebrates. Humans usually have 33 vertebrae, with about 10 of them being fused in groups, leaving about 23 that articulate. Each one is separated from its neighbors by a disk of cartilage. Therefore, humans have some ability to stretch. With age, humans tend to shrink as the disks and bones degrade. The giant snakes we are discussing here have 250–300 articulating vertebrae. If each joint (one at each end of the vertebrae) has just two millimeters of stretching potential in an adult giant snake, that is a very conserva-

31 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes tive 100–120 cm stretch. Thus, any large snake can produce multiple measurements depending how much its vertebral column and body has stretched or contracted. This suggests to us that any measurement of a live snake’s length is likely to have a significant margin of error.

Scrub Python, Simalia kinghorni

This giant snake has always been underrated because its mass/length ratio is low compared to other giant snakes. Scrub Pythons likely reach or exceed six meters in length. Lumholtz (1889) reported an adult length of 6.1 m. Barrett (1950) at Brown’s Bay Zoo in Cairns described a 5.94 m specimen and another 6.71 m spec- imen. McPhee (1959, 1966) considered the average size to be 4.26 m, an authenti- cated maximum length of 7.01 m, and an alleged maximum of 8.53 m. However, he provides no specifics. Worrell (1963:97) discussed a specimen killed at Greenhill (near Cairns) and measured by L. Robichaux at 8.53 m and considered specimens over 4.57 m uncommon. Dean (in Pope, 1967) also mentions a 7.23 m specimen. See Table 2-2 for a summary of sizes reported in the literature. Fearn and Sambono (2000a) summarized the size records for this species from the literature. Maximum sizes ranged from 8.5 meters (Worrell, 1963) to 4.88 m (Anonymous, 1966). However, they obtained a specimen from Cairns, with a body length of 4.9 m and a 0.751 m tail, for a total length of 5.651 m and a weight of 24 kg. Murphy and Henderson (1997) considered this snake to be a near-giant. However, there is some evidence to suggest that the species does, in fact, exceed 6.1 m. Bick- ford (2004) described a specimen that he collected from Chimbu Province, Papua New Guinea. The SVL (Snout to Vent Length) was about 5.9 m, and he measured it twice. Based on records and photos, we must accept that this python may reach or exceed seven meters.

Burmese Python, Python bivittatus

The maximum size of this species is plagued with the same inaccuracies as other giant snake records: mismeasurements, overestimations, and misinterpretations. Reed and Rodda (2009) followed Ernst and Zug (1996) and Murphy and Henderson (1997) in acknowledging that specimens over five meters are rare, but state that Bellosa (2003) reported an 8.22 m, 182.8 kg specimen in Gurnee, Illinois in 1998. The snake was named “Baby.” This specimen is responsible for most of the reports of Python bivittatus exceeding 6.1 m, including the Guinness World Records published between 2003 and 2006. The snake had been weighed on Fox TV at 182.8 kg. Her length was measured using a cloth tape run down her back, and she was measured at 8.22 m. However, upon her death, she was measured with a steel tape measure in a lab and found to be 5.74 m. See Barker et al. (2012) for more details on the shrinking record.

32 Copyrighted Material Some pages are omitted from this book preview. Chapter 2 - Size and Shape

Table 2-2 Scrub Python, Simalia kinghorni Length (m) Reference 5.5 Skardon, 1938 7.01 Thompson, 1949 5.49 Barrett, 1950 (Brown’s Bay Zoo, Cairns) 6.71 Barrett, 1950 (photo from S. A. Newton) 6.1 Worrell, 1951 8.5 Worrell, 1954 7.225 Dean, 1954 6.4 Kinghorn, 1956 7.6 Worrell, 1958 5,18 Worrell, 1963 4.88 Williams, 1966 7.01 McPhee, 1966 5.5 Frauca, 1973 5.5-6.0 Parker, 1982 5.18 Firth & Firth, 1991 5.5 Fearn, 1998 5.6 Fearn and Sambono, 2000 Table 2-2 The size records for the Scrub Python, Simalia kinghorni. Much of this data is based upon Fearn and Sambono, 2000a. Also, see Bickford (2004).

Table 2-3 Summary of reported sizes for the Burmese Python, Python bivittatus. Length (m) Weight (kg) Location Reference 5.74 captive Barker et al., 2012 4.73 52.2 Chitwan National Park, Dhungel, 1983 5.56 United Provinces Prince in Wall, 1921:68 5.18 Fife-Cookson, 1887 6.4-6.7 Khasyia Hills Forsyth, 1911 5.84 Cooch Behar, in Wall, 1911:452 5.18 Darjeeling Wrenicke, 1955:134.

Records of Python bivittatus that are 5.18 m or more are well represented both in captive and wild snakes, with rare records exceeding 5.48 m in length. Based on seeing many highly esteemed and valued captive wild-caught giant Burmese, I (TC) has never seen even one that was 5.4 m long. Most were 4.87–5.18 m long. Cynthia, a large Burmese I once had, grew to just under 5.18 m after I had her for 20 years. A friend had her for the decade before. He brought her home with him from . Captive Burmese grow very rapidly to about 3.64–4.26 m. The rate of growth after that dramatically slows. In July 2015 a Burmese Python was captured in Shark Valley, ; it was found to be 5.56 m and weigh 60 kg. Table 2-3 summarizes the other size records for this species. We do not believe that P. bivittatus reach much over 6.1 m in length. 33 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes

Indian Python, Python molurus molurus

Because of the confusion between this snake and Python bivittatus, the size of P. molurus has been greatly exaggerated by some authors. It is possible, but unlikely, that it exceeds 5 m. Table 2-4 shows sizes reported for this snake. Table 2-4 Summary of reported sizes for the Indian Python, Python m. molurus. Length (m) Location Reference 2.88 Acharjyo and Mirsa, 1975 4.35 Acharjyo and Mirsa, 1975 7 Deoras, 1965 4.57 Pakistan Minton, 1966:117 5.48 Ashambu Hills, India Ferguson, 10:69 5.56 United Provinces Prince in Wall, 1921:68 5.18 Fife-Cookson, 1887 5.79 , India Jerdon, in Wall, 1911 5.8 Assam Cooch Behar, in Wall, 1911 5.18 Darjeeling Wrenicke, 1955:134.

Ceylonese Python, Python m. pimbura

Relatively little is known about the size of this snake. Table 2-5 summarizes the literature records for the largest specimens. Note that these are mostly old records reported in books about traveling in Ceylon (now ). We believe that Ceylonese Pythons attain about the same size as Indian Pythons or slightly smaller. It is doubtful they exceed 5 meters long. Table 2-5 Sizes reported for the Ceylonese Python, Python m. pimbura. Length (m) Reference 5.79 Tennent, 1861:153 5.18 Tennent, 1861:304 5.79 Jerdon, 1853 6.7 Percival, 1805:311.

South African Python, Python natalensis

The long-standing confusion between Python sebae and Python natalensis makes it difficult to sort out the tangle of comments in the literature on these two species. Python natalensis was not recognized as a valid species until Broadley (1999) sepa- rated the two . Clarence Abercrombie told us that he has seen 6-meter and possibly 7-meter individuals of Python natalensis in Zimbabwe in recent times. 34 Copyrighted Material Some pages are omitted from this book preview. Chapter 2 - Size and Shape

North African Rock Python, Python sebae

Specimens near six meters were said to be common in (Chirio and LeBreton 2007). According to Pope (1961), the greatest, seemingly trustworthy total length known was 9.81 m, based upon a specimen found in a garden hedge at Bingerville (05°21’N 03°54W, ). The python was measured by Béart in 1932 according to Villiers (1950, 1975). The snake was also discussed and accepted by Minton and Minton (1973). However, Branch (1984) and Spawls et al. (2002) agreed that this record was anecdotal and without any tangible evidence to support it. A slightly more credible record of a 7.5 m specimen from a research station in the Ivory Coast, was discussed by Doucet (1963) but again, the evidence is absent. Starin and Burghardt (1992) reported that in Gambia 52 python sightings were made on 45 different occasions. The pythons ranged in size from 0.5 m to about 7.5 m in total length. Based on 50 sightings, the median and modal total length was 3 m with a mean of 3.3 m (SD = 1.3); 20% of the sightings were of animals greater than 4.5 m. However, there is considerable variation in weight according to condition and sex, the females being proportionately heavier than the males. See Table 2-6 for a summary of published size records. The North African Rock Python likely rivals the Green Anaconda and possibly the Reticulated Python for length. The largest one TC had was a female that measured at 5.33 m and weighed 70.3 kg but was not fat. We don’t completely accept the old record from Bingerville, West Africa, of a 9.93 m Python sebae killed in a schoolyard because of the lack of documentation. However, maximum size in the 7.5–8 m range is likely given the Starin and Burghardt (1993) paper. Of course, this is our guesswork, and we could always be wrong. There is a difference in size between the Northern and Southern African Rock Pythons. Specific localities of both may produce larger or smaller snakes. The West Africa P. sebae seem to be much larger.

Table 2-6 Size reported for the North African Rock Python Python sebae. Length (m) Reference 9.8 Beart, 1932 in Doucet, 1961 7.5 Doucet, 1963:227 9.45 Duncan, 1847:157 12.2 Kingsley, 1897:546 7.3 Lonnberg, (1911) 7.5 Starin and Burghardt, 1993:53 7.85 Stucki-Stirn, 1979:50 4.93 Tjader, (1910:233) 11.0 Van Rompaey, (1985:250)

35 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes

Reticulated Python, Malayopython reticulatus

The Reticulated Python probably reaches a maximum length of 8 to 8.5 meters. Our reason for this estimate is that the oft-cited maximum of 10 meters has not been docu- mented. Measuring a snake that size, even if the snake cooperates, is difficult. TC has seen one living Reticulated Python in the [near Jakarta] that was said to have been collected in Sumatra. The owner claimed it was nine meters long. It was more like 7–7.5 meters. TC offered the owner $5,000 for it in 1988 but was turned down. $5,000 was a princely sum in Indonesia. My thought was that snakes of this size were very rare, even in 1988. Writers discussing the Reticulated Python always want to round up, not round down, on the size reported. There is a story of a giant python from Kedah, that has not received much attention. We note it here because it is told in a calm and straightforward way, but it ends with lost evidence, which is typical of many exceptionally-sized snake reports. Boris Hembry (2011) was a German rubber planter in Malaysia prior to World War II. In 1931 Hembry was eating breakfast when his workers reported a large snake nearby. He wrote, “…it was a good 12 inches across, and when we

Table 2-7 Sizes reported for the Reticulated Python, Malayopython reticulatus. Length (m) Weight (kg) Reference 6.90 Anon., (1903:149) 7.31 Anon., (1930:89) 6.01 Flower, (1899:655) 7.8 112.4 Hagenbeck, (1910:189) 8.83 Cross in Mayer (1920:845) 8.68 145.2 Barton and Allen 10.05 Raven, (1946:38-39) see discussion 6.01-7.92 Ridley, (1899:196) 8.22-8.53 Smith, (1943:110) 7.56 114.3 Barker and Barker, (1997:53) 7.26 90.7 Barker and Barker, (1997:53) 9.14 Cantor, 1847:57 6.95 26.7 Fredriksson, 2005 113.2 Hagenbeck, (1910:189) 7.62 138.3 Pope, (1961:163) 124.7 Anon. Reuters, Nov 24, 2002 170.1 Worcester, (1898:115) () 9.0 Sody, 1941 6.55 90.7 Ussher (1979:180) 8.2 Wray in Flower, 1899:655

36 Copyrighted Material Some pages are omitted from this book preview. Chapter 2 - Size and Shape eventually straightened it out, it measured 33 feet 6 inches.” [10.24 m]. The laborers had injured the snake, and they moved it to the plantation owner’s house. It took ten men to carry it. A local hunter was brought in to kill the snake. Hembry describes taking photographs, all of which were lost. The snake was skinned, and the plan- tation owner’s wife had the skin. Hembry suggests that the skin may have been brought back to Zurich. Maximum-sized Reticulated Pythons are likely in the 8–8.5 meters range. In discussing this python, authors often tend to follow others in their acceptance of maximum-record Reticulated Pythons. These do not hold up under scrutiny. The maximum size is likely 8-9 meters.

Green Anaconda, Eunectes murinus

The Green Anaconda has been the subject of remarkably imaginative specula- tion. Based on a Green Anaconda reliably measured by Bill Lamar, we suspect 7.62 meters as the maximum reported size for Eunectes. It has been TC’s experience that the largest Anacondas seem to come from gallery forest in tropical savannah and ox-bow lakes. All of his earlier imports of large anacondas came from these types of locales. TC always looked at old pictures when visiting places with giant snakes. Even the old pictures and stories of the biggest anacondas depicted them in these two major habitat types. We think that anacondas may indeed get larger in these areas, as the savannah offers a bounty of large endotherms like deer and . Additionally, the ox-bow lakes usually have the heaviest concentrations of caiman of all size classes. It seems likely that the largest anacondas come from such habitats because of an abundant food supply. Table 2-8 Size reported for the Green Anaconda, Eunectes murinus. Length (m) Reference 14.0 Amaral, (1948:237) 9.14 Bancroft, (1769:203) 9.15-12.19 Barbour, (1926) 8.84 Beebe, (1946) 7.01 Blomberg, (1956:97) 7.62 Ditmars, (1931:43) 7.31 Fountain, (1904:107) 11.5 Lamon, (in Dunn, 1944) * 10.25 Medem, (in Oliver, 1958) 9.75 Mole, (1924:23) 11.28 Quelch, (1898:297) 10.36 Roth, (in Oliver, 1958) 11.58 Up de Graff, (1923:71)

37 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes

A snake’s size is determined by its ancestry and its development. Different parts of the body grow at different rates, producing the variation we see in body proportions. At the same time, the timing of when an organ or section of an animal’s body starts to grow (heterochrony) can have a dramatic impact on what an organism looks like. We review the criteria for determining longest snakes and the problems associated with obtaining accurate measurements. A whole snake can be measured several times with different results. Large snakes and snakes that are uncooperative are more difficult to measure accurately. Measurements of snakeskins are overestimates of the actual size of the snake by at least 20%. Snakes don’t get as big as popular culture may lead us to believe.

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38 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes radiation in two primary wavelengths, one of which (8–12 μm) matches the infrared emission of targeted prey (Grace et al. 1999). Goris (2011) went farther and suggests the infrared organs of boas and pythons, as well as pit vipers, need to be thought of as eyes. They function not by a photo- chemical reaction as light-receiving eyes, but by heat generated in the receptors or the terminal nerve masses (TNMs) by electromagnetic radiation. Pythons and pit vipers have the TNMs located in a pit, and the pit opening acts like an aperture of a pinhole camera, a virtual lens. This allows the snake to focus the heat and permits the receptors to encode the movements of an infrared source sufficiently for the brain to form an image. Many booid snakes possess TNMs identical to those of the pythons, but they lack an opening that could serve as a lens. All TNMs are irrigated by a dense network of capillaries that act as a heat regulator and mimic the role of the photochemical cycle in eyes. Thus, the pits are an integral part of the snakes’ visual system, which makes use of the longer waves of the electromagnetic spectrum for which there are no appropriate photoreceptive pigments in nature; they do everything the eyes do. They are often treated as a sixth sense that is used only for the detection and acquisition of prey, but this idea is misleading. Boas, pythons, and pit vipers see the world using both the visual and the infrared spectra.

Figure 3-2. Infrared sensing organs have evolved in the booids, pythoninds, and pit vipers. Booids and pythonids have the pits located in the labial scales, while pit vipers have the pit in the loreal region of the face. The pits collect heat (infrared energy) that produces an image in the snake’s brain. The tongue collects molecules that are analyzed by the vomeronasal system (VNS), and the nares also collect chemical information for the olfactory system. The snake is the Python, Malayopython timoriensis. JCM

50 Copyrighted Material Some pages are omitted from this book preview. Chapter 3 - Snake Origins & Biology

Active Foraging and Ambush Snakes are frequently classified as foragers or sit-and-wait hunters. These two behaviors are at opposite ends of a continuum. Snakes may alternate methods or use variations of these hunting methods, depending on the habitat, kind of prey, the thermal environment, their reproductive status, and the snake’s exposure to preda- tion risk. Species hunting from ambush tend to be sedentary, infrequent feeders; they hunt for extended periods of time, move relatively short distances, have stout bodies, grow and mature slowly, females have relatively smaller litters. They tend to be cryptic in their coloration and behavior. Active foraging species tend to be active, hunt for relatively short time periods, have a high metabolic rate, grow fast, and have smaller litters. Their body is slender, and they have a high risk of predation. This is a simplistic dichotomy, and the giant snakes don’t fit neatly into one or the other. This is likely due to the generalized nature of their diet and habitats. While all of them capture prey from ambush, they will also wander over the landscape in search of food. This flexible hunting behavior is one of the attributes that makes them successful. A term we have coined for this is “ambush foraging.” A giant snake may sense prey that is too far away to ambush, so the snake moves forward taking advan- tage of every shadow or prop to get close enough to attack, unseen by the prey. One of the authors (TC) has seen this routinely in captive giants. In photos of giant snakes catching prey in seemingly open areas, we believe this method is used frequently. Food and Feeding Snakes are obligate carnivores. Their jaws, teeth, and musculature are adapted for swallowing whole prey. (Two known exceptions tear their soft-shelled crustacean prey into pieces.) Some snakes are known to have specialized for particular prey: fish eggs, reptile eggs, nestling birds, worms, or geckos. Others are generalists, preying upon any animal they can overpower. All of the giant snakes fall into the latter category. The strike of a snake may differ with the situation. A feeding strike usually occurs when the snake opens its mouth during the strike and embeds its teeth into the prey, immediately following the strike by throwing one or more coils of its body around the prey. Defensive strikes may occur with the mouth closed or partially closed and are not followed by constriction. The snake may, however, deliver a warning bite where the snake quickly bites and releases the prey. Snake evolution has taken several pathways when it comes to killing prey. Many snakes have toxic molecules in venom for subduing and killing prey. The venom may be delivered in several ways, ranging from a highly efficient enclosed fang that trans- ports venom under high pressure into the prey, to open grooved fangs that provide venom under low pressure, to venom apparently seeping into a bite wound with no specialized fangs at all. Other clades have evolved behavior that involves pressure

51 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes from the snake’s body to hold the prey down and swallowing the prey alive. Yet other snakes use constriction, wrapping one or more coils of their body around the prey and tightening the coils until blood flow to the heart stops, and the prey dies. The force exerted by constriction is directly related to the diameter of the snake’s body. Snakes with the most significant body diameter apply the most force (Penning et al. 2015). Frank Wall (1912) may have been the first to propose that death by constriction results from circulatory arrest. Constricting snakes are able to exert enough force on an animal’s body to prevent the return of venous blood to the heart. Others had proposed that death from constriction resulted from suffocation, but death by suffo- cation takes longer and might increase the chances that the prey will injure the snake. Circulatory arrest can result in rapid heart failure and death. It is also possible that constriction can result in spinal fractures that kill the prey quickly and probably reduce the risk of injury to the snake. However, when constricted prey is examined for bone damage, broken bones are rarely found. The amount of force exerted by constricting snakes was measured in 12 species by Moon and Metha (2007). Not surprisingly, larger snakes exerted more force than smaller snakes and used a force directly related to the diameter of the trunk and the number of loops. They proposed that snakes that exert a high amount of force prob- ably kill prey by a spinal fracture and by dislocation of vertebral joints. Snakes that exert more moderate forces may kill prey using circulatory arrest, and small snakes that exert less force may kill prey with suffocation. Of interest is that the snake doing the constriction may also reduce blood flow in its own muscles while constricting prey. Snakes have low arterial blood pressure but are tolerant of anoxia. They can use anaerobic metabolism during constriction, so the anoxia does not result in adverse effects. They predict that large constricting boas and pythons may be able to exert forces that approach 900kPa (= about 130.5 pounds/square inch). Snakes use their epaxial muscles (muscles that are dorsal to the transverse processes of the vertebrae) for constriction, and Lourdais et al. (2005) examined these muscles in the Columbian Rainbow Boa (Epicrates maurus) using MRI (magnetic resonance imaging) and caliper measurements. They measured the epaxial muscu- lature in snakes, their performance in escaping a predator, and their ability to handle prey. Snakes were fed after a fasting period and had their musculature and behavior measured. They found epaxial muscles were a determinant of strength intensity. Snakes were fed for six months after the fast and showed an increase in the width of the epaxial muscles along the length of the body; muscle gain was greatest in the posterior body. During a fast, this region lost the most protein to fuel metabolism. Penning et al. (2015) studied constriction performance in Malayopython reticu- latus and Python bivittatus and propose that prey death by constriction occurs by a mechanism other than suffocation. In both species, constriction pressure increased significantly with snake diameter. Constriction exerts forces dramatically higher than

52 Copyrighted Material Some pages are omitted from this book preview. Chapter 3 - Snake Origins & Biology their prey’s blood pressure, stops blood from circulating, and results in rapid prey death. The snake likely stops blood flow to the heart and may disrupt brain function. All of the boas and pythons employ constriction to kill their prey. Contrary to popular opinion, constriction does not usually break bones. Once the prey stops moving the snake lessens its coils and starts swallowing the prey’s head, working its jaws around the prey and slowly pulling it into its esophagus. Conventional wisdom has been that smaller snakes tend to eat smaller prey while larger snakes tend to eat larger prey. The correlation between the body size of a snake and the body size of its prey could result from a variety of factors. Indeed, encounter rates would be one of them. Small prey would typically be expected to be more abun- dant than large prey, yet some large snakes seem to ignore small prey completely. The snake may have a specific preference for a particular of prey that would result in this phenomenon. Or the snake may choose prey based upon its ability to capture, subdue, and handle it. Giant snakes can swallow prey that is equal to or greater than their body weight. Eating such prodigious meals presents several challenges, including the following: digesting the prey before it decomposes in its digestive system, escaping a predator after it has consumed a large meal, and preparing its digestive system to deal with substantial meals. The snake must have a secluded shelter with a temperature gradient favorable to digestion. During digestion, they are unable to defend themselves. The extra weight of the prey prevents their moving away quickly. This is why snakes regurgitate food when disturbed or captured. While a ratsnake throwing up a may not take much time or effort, a giant python that has eaten a deer (sometimes with antlers) and weighs 41 kg faces a dilemma if it is attacked by a predator. Giant Reticulated Pythons are often associated with cave systems. We believe this preference is to protect themselves from predators and to achieve the body temperatures they require to facilitate digestion. Secor and Diamond (1997) investigated the adaptive responses of juvenile Burmese Pythons to meal sizes. They found larger meals took longer to pass through the gut; oxygen consumption rates rose to up to 32 times fasting levels and remained significantly elevated for up to 13 days. This required the snake to expend 29-36% of the energy they ingested. They also found dramatic changes in blood chemistry. Within a day after ingestion, the intestinal mucosal mass more than doubled, and the masses of the intestinal serosa, , stomach, pancreas, and kidneys also increased. Intestinal absorption rates of amino acids and of D-glucose increased by up to 43 times fasting levels, whereas uptake capacities increased by up to 59 times fasting levels. The magnitudes of many of these responses increased with meal size, while other responses plateaued at meals equal to 25% of the snake’s body mass. Pythons undergo a wide array of responses after eating, many of which differ in degree to the

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Figure 3-3. A South African Python, Python natalensis, constricting and swallowing an Impala. Snakes have the remarkable ability stretch their jaws around their prey and pull the food into their gut using their teeth. Photo by Tal Feinberg. 54 Copyrighted Material Some pages are omitted from this book preview. Chapter 3 - Snake Origins & Biology meal size. This research leads to many more projects to understand how pythons modify their physiological response to large meals. The molecular basis for extreme morphological and physiological adaptations to large meals in snakes was studied by Castoe et al. (2013). They sequenced the genome of the Burmese Python and discovered large numbers of rapidly-evolved genes that are linked to extreme characteristics in snakes. Traits like a rapid increase in metabolism and organ growth after feeding allow the snake’s digestive system to handle the large prey. The researchers also found that after Burmese Pythons ate, they experienced massive changes in gene expression linked to 35–100 percent size increases in their heart, small intestine, liver, and kidneys in just 24–48 hours. Snake metabolism is among the lowest of any vertebrate, but it is ramped up significantly after eating. In 2015, a group of biologists found a 3.35 m, 14.2 kg Burmese Python in Florida’s Collier Seminole State Park. It had swallowed a white-tailed deer fawn weighing 15.8 kg (Bartoszek et al. 2018). The report suggests that this is the largest python-to- prey ratio documented to date. However, Shine et al. (2012) reported prey that was between 1.1 and 1.4 times the mass of the Reticulated Pythons that had consumed them. Before that, Lederer (1944) had described a female Reticulated Python that was 7.5 m and weighed 47.5 kg. This snake ate a that weighed 54.5 kg. (The predator to prey ratio was 1.147.) To the best of our knowledge, this was the first documented evidence that a python could consume prey greater than its mass. Ecdysis Snakes periodically shed their skin in a process termed ecdysis, a trait shared with other squamates. The events that take place prior to and during ecdysis are complex. A mature epidermis contains a keratinized layer of β-keratin composed of multiple layers of cells. A second layer below this one is composed of multiple layers of keratin between lipid material above and below the second layer. Next is a layer of α-keratin where the older keratin interfaces with a new α-keratin layer that is maturing. All of these cells are produced by the lower stratum germina- tivum found just above the dermis. Before shedding, the lower β-keratin becomes mature, and the two layers separate. The shed is composed of a β-keratin and an α-keratin layer. Ecdysis may be triggered by growth, damage to the skin, ecotoparasites, or some other factors. Given that a snakeskin removed from a dead snake can stretch dramatically (at least 20%), and that snakeskin stretches dramatically when eating large prey, ecdysis is a poorly understood phenomenon in snakes. Any injury will prompt unusual numbers of shed cycles even without feeding as a means to heal. TC feeds his captive pythons seven to nine times per year, and each feeding seems to throw them into a shed cycle. The size of the prey does not seem to matter 55 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes in terms of triggering ecdysis. This leads us to believe that prey selection is critical to large pythons. Albeit being an opportunistic apex predator, it would seem unlikely a python would pass up a small meal. Since food is directly related to the size they can attain prey, selection may be more important than we now recognize. This applies more to older, larger snakes than to young ones that are continually shedding when fed. Neonate Reticulated Pythons will shed 12–16 times in their first year if they are “power fed” (a term that refers to frequent feeding for maximum growth). Most adult giant snakes will not feed when in a shed cycle, although neonate giant snakes frequently will eat in ecdysis. Power feeding of any giant snake is done to get the maximum size in the shortest amount of time. This is done to facilitate the ability to breed sooner and to produce more eggs, as the female python is bigger than average for its age. Power feeding seems to drastically shorten the lifespan of these giant snakes due to renal and hepatic medical issues. Diel Color Change Diel color change is well known in squamates (Boback and Siefferman 2010). All boas and pythons undergo a physiological color change, including the giant snakes. These sometimes dramatic changes are used to increase their cryptic appearance from day to night and related to enhancing thermoregulatory abilities. Captive boas and pythons undergo some extraordinary color changes in a very short time. The giant snake that undergoes the greatest color change is the Reticulated Python. They Day Night

Snake A Snake B Snake A Snake B Figure 3-4. A comparison of the day and night coloration in two Candoia bibroni. Photos by Jay Wagner.

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Chapter Four Pythonids – An Overview

73 Copyrighted Material Some pages are omitted from this book preview. Chapter 4 - Pythonids – An Overview

What makes a python a python? Using morphology to answer this ques- tion would result in a long list of anatomical traits that most readers would find unfamiliar and confusing. It could read something like this partial statement from Underwood’s (1967:169) Classification of Snakes: “Prefrontals approach one another at midline. Movable articulation between snout and braincase. Dorsal end of postor- bital bilobed. Levator anguli oris muscle lost. Body of pancreas lobed.” Instead, the simpler, cleaner answer and one that can be tested is simply—ancestry. All pythons share a common ancestor. The 18th century naturalist (a.k.a. Carolus Linnæus) became a professor of medicine and botany at Uppsala University. He developed a system for naming plants and animals, the binomial system that is still used today. In 1758, he described Coluber molurus, the first python described by science. Since the start of the 21st century, 12 new species of pythons have been described, and more are likely to follow. Some of these will be the result of raising subspecies to species level, others will be new discoveries, and most will be from the Indonesian Archipelago and Australasia region. Pythonidae consists of eight clades which mostly agree with the ten recognized genera. This arrangement and the names are always subject to change as data is added, so there is a good chance that by the time this goes to press there will be some differences. There are two unlikely sisters to the family Pythonidae, the Asian Sunbeam snakes (family Xenopeltidae) and the Mexican Sunbeam Snake, bicolor (family Loxocemidae). Molecular studies consistently show these two families as the sister to the pythons. Thus, the closest living relatives of the Pythonidae appear to be the Mexican Sunbeam Snake, Loxocemis bicolor and the Southeast Asian Sunbeam Snakes in the genus (Vidal & Hedges 2007; Wiens et al. 2008; Pyron et al. 2013). This is in stark opposition to the idea that boas and pythons were each other’s closest relatives and sister taxa. The single genus in the Asian Sunbeam snake family contains two species slightly less than a meter in length. They are fossorial and semiaquatic, have premaxillary teeth, a long left lung, and lay eggs. One species, , is widespread, ranging from India to the . The other, X. hainanensis, is restricted to the island of Hainan. The widespread species is in all likelihood a complex of species. Sunbeam snakes seem to lack the ability to constrict prey, and they have entirely lost 75 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes the pelvic girdle. Pythonids constrict prey, have a remnant pelvic girdle, and most feed on other reptiles. The Mexican Sunbeam Snake, Loxocemus bicolor is the single member of the family Loxocemidae. It inhabits Central America’s west coast and resembles the xenopeltids in general appearance, but it has a remnant pelvic girdle, premaxillary teeth, and a supraorbital bone. Loxocemus hunts in burrows, killing lizards and rodents by constriction, but it is also well known for eating turtle eggs. The family Pythonidae is composed of about 44 extant species in ten genera restricted to the Eastern hemisphere. They range in size from the 700 mm Pigmy Python, Antaresia perthensis, of to the Reticulated Python, Malay- opython reticulatus, which may reach or exceed 9,000 mm. Five of these species range north of the Equator (Python bivittatus, P. molurus, P. sebae, P. regius, and Malayopython reticulatus). The others are restricted to land masses south of the equator. The family may have originated in Laurasia sometime between 70–43 Ma (Reynolds et al. 2014).

Figure 4-1. Scale nomenclature for python head scales. The Reticulated Python is on the left; the Boa Constrictor is on the right. Note that the head scales on the Boa Constrictor are divided into much smaller scales than those on the python. This arrangement does not hold true for all boas and pythons. A. rostral, B. internasal, C. nasal, D. prefrontal, E. eye, F. supraocular, G. frontal, H. occipitals, I. loreals, J. preocular, K. upper labials, L. postoculars, M. lower labials.

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The Afro-Asian Python Clade, Genus Python The genus Python contains ten extant species and three extinct species. The sister to all of the members of this clade is the Royal Python or , Python regius, a central African species inhabiting savanna and forest-edge habitats. Royal pythons rarely exceed two meters (Reynolds et al. 2014). The Afro-Asian Python clade members can be distinguished from other clades by a longitudinally divided frontal scale, 56 or more dorsal scale-rows around the anterior body, a maximum number of mid-body scale-rows of more than 54; shallow infrared sensing pits present in lower labials 2–4 or 2–5; a suture present in the dorsal margin of the rostral infrared sensing pit; and rostral infrared sensing pits deeper than the lower labial pits. Luiselli & Angelici (1998) studied the food habits of the Royal Python in south- eastern . Female pythons were significantly longer than the males. Both sexes preyed exclusively upon birds and mammals, but there were significant inter- sexual differences in terms of dietary composition. Males preyed more frequently upon birds (70.2% of the total prey), whereas females preyed more frequently upon mammals (66.7% of total prey). An ontogenetic change in the diet occurs in both sexes. Specimens less than 70 cm total length preyed almost exclusively upon small birds (nestlings and immature individuals). Large specimens, more than a meter long, preyed almost entirely upon small mammals. The authors suggest that the two sexes differ in terms of their natural history; males are more arboreal than females, and it is this behavioral difference that explains the observed differences in diet. Given that the Royal Python is the sister to all other pythons, it is not surprising that other members of the clade also have smaller males and larger females, sexual differences in diet, and ontogenetic shifts in prey preferences. These traits appear to be ancestral. The short-tailed python clade is composed of four species of heavy-bodied, short-tailed snakes that are semiaquatic and usually associated with forested streams. The clade includes four species Borneo( Python (Python breitensteini), Blood Python, (Python brongersmai), Sumatran Python (Python curtus), and the Mon Python (Python kyaiktiyo) that occur in Indochina and the Greater . Shine et al. (1999) studied the two Sumatran species (P. curtus and P. brongersmai) and found that females of both species are larger than males. Both species feed almost exclusively on rodents and have adapted well to living on oil palm plantations. Reproduction is seasonal, with females reproducing every other year with clutches of 12–16 eggs.

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The distribution of the short-tailed python clade was examined by Zug et al. (2011), and they suspected the distribution of these snakes might have been altered by humans transporting the snakes for commercial trade. Python brongersmai has the broadest distribution, ranging from northern peninsular Thailand to Bangka and Belitung Islands off the southeast coast of Sumatra. The northern disjunct populations of P. brongersmai were described as the new species P. kyaik- tiyo from Mon State, . The species is known only from the type locality on the western face of the Tenghyo Range. The Borneo Python is widespread on the island.

Figure 4-2. The Angolan Python, Python anchietae, is found in southern Angola and northern . It inhabits rocky outcrops and areas of rocky terrain in open brush or , using rock crevices for shelter. Its diet is mostly small mammals and birds. Females produce small clutches of four to five eggs. JCM

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The Indian and Burmese Pythons, long confused with each other, are sister species. The Burmese Python, Python bivittatus, and the Indian Python, Python molurus, have complementary and slightly overlapping distributions, with a combined range extending from Pakistan’s Indus River Valley to southern . Disjunct populations of the Burmese Python are present in Sichuan, China, and Java, Indonesia. The Indian Python, however, does not occur beyond eastern India. These large to giant species will be discussed in detail in another chapter. The Escarpment Python or Angolan Python, Python anchietae (Figure 4-2), was recovered as the sister to the African Python, Python sebae, and this group probably also includes the South African Python, Python natalensis. However, P. natalensis was not included in the study by Reynolds et al. (2014). Body size differences are dramatic in this clade, and it suggests that large body size may have evolved twice independently within the genus. Rocky outcrops on mountainous terrain and brushy plains form the habitat for the Escarpment Python, Python anchietae. It is restricted to southern Angola and Namibia along 1,250 km of the Great Escarpment, with most of its distribution in central Namibia. The distribution appears to be limited to between 750–1,600 m above sea level (Auliya 2010). The large African species will be discussed in a separate chapter. The extinct members of the genus are Python europaeus from the Middle of (13.7–16.9 Ma), Python maurus from the Middle Miocene of Morocco (11.1–12.8 Ma), and Python sardus from the Middle Miocene of Italy (11.1–13.7 Ma). The Indo-Malaya Python Clade, Genus Malayopython Malayopython contains four extant taxa: Malayopython reticulatus, M. r. jampeanus, M. r. saputrai, and M. timoriensis. These are large snakes; all exceed two meters. The species in this genus were long considered to be members of the genus Python. The Malayopython clade can be distinguished from other clades by having shallow anterior supralabial pits and deeper posterior lower labial pits (Python has deep anterior supralabial pits and shallow lower labial pits). Malayopython timoriensis reaches about three meters in total length. It occurs in the (, , Solor Adonara, Lomblem, Pantar, and their satellites). Females lay small clutches of 5–8 eggs. This is a poorly-known species (see de Lang, 2011 for a summary), and it should be noted that it does not occur on the island of Timor. Malayopython reticulatus occurs throughout the islands of the Indonesian Archipelago and may have an isolated population in northeast India. The clade will be discussed in detail in a later chapter.

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The Carpet & Tree Python Clade, Genus Morelia Six extant taxa and one fossil form are in the genus Morelia, a clade of pythons restricted to Australia and New Guinea. This is a diverse clade of medium to large pythons that use a variety of habitats. The Rough-scaled Python, Morelia carinata, of northwestern Australia is morphologically distinctive, recently described, and the sister to the rest of the clade. Porter et al. (2012) reported on seven clutches of eggs totaling 71 viable offspring over a period of six years. Captive specimens are seden- tary, spending much of their time tightly coiled in arboreal or saxicolous ambush sites. Females produce clutches of 10–14 eggs, which hatch as large, slender-bodied offspring about 406 mm SVL and 16.9 grams. Growth is rapid; captive males attain sexual maturity about 1,000 mm SVL and 18 months of age; females mature at 1,400 mm and 30 months of age. Hatchlings reluctantly accept mammalian prey unless they had been scented with frogs or birds, suggesting that frogs may be important in the diet of juvenile snakes. The Centralian Python, Morelia bredli, is the sister to the Diamond Pythons, Morelia spilota, M. s. variegata, and M. imbricata. It is restricted to the arid, south- central region of the . They may exceed two meters in length and remain poorly known. The Carpet and Diamond Pythons occur in numerous habitats, ranging from deserts to . They may reach four meters in total length, are often nocturnal, and they feed on vertebrates. Female Southern Carpet Pythons, Morelia imbricata, grow to more than twice the length and more than 10 times the mass of adult males. Mean adult size is 1.04 m SVL (0.305 kg) for males versus 2.14 m SVL (3.9 kg) for females. This size difference is a consequence of males ceasing to grow due to a reduced rate of feeding. Captive males display low feeding rates, suggesting their “dwarf” size reflects genetic control rather than local prey availability. Observations of free-ranging snakes suggests males do not engage in combat during the mating season, and that larger body size does not enhance male mating success. These results agree well with previous inter- pretations of the relationship between mating systems and sexual size dimorphism in snakes (Pearson et al. 2002). The Diamond Python, Morelia s. spilota, was studied in the field near Sydney, NSW by Slip and Shine (1988) and found to be sedentary in summer and fall and more active in spring, with males moving long distances on a daily basis. In the winter they sheltered in rocky crevices. Home ranges were large, about 124 ha. This species displays the greatest geographic variation in sexual-size dimorphism recorded for any vertebrate species (Pearson et al. 2002). The status of some subspe- cies needs clarification. Cryptic sibling species within the Green Tree Python were found by Rawl- ings & Donnellan (2003). The two species have a genetic divergence of about 7% in mitochondrial DNA (cyt b gene) between the northern and southern lineages. 80 Copyrighted Material Some pages are omitted from this book preview. Chapter 4 - Pythonids – An Overview

Figure 4-3. The Green Tree Python, Morelia viridis. This species is distributed throughout main- land New Guinea, its offshore islands, eastern Indonesia, and the northeast of Australia. It is found between sea level and an altitude of 2,000 m above sea level and is restricted to moist forests from lowland to mid-montane altitudes. JCM

They are separated by the Central Mountain Range that divides New Guinea in an east-west direction. So, there are two species, one from north of the central cordil- lera, and the other from the south, including the Aru Island and Australian popula- tions. Nevertheless, within the southern lineage, the Australian material formed a well-supported clade, whereas material from Aru Island clustered with that from Merauke and Timika. The name azureus Meyer 1874 is available for the northern lineage, having its type locality on Biak Island; since the types were lost, Barker et al. (2015) designated a neotype. The Southern Green Python (Morelia viridis), and the Northern Green Python (Morelia azurea), are highly arboreal. Wilson et al. (2006) radio-tracked 27 indi- vidual Southern Green Pythons (17 females and 10 males) for up to 18 months, locating snakes day and night. The home range size for adult females was about 6.5 ha and was correlated with snout–vent length. Adult males and juveniles did not have stable home ranges, but adult females did have stable home ranges that overlapped with other females and juveniles. Adult males passed through the terri- 81 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes tories of adult females. Males may increase the rate at which they encounter mature females by roaming instead of holding a home range or territory. In captivity, males fight in bouts that includes biting. It is likely that continual movement by mature males have a lot to do with territoriality as well. We think this is to ensure genetic diversity as well. The single known fossil species that belongs to this class is the extinct Riversleigh Python, Morelia riversleighensis†. It is known from the Middle Miocene (10.4–16.3 Ma) from Riversleigh, Queensland, Australia. The Dwarf Python Clade, Genus Antaresia Five extant taxa of questionable relationships are in the genus Antaresia. They are small, usually less than one meter in length, and widespread across Australia. The dwarf pythons have been considered sisters to Morelia, Aspidites, and Leiopython in various analyses. Children’s Python, Antaresia childreni, and the Large-blotched Python, Antaresia stimsoni orientalis, appear to be sisters and are probably related to A. s. stimsoni. The , Antaresia maculosa, and the , Antaresia perthensis, may also have a sister relationship. Antaresia are small, with enlarged, symmetrical head shields, infrared sensing pits on lower labials, but none on the rostral; and three or more loreal scales.

Figure 4-4. The Large-blotched Python, Antaresia stimsoni, ranges from the coast of Western Australia through central Australia as far as the Great Dividing Range. It is an arid land species inhabiting rough terrain with rocky outcrops, eucalyptus woodlands, arid shrublands, and deserts. A nocturnal , it consumes lizards, frogs, and mammals. JCM 82 Copyrighted Material Some pages are omitted from this book preview. Chapter 4 - Pythonids – An Overview

Stimson’s Python occurs throughout Australia’s arid zone. McDonald et al. (2011) studied this species in the MacDonnell Range’s bioregion of the Northern Territory. They found that it remains active over a broad range of air temperatures and maximizes activity after rainfall when humidity is high and ground-dwelling frogs are abundant. It is inactive during the coldest months of the year. Antaresia stimsoni uses a range of vegetation types but prefers riparian woodland. Diet includes a variety of terrestrial vertebrates, and sexual dimorphism is unknown. Of interest, Lourdias et al. (2008) found that A. childreni embryos are in an advanced stage of embryonic development at oviposition, and that thermoregulation has a significant influence on reproduction, with reproductive females maintaining higher and less variable body temperatures than non-reproductive females. The Anthill Python, Antaresia perthensis, is the smallest of the pythons, only reaching 50 centimeters in length and weighing only 200 grams. This tiny python often inhabits and ant mounds. The lay small clutches of two to four very large eggs. Reynolds et al. (2014) suggest that Antaresia and Morelia are paraphyletic. Barker et al. (2015) attribute this to molecular characters simply not yet satisfactorily recov- ering the relationships of the Indo-Australian pythons. Here we are retaining each as a distinct clade.

Figure 4-5. The Olive Python, Liasis olivaceus, inhabits Western Australia, the Northern Territory, and Queensland. Olive Pythons inhabit rocky streams, gorges with pools of water, caves, and rock crevices, and occasionally hollow logs and burrows under rocks. The diet includes birds, mammals, and other reptiles. It ambushes prey along game trails and from the water. JCM 83 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes

The Water Python Clade, Liasis + Apodora The Australasian – Eastern Indonesian genus Liasis contains six extant and one extinct . The Papuan Python, Apodora papuana, is the sister to the Water Pythons, Liasis, and is monotypic. The other members are the Water Python Liasis( fuscus), the Freckled Python (Liasis mackloti), the Western Olive Python (Liasis olivaceus barroni), the Olive Python (Liasis o. olivaceus), Python (Liasis dunni), and the Savu Python (Liasis savuensis). The extinct Liasis dubudingala is known from the Lower Pliocene (4.5–4.0 Ma) at Down’s Bluff Station, in northeastern Queensland. These tend to be medium to large snakes with plate-like scales on the crown, infrared sensing pits absent from the rostral, but present on lower labials, and one or two loreal scales. Most are uniform in color and lack a pattern, with the exception of scattered spots. Plasticity of the Water Python’s life history traits were studied by Madsen & Shine (1999) along the Adelaide River floodplain in tropical Australia. Subpopulations of the pythons separated by 2 km differed in the timing of reproduction, survival rates, cost of reproduction, and in the frequency of reproduction. The differences appear to result from nest-site characteristics. Female Water Pythons used two types of nest sites: those with relatively low, variable temperatures, often in the hollows of paperbark root systems on the edge of the floodplain (cool nests), and those with higher, constant temperatures in burrows dug by large varanid lizards in the higher, drier ridges (hot nests). Cool nests delayed reproduction and reduced survival rates of hatchlings in at least one year during the study. Females laying eggs in cool nests attended the clutch throughout the two- month incubation period. In contrast, females laying eggs in hot nests deserted the clutch within a few days of egg-laying. Females attending eggs did not feed and were emaciated by the end of incubation. Many cool-nesting females died from starva- tion or predation. Surviving cool-nesting females required two years to replenish their energy reserves before producing another clutch. Hot-nesting females had higher rates of survival, and most could reproduce the following year. Most females showed nest-site fidelity in successive clutches, but some moved between hot and cool nest sites. The Scrub Python Clade, Genus Simalia The genus Simalia contains seven extant taxa: The Amethystine Python Simalia( amethistina), the Black Python (Simalia boeleni), the Southern Moluccan Python (Simalia clastolepis), the Scrub Python (Simalia kinghorni), the Tanimbar Python (Simalia nauta), the Oenpelli Python (Simalia oenpelliensis), and the Python (Simalia tracyae). 84 Copyrighted Material Some pages are omitted from this book preview. Chapter 4 - Pythonids – An Overview

The Scrub Python clade can be distinguished from Apodora, Aspidites, Antaresia and Liasis by the presence of two large, deep infrared-sensing pits on the rostral scale and well-developed pits on upper labials 2–5. Aspidites and Bothrochilus have no infrared-sensing pits on the rostral or upper labials. Antaresia and Liasis have no rostral infrared-sensing pits. Apodora has shallow pits on the rostral and upper labials 2–3. Leiopython has a rostral pit and may have pits on the first 2–3 upper labials. Members of the Scrub Python clade have more than four scales between the loreal and upper labials; Bothrochilus and Leiopython have one or two. Scrub Python clade members can be distinguished from Morelia by the plate-like supraocular and frontal scales and one or more pairs of parietal scales. The exception is the Oenpelli Python, Simalia oenpelliensis, which has small parietals and irregular scales posterior to the supraoculars that contact the large frontal. The only large scales on the crown of Morelia are the internasals and prefrontals, the exception being M. carinata, with a single round frontal between the eyes, separated from the supraoculars

Figure 4-6. The Black Python, Simalia boeleni, is found in Indonesia’s in the Wissel Lakes region, as well as in the Papua New Guinea provinces of the Eastern Highlands, Central, and Morobe, and on Goodenough Island. It is a montane forest species found at elevations above 1,000 m. Adults can exceed 3 meters. Photo by Ari Flagyl. 85 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes

The Black-headed Python Clade, Aspidites The genus Aspidites contains two extant taxa: the Black-headed Python (Aspidites melanocephalus), and the Woma (Aspidites ramsayi), and they are the sister to the Ringed Python clade. Both Aspidites exceed two meters. They can be distinguished from other pythons by the absence of infrared sensing pits, a head barely distinct from the anterior body, and a crown with plate-like scales. They are endemic to Australia, feeding mostly on reptiles, but they will also take birds and mammals. Bruton (2013) radio-tracked the Woma, A. ramsayi, in southwestern Queensland and found that while it is a burrowing species, it will climb to prey on sleeping lizards (Pogona barabata). It also feeds on Sand Goannas, Varanus gouldii, and Yakka , Egernia rugose. Active foraging behavior, as well as ambush behavior, is used by this species. This may be the only python that excavates its own burrow. Black- headed Pythons bask with their head out of the burrow, thus the black pigment may facilitate thermoregulation (Shine 1991).

Figure 4-7. The Woma, Aspidites ramsayi, was at one time common throughout Western Austra- lia’s sandplains; it has become critically endangered in some regions. Adults are about 1.5 m. The Woma lacks heat-sensing pits. The range in Southwest Australia extends from Shark Bay, along the coast and inland regions, and was previously common on sandplains. Much of its prey is caught in burrows, where it uses a loop of its body to kill its food. JCM

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The Ringed Python Clade, Bothrochilus + Leiopython The Bothrochilus clade contains seven extant taxa: Ringed Python (Bothrochilus boa), Northern White-lipped Python (Leiopython albertisii), Biak White-lipped Python (Leiopython biakensis), Karimui Basin White-lipped Python (Leiopy- thon fredparkeri), Huon Peninsula White-lipped Python (Leiopython huonensis), Southern White-lipped Python (Leiopython meridionalis), and the Wau White- lipped Python (Leiopython montanus). All are endemic to New Guinea. Reynolds et al. (2014) suggested that Bothrochilus should hold all of these species, Barker et al. argue against this action, pending further analysis. The Bismarck Ringed Python, Bothrochilus boa, has a maximum size of about 1.7 m and is endemic to the (Papua New Guinea). Natusch & Lyons (2012) surveyed Indonesian wildlife traders for ecological information on Leiopython. Leiopython albertisii and L. meridionalis exhibit similar ecological attri- butes; however, L. meridionalis has a longer and wider head than L. albertisii. White- lipped pythons prey mainly on mammals, although smaller individuals feed on lizards. Males and females are similar in size, or females are just slightly larger than

Figure 4-8. D’Albert’s Python, Leiopython albertsii, occurs at low elevations throughout New Guinea and on some satellite islands. Habitats include rainforests and swamps, as well as disturbed areas. They will take refuge in water when disturbed. The diet includes mammals and birds that are captured at night by ambushing on the forest floor. Photography Bill Love.

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males. Reproduction appears to be seasonal, with egg-laying and hatching occur- ring between December and March. Clutch sizes range from 5–17. The Northern White-lipped Python is more often in trade, and traders differentiate between the two species on the basis of color; however, both were traded under the name L. albertisii. Pythonids are Eastern Hemisphere snakes composing eight clades. Their greatest diversity is in Australasia, but it is unclear if this is where they originally evolved. All pythons are constrictors and use both active foraging and sit-and-wait predation. They are all oviparous with the female often attending the eggs, and in some cases generating body heat to control embryonic development.

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Chapter Five Giant Constrictors of Australasia

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Chapter Six Giant Pythons in the Afro-Asian Clade

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Chapter Seven The Reticulated Python Clade

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Chapter Eight Booid Snakes, an Overview

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Chapter Nine The Boa Constrictor Clade

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Chapter Ten The Anaconda Clade, Giant Aquatic Boas

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Chapter Eleven Giant Snakes in Captivity

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Chapter Twelve Invasive Giant Snakes in Florida

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Chapter Thirteen Extinct Giant Snakes

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Giant snakes existed in deep time. The evidence is often just a few excep- tionally large vertebrae, but sometimes there is also a partial skull. Ancient remnants supply tantalizing evidence of paleo giant snakes. Snakes larger than the living species crawled and swam their way through prehistoric Earth. Below is a list of extinct species that likely attained 6.1 m.

Figure 13-1. A phylogenetic tree with the fossil snakes included. From Willson et al.

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Appendix 1. Some DNA Basics The genetic code for all organisms is found in the molecule deoxyribonucleic acid, usually shortened to DNA. While the acronym DNA has infiltrated our everyday lives in criminal investigations and medical research, it is also a powerful tool used by biologists to unravel the history of life. DNA occurs in two locations in the cell, the mitochondria (mtDNA) and in the nucleus (nDNA). Most advanced organisms (eukaryotes) have about a meter of nDNA in each cell, and it is usually divided into small packets called chromosomes. The entire complement of DNA in an organism is referred to as its genome. DNA is a linear molecule coiled into a double helix, perhaps most easily thought of like a ladder-shaped molecule that is twisted into a tightly coiled structure. Each side of the ladder is a strand of many nucleotides. On the outside of each side of the ladder is a backbone composed of 5-carbon sugars and phosphate molecules; the steps of the ladder are the nucleotides. There are four kinds of steps, and the sequence of the steps is the code. A sequence of three steps forms a codon that codes for an amino acid. The mitochondrial DNA is sometimes written mtDNA. When commonly discussing DNA the term genome is applied, and when used in this way refers to the nuclear genome (nDNA). The mtDNA is always inherited from the female parent. Usually, half of the chromosomes (one member of each pair) is inherited from each parent in sexually reproducing organisms. The DNA code contains four kinds of nucleotides, which are represented by four letters A-T-C-G. These letters can be thought of as the DNA alphabet, and each DNA word (a codon) is composed of three of those letters. Most codons code for one of twenty-three amino acids, and since there are 64 possible codons, most amino acids are coded for by multiple codons. Thus, the genetic code is a bit sloppy. Amino acids are the building blocks of proteins, so a string of DNA codons tells the cell what amino acids need to be placed at a particular location in a protein. Place an amino acid in the wrong place in the sequence and the protein may not function correctly. About 70% of the dry weight of living matter is protein. Proteins are important in cells, because they not only form structures, but they carry out and regulate cell processes. One way to think about genes is that they are the instructions for making proteins. A section of DNA acts as a template for making a strand of messenger RNA (ribonucleic acid). The messenger RNA leaves the cell’s nucleus and goes to a ribosome in the cell. The ribosome then matches the RNA codons with the correct amino acid to build a chain of amino acids (a polypeptide) that the cell will eventu- ally fold into a protein (or in some cases leave it as a polypeptide).

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Appendix 2. How the Anaconda Got its Name A storm at sea on 19 November 1659 broke the mast on the ship Anne and stranded its crew in Ceylon. The ship was in the service of the British East India Company and under the command of Robert Knox. Knox was accompanied by his son of the same name. The crew of the Anne was detained by the king of Ceylon, Raja Sinha. Political tensions between European countries and the king of the South Asian island nation resulted in Anne’s crew being forbidden to leave the island by Sinha. Knox and his men were free to move about the country but could not leave. Eventually father and son contracted malaria. The older man died, and the younger Knox lived on the island for almost 20 years until he finally escaped. In his meticulously written and thoughtful account of life in Ceylon, Robert Knox described a large snake. He wrote, “The Pimberah, the body whereof is as big as a man’s middle, and of a length proportionable. It is not swift, but by subtlety will catch his prey; which are Deer or other Cattel; He lyes in the path where the deer use to pass, and as they go, he claps hold of them by a kind of peg that growes on his tayl, with which he strikes them. He will swallow a Roe Buck whole, horns and all; so that it happens sometimes the horns run thro his belly, and kill him” Pimberah was, and is, the well-established Sinhalese name for the species now known to science as Python molurus or Python molurus pimbura, the only large python to inhabit Sri Lanka. As we will see, Knox’s account is important for a quite unexpected reason. The first appearance of the name “anaconda” in the English language was in a tall tale told by “R. Edwin” (Boyle 2008). What follows is the unusual story of the origin of the name “anaconda,” the largest extant snake. It is a story of how a simple clerical error becomes a fact and gets twisted and turned in the human brain, only to emerge as new information that becomes widely accepted knowledge. It was a warm, humid day on the island of Ceylon (now known as Sri Lanka) when the traveler observed an enormous snake in ambush posture, hanging from the crown of a palm tree. He alerted a group of men to the snake’s presence and soon a decision was made to hunt down the snake. On horseback, and armed with firearms, they rode to the serpent. They found the snake to be larger in diameter than a man’s waist, very long, and exceptionally nimble. The men, undetected by the snake, aimed their rifles and fired, but missed the snake completely. The snake ignored the dozen or so bullets that flew past and the men returned home, resolute to continue the hunt the next day. The traveler, R. Edwin, wrote, The Ceylonese seemed to know the creature well; they call it anaconda and talk of eating its flesh when they caught it,” wrote the traveler. The following morning a hundred men returned to the tree and found the snake 289 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes

Appendix 3. Giant Snakes in Oral Traditions Giant snakes have a long held a position in folklores and mythologies, including popular modern culture. Consider the outlandish, but famous Anaconda movie franchise that lasted more than a decade. Murphy and Henderson (1997) catego- rized snakes of exaggerated sizes with supernatural powers as supersnakes. These fanciful serpents frequently exceed 50 feet (15.2 m), have eyes the size of plates, move with lightning speed, and are the likely product of confusion of real snakes with those of mythologies. Supersnake and mythological snake stories from Africa are of greater relevance because it seems probable they are closest to the ancestral supersnake story, given the African origin of Homo sapiens. As humans moved out of Africa to colonize Eurasia, apparently numerous times, they carried their belief systems and folklores with them, and it is probable that the remnants of the mother of all supersnake stories lie within extant African cultures. The hunter-gathers and early agricultural peoples of southern Africa painted large snakes in their rock art. Hoff (1997) determined that the Khoekhoen refer to large snakes with supernatural powers as Great Snakes. Interviews with members of the Khoekhoen, the Northern Cape and southern Namibia descendants of the /Xam, a culture and people from the Upper Karoo previously believed extinct, provided the basis for Hoff’s work. The Khoekhoen believe in huge, train-sized snakes they call Great Snakes, which are divided into two categories: aquatic and terrestrial. The terrestrial Great Snake is the Veldt Snake, which likes water, predicts rain, and is active in the veldt after rain. Killing a Veldt Snake will cause drought or torrential rains and lightning. To the Khoekhoen, Great Snakes are real supernatural beings, not just part of their folklore and live south of Namibia, while to the north they are replaced by other snakes. The /Xam do not believe in the Veldt Snake, but the Water Snake is important in their belief system; it is a color and shape shifter, so that it can catch people. To the /Xam, the Water Snake can instill sexual potency and health by licking a person or spitting into their mouth. But the Khoekhoen also believe the Water Snake preys on humans who venture too close, and it may enter a person’s home to catch someone he is in love with. It kills by stabbing the victim with its tongue, removing a person’s brain by inserting its tongue into their nose, and sucking out a person’s blood. Water Snakes also seduce women and take them underwater for copulation. The Water Snake does not like menstruating women and may cause women in estrus to break out in blisters. They also have the power to change a person into a frog. Hoff suggests that the /Xam’s more positive view is older and that both groups initially regarded the Water Snake as an active force. The is a name coined by British anthropologist Radcliffe- Brown (1926) for a collection of Aboriginal names applied to mythical serpent-like creatures. These names were associated with a similar set of beliefs and related to

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Appendix 4. Attacks and Deaths from Constrictors Naturalist and writer Roger Caras (1964) was skeptical about constricting snakes being able to kill a human, and he was unable to locate anyone who had witnessed the death of a human in the coils of a constrictor. This list is not meant to be exhaustive.

Green Anaconda, Eunectes murinus Bates 1863:236 reported an attack by an anaconda on a 10-year-old boy playing in shallow water. The boy was rescued by his father. Blomberg (1956) reported two attacks by anacondas. A man on the Napo River in was killed while swimming; his body was found downstream. Apparently, the snake was not large enough to swallow him. A second attack occurred at the mouth of the Yasuni River: a 13-year old boy was attacked, swallowed, and later regurgitated. The boy’s father killed the snake. 2007, February 9, Cosmorama, Brazil. Eight-year old Joaquim Pereira was attacked by a wild 16-foot anaconda. His grandfather saved him by beating and stab- bing the snake. (AP story).

Reticulated Python, Malayopython reticulatus 1910. Carl Hagenbeck described four of these large pythons simultaneously attacking one of his sons as he entered their cage. He survived the attack, but the pythons were not easily subdued. 1911, July 2, Northwest Bangladesh. The Sunday Standard. A man half swallowed by a python was killed during a rescue attempt by his neighbors. The snake was said to be 10 m. Both the man and the snake died. [Hamadryad 1977, 2(3):5]. 1921, July 13, Island of Salibathoe (north of Sulawesi). A 14-year-old boy went into his family’s garden to pick vegetables. He did not come home. His parents searched for him but did not find him. On July 15 village men searched for him. One man found vegetables lying on the ground. Two meters away they found blood, and another 10 meters away they found a large snake. The snake was killed, cut open, and the boy was found. (Kopstein, 1927). 1965, September. Reuters reported a 10-year-old girl stepped on a 20-foot reticulated python. The snake attacked. A gardener saved the girl. 1965, September. Reuters reported an 8-year-old boy swallowed by a reticulated python in Ye Village in lower Burma. The boy’s body was retrieved when villagers killed the snake. 1978, Kentucky, USA. A 33-year old was killed by a captive 3.8 m M. reticulatus (McCarty et al. 1989).

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Glossary ambush stalking — A behavior in which a giant snake is alerted to prey that is too far away to reach, so the snake stalks the prey using available cover (including shadows) to conceal their presence until the snake is within striking range. amplified fragment length polymorphism (AFLP) — a method for detecting polymorphisms in DNA. The procedure involves three steps: (1) digestion of cellular DNA with one or more restriction enzymes and ligation of restriction half-site-specific adaptors to all restriction fragments; (2) selective amplification of some of these fragments with two PCR primers; (3) electrophoretic separation of amplicons on a gel matrix so the band pattern can be visualised. automixis — A form of parthenogenesis in which two haploid gametes from the same meiosis combine. clade — A group of organisms evolved from a common ancestor. It includes that ancestor. chromosome — A package of nucleic acids and protein found in the nucleus of most living cells; it carries genetic information in the form of genes. crown group, -group, stem group — It is not necessary for a species to have living descendants in order for it to be included in the crown group. Extinct side branches on the family tree that are descended from the most recent common ancestor of living members will still be part of a crown group. For example, if we consider the crown-snakes (i.e. all extant snakes and the rest of the family tree back to their most recent common ancestor), extinct side branches like the Titanoboa are still descended from the most recent common ancestor of all living snakes, so fall within the snake crown group. In phylogenetics, the crown group of a collection of species consists of the living representatives of the collection together with their ancestors back to their most recent common ancestor, as well as all of that ancestor’s descendants. Thus, it is a clade, a group consisting of a species and all its descendants. Often, the crown group is given the designation “crown” to separate it from the group as commonly defined. Snakes are traditionally defined by their traits and contain fossil members that lived before the last common ancestors of the living groups or, like the Sanajeh indicus, that was not descended from that ancestor. Crown-snakes therefore differ slightly in content from the common definition of snakes. A pan-group or total group is the crown group and all organisms more closely related to it than to any other extant organisms. In a tree analogy, it is the crown group and all branches back to (but not including) the split with the closest branch to have living members. The pan-snakes thus contain the living snakes and all

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(fossil) organisms more closely related to snakes than to lizards (their closest living relatives). Therefore, pan-group snakes would include all squamates and their close relatives like the Rhynchocephalia. A stem group is a paraphyletic group composed of a pan-group or total group minus the crown group itself; it therefore contains no living members of the pan- group. This leaves primitive relatives of the crown groups, back along the phylo- genetic line to (but not including) the last common ancestor of the crown group and their nearest living relatives. It follows from the definition that all members of a stem group are extinct. The “stem group” is the most used and most important of the concepts linked to crown groups, as it offers a purely phylogenetic route to classify fossils that otherwise do not fit into systematics based on living organisms. divergence date — A date for an evolutionary divergence event, such as the split between lizards and snakes. An estimate of the timing of this event can be obtained by examining the fossil record or by correlating this particular instance of evolu- tionary divergence with some geological event of known antiquity, including such things as the formation of a mountain range that split the geographic range of a species in two, thus initiating a process of speciation. Once the evolutionary rate is calculated using a calibration, this calibration can then be applied to other organisms to estimate the timing of evolutionary events. feeding bite — A bite that is usually very serious with any python over 3 meters. When it is directed at an animal, it is often followed by the snake’s coiling around and constricting prey. If it is directed at a human, the behavior may be a mistaken attempt at predation. The bite attempt usually does not last long, once the python realizes the mistake. It is important NOT to fight the bite. That will only make it worse, and if it is a large python, it will make them bite and constrict harder and longer. Florida Burmese Pythons — this specifically refers to the feral population of Python bivittatus that inhabit extreme southern Florida. gene — Used informally, it is a unit of heredity that is transferred from a parent to offspring and is thought to determine some characteristic of the offspring. Technically, a gene is a distinct sequence of nucleotides forming part of a chro- mosome. The sequence of nucleotides determines the order of amino acids in a protein that a cell may manufacture. genome — All of the DNA present in a cell. Most cells contain a complete genome. In a eukaryote organism, it occurs in the nucleus of the cell and a small portion occurs in the mitochondria. In prokaryote cells, it is occurs loose inside the cell. — a taxonomic group that does not form a monophyletic clade. They are perhaps best thought of as collection of remnants of ancient snake lineages that have mostly become extinct. It includes two families now considered the

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Amerophidia (Neotropical Pipe Snakes, Aniliidae, and the Dwarf Boa Tropi- dophiidae), three families now considered Uropeltoidea (Asian Pipe Snakes, Cylindrophiidae; the Dwarf Pipe Snakes, Anomochilidae; and the Shield-tailed Snakes, ); three families now considered Pythonoidae (the Pythons, Pythonidae, The Mexican Burrowing Python, Loxocemidae, and the Sunbeam Snakes, Xenopeltidae), and the , which contains five families (the Neotropical Boas, the Boidae; the West African Burrowing Boa, Calabariidae; the New World Dwarf Boa Clade, Charinidae; the Eastern Hemisphere Sand Boa Clade, Family Erycidae; and the Pacific Boa Clade, Candoiidae.) Monster of God — the title of a book written by environmental author David Quammen. parthenogenesis — Reproduction involving only one female. No sperm is required for fertilization. phylogeny — The study of the evolutionary history and relationships among indi- viduals or groups of organisms (species, or populations). These relationships are discovered through phylogenetic inference methods that evaluate observed heri- table traits, such as DNA sequences or morphology. The result of these analyses is a phylogeny or a phylogenetic tree. phylogenetic tree — This is a diagrammatic hypothesis about the history of the evolutionary relationships of a group of organisms. The tips of a phylogenetic tree can be living organisms or fossils and represent the end point for fossil lineages or the present in living lineages. Phylogenetic analyses have become central to understanding biodiversity, evolution, ecology, and genomes. The nodes where the branches split represent the point at which a common ancestor gave rise to two sister taxa. (See "Figure 13-1. A phylogenetic tree with the fossil snakes included. From Willson et al." on page 279.)

Figure G-1. How to interpret a phylogenetic tree.

309 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes power feeding — Feeding the neonate as much as possible to achieve maximum growth and sometimes, early reproduction. This ‘’power feeding’’ dramatically shortens the lifespans, particularly when as adults the snakes are fed more than 7–9 times annually. Most giant snakes kept this way die, usually at 10 years of age or so. The famous, now dead, giant Retic called Twinkie is the best example I can think of. We think giant snakes likely live more than 40 years. Sisters — A or sister taxon is a phylogenetic term denoting the closest relatives of another given unit in an evolutionary tree. The expression is most easily illustrated by a cladogram in Figure 63: A, B, and C each represent a taxon. The sister group to A is B; conversely, the sister group to B is A. Groups A and B, together with all other descendants of their most recent common ancestor, form the clade AB. The sister group to clade AB is C. The whole clade ABC is itself a subtree of a larger tree, which offers yet more sister group branches that are related, but farther removed from the leaf nodes, such as A, B, and C. In cladistics, A, B, and C may represent specimens, species, taxon-groups, etc. If they represent species, the term sister species is sometimes used. The term “sister group” is used in phylogenetic analysis, and only groups identified in the analysis are labeled as sister groups. An example is in birds, whose sister group is commonly cited as the crocodiles, but that is true only when dealing with extant taxa. The family tree is rooted in the dinosaurs, making for a number of extinct groups branching off before coming to the last common ancestor of birds and crocodiles. Thus, the term sister group must be seen as a relative term, with the caveat that the sister group is the closest relative only among the groups/ species/specimens that are included in the analysis. species — A species is the basic unit of biological classification and a , as well as a unit of biodiversity, but satisfactory definitions are elusive. There are probably a hundred species definitions, and most are distinguished by subtle differences from the others. Scientists and conservationists require a species definition that allows them to work, regardless of the theoretical difficul- ties. If Linnaeus had been correct and species were fixed and clearly distinct from one another, there would be no problem. However, evolutionary processes cause species to change continually and to grade into one another. Often a species is defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. While this definition is often adequate, when looked at more closely, it is problematic. For example, with hybridization, in a species complex of hundreds of similar microspecies, or in a ring species, the boundaries between closely-related species become unclear. Among organisms that reproduce only asexually, the concept of a reproductive species breaks down, and each clone is potentially a microspecies. Problems also arise when dealing with fossils, since reproduction cannot be examined. The

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concept of the chronospecies is therefore used in paleontology. Other ways of defining species include their karyotype, DNA sequence, morphology, behavior, or ecological niche. As a practical matter, species concepts may be used to define species that are then used to measure biodiversity, though whether this is a good measure is disputed, as other measures are possible. For this book we have used the evolutionary species concept. An evolutionary species is an evolutionarily divergent lineage, one that has maintained its heredi- tary integrity through time and space and is on a trajectory where it will have a fate separate from its closest relatives. Unlike the biological species concept, the evolutionary species does not rely on reproductive isolation, so it is independent of processes in other concepts. It works for asexual lineages and can detect recent divergences, which a concept based on morphology cannot. However, it does not work in every situation. The concept ultimately leads to splitting existing species and suggests that biodiversity is richer and more diverse than previously thought. The evolutionary species, suggested by George Gaylord Simpson in 1951, is “an entity composed of organisms which maintains its identity from other such enti- ties through time and over space, and which has its own independent evolu- tionary fate and historical tendencies.” This differs from the biological species concept in embodying persistence over time. The evolutionary species concept is very similar or identical to Willi Hennig’s species-as-lineages concept and works for both asexual and sexually-reproducing species. All species are given a two- part name, a “binomial.” The first part of a binomial is the genus to which the species belongs. The second part is called the specific name or the specific epithet (in botanical nomenclature, also sometimes in zoological nomenclature). For example, Boa constrictor is one of four species of the genus Boa. tree of life — wrote, “The affinities of all the beings of the same class have sometimes been represented by a great tree... As buds give rise by growth to fresh buds, and these if vigorous, branch out and overtop on all sides many a feebler branch, so by generation I believe it has been with the great Tree of Life, which fills with its dead and broken branches the crust of the earth, and covers the surface with its ever branching and beautiful ramifications.” This statement has given rise to representing the evolution of organisms with a tree-like diagram. The tree trunk represents the common ancestors, the branches represent lineages, and the ends of the branches represent species or individual organisms. warning bite — A strike and bite that is not usually serious, even with larger pythons. Usually it is more like a simple “hit” with wide-open mouth, then an immediate release and pullback that causes little damage under normal conditions. Most of the time it is a hit, rather than an actual bite.

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References Acharjyo L, Misra C G. 1980. “Growth rate of Indian Python, Python molurus molurus (Serpentes Boidae), in captivity with special reference to age at first egg-laying,” Journal of the Bombay Natural History Society 77:344–350. Acharjyo LN, Misra R. 1976. “Aspects of reproduction and growth of the Indian Python, Python molurus molurus, in captivity,” British Journal of Herpetology 5:562–565. Ackerman L. 1997. The Biology, Husbandry, and Health Care of Reptiles, Vol II. (pages 272-288) TFH Publishing, Neptune City Aglionby J. 2004. No title. The Guardian, January 5, 2004. Akani GC, Eyo E, Odegbune E, Eniang EA, Luiselli L. 2002. “Ecological patterns of anthropogenic mortality of surburban snakes in an African tropical region,” Israel Journal of 48:1–11. Akani GC, Luiselli L, Politano E. 1999. “Ecological and conservation considerations on the reptile fauna of the eastern Niger Delta (Nigeria),” Herpetozoa 11:141–153. Albino AM. 1991. “Las serpientes del Santacrucense y Friasense de Argentina,” Ameghiniana 28 (3-4), 402. Albino AM 1993. “Snakes from the Paleocene and Eocene of Patagonia (Argentina): paleoecology and coevolution with mammals,” Historical Biology 7, 51-69. Albino AM. 2000. “New record of snakes from the of Patagonia (Argentina),” Geodiversitas 22, 247-253. Albino A. 2007. “Lepidosauromorpha,” Pages 87-115. In Patagonian Mesozoic Reptiles (Z. Gasparini et al. eds), Bloomington: Indiana University Press. Albino A. 2011. “Evolution of Reptiles in Patagonia based on the fossil record,” Biological Journal of the Linnean Society 103:441–457. Alexander GJ. 2007. “Thermal biology of the southern African python (Python natalensis): Does temperature limit its distribution,” Pages 51–75. In R. W. Henderson and R. Powell (eds.) Biology of the Boas and Pythons, Eagle Mountain Publishing LC. Alexander GJ. 2018. “Reproductive biology and maternal care of neonates in southern African python (Python natalensis),” Journal of Zoology. Alexander GJ, Marais J. 2007. A Guide to the Reptiles of Southern Africa, Struik. Cape Town. Allen R. 1963. The anacondas. Ross Allen’s Reptile Institute Bulletin No. 14. Alves R, Filho G. 2007. “Commercialization and Use of Snakes in North and Northeastern Brazil: Implications for Conservation and Management,” Biodiversity and Conservation, 16(4):969-985. Amaral A do. 1944. “Notas sõbre a ofiologia neotrópica e Brasílica. VI. Formas de boíneos de recente registo,” Papeis Avulsos, São Paulo 10:41–48. Amaral A do. 1948. Serpentes gigantes, Bol. Mus. Paraense Emilio Goeldi 10:211-237. Amaral A. do. 1976. Brazilian Snakes: A Color Iconography, Univ. Sao Paulo. Ahmed A. 2006. “Conservation Project for “Python Extinctive (sic) Species in District Sialkot, Punjab. WWF – Pakistan,” 8 pages, Project Number 50039601. Ahmed MF, Das A, Dutta SK. 2009. Amphibians and reptiles of northeast India, a photographic guide. Aaranyak, Guwahati. Andrew AL, Perry BW, Card DC, Schield DR, Ruggiero RP, McGaugh SE, Choudhary A, Secor SM, Castoe TA. 2017. “Growth and stress response mechanisms underlying post-feeding regenerative organ growth in the Burmese python,” BMC genomics 18(1):338. Andrews CW. 1901. “Preliminary note on some recently discovered extinct vertebrates from Egypt (Part II),” Geological Magazine (Dec. 4) 8:434-444. Andre E. 1904. A Naturalist in . London: Smith, Elder and Company. Archarjyo LN and R. Mirsa 1975 (1976). “Aspects of reproduction and growth of the Indian Python, Python molurus molurus in captivity,” British Journal of Herpetology 5:562-565. Auerbach RD. 1987. The Amphibians and Reptiles of . Gaborone Printing Works, Botswana, 295 pp. Auffenberg W. 1994. The Bengal Monitor. Gainesville: University Press of Florida, 560 pp. Augstenová B, Pokorná MJ, Altmanová M, Frynta D, Rovatsos M, Kratochvíl L. 2018. “ZW, XY, and yet ZW: sex chromosome evolution in snakes even more complicated,” Evolution. 2018

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Woodruff, D. S. and L. M. Turner. 2009. “The Indochinese-Sundaic zoogeographic transition: a description and analysis of terrestrial species distributions,” Journal of Biogeography 36:803-821. Wootson C. 2017. “Giant pythons keep attacking people in Indonesia—and humans might be to blame,” Washington Post October 4, 2017. Worcester DC. 1898. The Philippines Islands and their people, The Macmillan Company, London. Wrenicke CTJ. 1955. “Pythons,” Journal, Bombay Natural History Society, 53:135-137. Smith-Woodward A. 1905. A Guide to the Fossil Reptiles, Amphibians and Fishes in the Department of Geology and Paleontology in the British Museum (Natural History). 8th Edition, London: British Museum of Natural History. Yi H, Norell MA. 2015. “The burrowing origin of modern snakes,” Science Advances, 1(10):e1500743. Yule H, Burnell AC. 1886. Hobson-Jobson, A Glossary of Colloquial Anglo-Indian Words and Phrases and of Kindred Terms, Etymological, Historical, Geographical and Discursive, London: John Murrray. 1021 pp. Zang L. 2015 (2016). New discoveries in Burmese Pythons, (In Chinese) Zhao E, Adler K. 1993. Herpetology of China: Society for the Study of Amphibians and Reptiles. 505 pages Zhong C. 1993. “First records for Ophisaurus harti and Python molurus bivittatus from Province, China,” Asiatic Herpetological Research 5:103–104. Ziegler T. 2002. Die Amphibien und Reptilien eins Tieflandfeuchtwald-Schutzgebietes in Vietnam, Munster: Natur und Tier- Verlag Gmbtt. Ziegler T. et al. 2007. “The diversity of a snake community in a karst forest ecosystem in the central Truong Son, Vietnam, with an identification key,” Zootaxa 1493:1-40. Zug GR., Ernst CH. 2004. Smithsonian Answer Book: Snakes, Washington: Smithsonian Books. 177 pages. Zug GR., Mitchell JC. 1995. “Amphibians and reptiles of the Royal Chitwan National Park, Nepal,” Asiatic Herpetological Research 6:172–180. Zug GR. et al. 1998. “Herpetofauna of the Chatthin Wildlife Sanctuary, north-central Myanmar with preliminary observations of their natural history,” Hamadryad 23:111–120. Zweifel RG, Norris KS. 1955. “Contribution to the herpetology of Sonora, Mexico: Descriptions of new subspecies of snakes (Micruroides euryxanthus and Lampropeltis getulus) and miscellaneous collecting notes,” American Midland Naturalist 54:230–249.

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339 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes Index A Boa constrictor constrictor 190 Boa constrictor longicauda 191 Acrantophis dumerili 182 Boa constrictor occidentalis 191 Acrantophis madagascariensis 183 Boa constrictor ortonii 192 African Python 79 Boa imperator 63 African Pythons 18, 138 Boa imperator imperator 193 Amblyrhynchus cristatus 31 Boa imperator sabogage 194 Amethystine Python 84 Boa nebulosa 194 Anaconda Clade, 178 Boa orophias 195 anajeh indicus† 283 Boa sigma 195 Angolan Python 78, 79 Boidae 177 Antaresia childreni 57, 82 Booid Snakes 169 Antaresia maculosa 82 Borneo Python 77 Antaresia perthensis 76, 82, 83 Bothrochilus boa 87 Antaresia stimsoni orientalis 82 Burmese Python 32, 55, 63, 70, 79, 108, Anthill Python 83 111, 117, 253 Apodora papuana 84, 103 Burmese Pythons 13 Argentine Boa Constrictor 191 Asian Pipe Snake 43 C Asian Sunbeam snakes 75 Aspidites melanocephala 91 Calabaria reinhardtii 171 Aspidites melanocephalus 86 Calabariidae 172 Aspidites ramsayi 86 Camfield Giant Snake 283 Australopithecus anamensis 7 Candoiidae 177 Central American banana boas 174 B Central American Boa 63 Central American Boa Constrictor 193 Ball Python 63, 77 Centralian Python 80 Ball Pythons 237 Ceylonese Python 34, 137 Banana Boa 174 cf Eunectes sp.† 280 Beni Anaconda 199, 200 Charinidae 174 Biak White-lipped Python 87 Chilabothrus 180 Bismarck Ringed Python 87 Chilabothrus inornata 131 Black-headed Python 86, 91 Children’s Python 57, 82 Black Python 84, 85 Chilobothrus striatus 57 Blood Python 77 Choctaw Giant Aquatic Snake 285 Blue-tongued Skinks 237 Chubut Eocene Boa 280 Bluff Downs Pliocene Python 286 Chubutophis grandis† 280 Boa constrictor 25, 27 Coluber molurus 75, 108 Boa Constrictor 185 Columbian Rainbow Boa 52 Boa constrictor amarali 188 Corallus 178 340 Copyrighted Material Some pages are omitted from this book preview. Index

Corallus caninus 43 G Corallus grenadensis 179 43 Garstin’s Giant Snake 281 Gigantophis garstini† 281 D Green Anaconda 18, 26, 27, 37, 64, 71, 199 DeSchauensee’s Anaconda 199, 200 Green Anacondas 13 Diamond Pythons 57, 80 Green Tree Python 43 Diel color change 56 Bank Tree Boa 179 patagonica 42 Dinosaur-eating Snake 283 H Dominican Boa Constrictor 194 Dumeril’s Boa 182 Hainan Python 122 Dwarf Boa Clade 174 Halmahera Python 84 H. erectus 7 E Hispaniolan Boa 57 Homo erectus 7 Eastern Hemisphere Sand Boa Clade Huon Peninsula White-lipped Python 175 87 E. beniensis 199 Hybrid Python 272 Ecdysis 55 E. deschauenseei 199 I 43 E. notaeus 199 Indian Python 34, 79, 110, 123, 125, Epicrates 180 126 Epicrates maurus 52 Indotyphlops braminus 61 Erycidae 175 Island Boa Clade 180 Escarpment Python 79 Eumeces murinus 64 J Eunectes 178 Eunectes beniensis 200 Jampea Reticulated Python 152 Eunectes deschauenseei 200 Eunectes murinus 13, 27, 37, 201 K Eunectes notaeus 211 Eunectes stirtoni 199 Karimui Basin White-lipped Python 87 Exiliboa placata 174 L F Lachesis muta 8 Facultative Parthenogenesis 58 Large-blotched Python 82 Florida Burmese Pythons 64 Leiopython albertisii 87 Flower Pot Snake 61 Leiopython biakensis 87 Freckled Python 84 Leiopython fredparkeri 87 341 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes

Leiopython huonensis 87 Morelia viridis 43, 81 Leiopython meridionalis 87 M. r. jampeanus 79 Leiopython montanus 87 M. r. saputrai 79 Lesser Sundas Python 151 M. s. variegata 80 Liasis dubudingala 84 M. timoriensis 79, 151 Liasis dubudingala† 286 Multiple Paternity 63 Liasis dunni 84 Myron 43 Liasis fuscus 15, 84 Liasis mackloti 84 N Liasis olivaceus barroni 84, 96 Liasis o. olivaceus 84 Naracoorte Giant Snake 282 Liasis savuensis 84 Neotropical Boa Clade 177 Long-Tailed Boa Constrictor 191 North African Python 18, 144 Loxocemus bicolor 75, 76 North African Pythons 271 North African Rock Python 35 M Northern Green Python 81 Northern White-lipped Python 87 Madagascar Tree Boas 57 Madtsoia bai† 281 O Madtsoia madagascariensis† 282 Malagasy Boa Clade 181 Oaxacan Boa 174 Malagasy Giant Snake 282 Oenpelli Python 84, 100 Malayopython reticulatus 16, 36, 52, 76, Olive Hissing Snake 66 79, 151 Olive Python 84, 96 Malayopython reticulatus jampeanus Orton’s Boa Constrictor 192 152 Malayopython reticulatus reticulatus 153 P Malayopython reticulatus saputrai 153 Malayopython timoriensis 79, 151 Pacific Boa Clade 177 Mali Eocene Snake 285 Palaeophis colossaeus† 285 Mangrove Snakes 43 Palaeophis grandis† 285 Marine Iguana 31 Palaeophis virginianus† 285 Mexican Sunbeam Snake 75, 76 Panamanian Banana Boa 174 Mexican West Coast Boa Constrictor Papuan Olive Python 103 195 Papuan Python 84 M. imbricata 80 Patagonian Giant Snake 281 Mon Python 77 P. brongersmai 77 Morelia azurea 81 P. curtus 77 Morelia bredli 80 Pearl Island Boa 194 Morelia carinata 80 Pigmy Python 76 Morelia imbricata 80 Pilbara Olive Python 96, 98 Morelia riversleighensis† 82, 287 P. kyaiktiyo 78 Morelia spilota 57, 80 P. molurus 76 342 Copyrighted Material Some pages are omitted from this book preview. Index

Potomac Eocene Snake 285 Rough-scaled Python 80 P. reg iu s 76 Royal Python 77, 91 P. reticulatus 18 Pritchard’s Rule 18 S Psammophis mossambicus 66 P. s eb ae 18, 76 Sanziniidae 181 Pterosphenus schucherti† 285 Savu Python 84 Pygmy Python 82 scramble competition 13 Python anchietae 78, 79 Scrub Python 13, 32, 71, 84, 91, 92 Python bivittatus 13, 15, 18, 32, 52, 63, Selayar Reticulated Python 153 64, 76, 79, 108, 233, 253 Shark River Eocene Snake 285 Python bivittatus bivittatus 109, 111 Short-Tailed Boa 188 Python bivittatus hainannus 122 Simalia amethistina 84 Python bivittatus progschai 109, 123 Simalia boeleni 84, 85 Python breitensteini 77 Simalia clastolepis 84 Python brongersmai 77, 78 Simalia kinghorni 13, 32, 84, 91, 92 Python curtus 77 Simalia nauta 84 Python europaeus 79 Simalia oenpelliensis 84, 100 Python kyaiktiyo 77 Simalia tracyae 84 Python maurus 79 South African Python 34, 54, 79, 142 Python molurus 79, 108, 110, 123 Southeast Asian blindsnake 112 Python molurus molurus 34 Southeast Asian Sunbeam Snakes 75 Python molurus pimbura 108, 137 Southern Carpet Pythons 80 Python molurus sondaica 108 Southern Green Python 81 Python m. pimbura 34 Southern Moluccan Python 84 Python natalensis 34, 54, 79, 142 Southern White-lipped Python 87 Python regius 63, 77, 237 Spotted Python 82 Python regiusa 91 Sulawesi Python 123 Python sardus 79 Sumatran Python 77 Python sebae 7, 18, 35, 79, 144 Sunbeam Snake 43

R T

Rainbow Boa Clade 180 Talisma Anaconda 280 Rattus colletti 15 Tanimbar Python 84 Red-Tailed Boa Constrictor 190 Tet’chien or St. Lucian Boa 195 Relative clutch mass 64 The Green Anaconda 201 Reticulated Python 18, 28, 31, 36, 76, The Green Tree Python 81 151, 153 Tiliqua scincoides 237 Reticulated Pythons 16, 131 Titanic Boa 280 Rhinotyphlops 14 Titanoboa 15, 183 Ringed Python 87 Titanoboa cerrejonensis† 65, 280 Riversleigh Python 82, 287 Treeboa Clade 178 343 Copyrighted Material Some pages are omitted from this book preview. Giant Snakes

Typhlopidae 14 Typhlops 14 Typhlops diardi 112

U unectes murinus 199 Ungaliophis 174 Ungaliophis continentalis 174 Ungaliophis panamensis 174

W

Water Python 84 Water Pythons 15 Wau White-lipped Python 87 West African Burrowing Boa 172 Western Olive Python 84 Wetar Python 84 wild bushmaster 8 Woma 86 Wonambi naracoortensis† 282

X

Xenopeltis 43 Xenopeltis unicolor 75 X. hainanensis 75

Y

Yellow Anaconda 199, 211 Yurlunggur camfieldensis† 283

344 Copyrighted Material Some pages are omitted from this book preview. Author & PhotographerGiant Snakes Information

John C. Murphy is a retired science educator with a lifelong interest in herpetology. His other books include: Tales of Giant Snakes; Amphibians and Reptiles of Trinidad and ; Homalopsid Snakes: Evolution in the Mud; Secrets of the Snake Charmer: Snakes in the 21st Century; Dogs and Snakes: avoiding the bite; A Field Guide to the Amphibians & Reptiles of Trinidad & Tobago and Arizona's Amphibians & Reptiles.

Tom Crutchfield has been a professional herpetologist, breeding and collecting reptiles his entire life. At one time he was one of the largest reptile dealers in the world. In addition, he has authored and coauthored several scientific papers and worked in situ on several out-of-country conservation projects.

346 Copyrighted Material Some pages are omitted from this book preview. Copyrighted Material Some pages are omitted from this book preview. Giant Snakes Giant Snakes A Natural History John C. Murphy & Tom Crutchfield

Snakes, particularly venomous snakes and exceptionally large constricting snakes, have haunted the human brain for a millennium. They appear to be responsible for our excellent vision, as well as the John C. Murphy & Tom Crutchfield anxiety we feel. Despite the dangers we faced in prehistory, snakes now hold clues to solving some of humankind’s most debilitating diseases. Pythons and boas are capable of eating prey that is equal to more than their body weight, and their adaptations for this are providing insight into diabetes. Fascination with snakes has also drawn many to keep them as pets, including the largest species. Their popularity in the pet trade has led to these large constrictors inhabiting southern Florida. This book explores what we know about the largest snakes, how they are kept in captivity, and how they have managed to traverse ocean barriers with our help.