tag:blogger.com,1999:blog-74430750878253689002024-03-12T23:04:40.146-06:00Life is short, but snakes are longSnake biology for everyoneAndrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.comBlogger92125tag:blogger.com,1999:blog-7443075087825368900.post-39629012731702423542018-09-09T13:24:00.000-06:002018-09-25T00:30:53.880-06:00Venom resistance in kingsnakes<div style="text-align: justify;">
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<tr><td class="tr-caption" style="text-align: center;">A kingsnake eating a rattlesnake</td></tr>
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Kingsnakes get their name because they eat other snakes, including venomous snakes like copperheads, cottonmouths, and rattlesnakes. They also eat lots of other kinds of prey, including non-venomous snakes, lizards, turtle eggs, and small mammals.<br />
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You often hear people say that kingsnakes are <b>resistant </b>or <b>immune </b>to the venom of copperheads, cottonmouths, and rattlesnakes. There is a subtle difference between the meaning of these two words.<br />
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Resistance is any physiological ability to tolerate or counteract the effects of a toxin or disease. Like many things in biology, resistance is not an all-or-nothing status, but a gradient. High enough resistance can result in immunity, where the toxin or disease has negligible or no effects.</div>
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<tr><td class="tr-caption" style="text-align: center;">A kingsnake eating a cottonmouth</td></tr>
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Individuals can acquire resistance through repeated exposure to low doses of a toxin. The immune system recognizes the toxin as foreign and attacks it. It forms a memory of each attack and stores the pattern for later, which makes later responses to the same toxin quicker and more effective. If the toxin dose is later increased, <a href="https://www.ncbi.nlm.nih.gov/books/NBK27158/#_A1377_">the memory is reinforced & may become stronger</a>. This is how antivenom is made, <a href="http://billhaast.com/serpentarium/index.html">how people become resistant to snake venom</a>, and also how vaccines against infectious diseases work.<a href="#1" name="top1"><sup>1</sup></a><br />
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It is not how kingsnake resistance to viper venom works. Kingsnake resistance is evolved rather than acquired. This means that kingsnakes are born resistant to venom. As far as we know, their resistance levels are fixed for life & don’t change with age or exposure. This has happened over a long time through natural selection, over many generations of kingsnakes. We don't actually have a very exact understanding of the physiological and molecular mechanisms behind how kingsnakes resist the toxic effects of viper venom. <a href="https://www.sciencedirect.com/science/article/pii/0041010171901413">At least some of their resistance comes from antibodies</a>—chemicals in their blood that interfere with the venom—<a href="https://www.jstor.org/stable/pdf/1565123.pdf">because mice injected with kingsnake blood survive viper venom better than those that aren't</a>, and the <a href="https://www.sciencedirect.com/science/article/pii/0041010171901413">chemical composition of kingsnake blood changes after exposure to viper venom</a>.</div>
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<tr><td class="tr-caption" style="text-align: center;">A kingsnake eating a western hognose snake</td></tr>
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Any time a weapon appears, there is potential for counter-weapons (i.e. resistance and immunity) to appear in response. This happens through a process called a co-evolutionary arms race<a href="#2" name="top2"><sup>2</sup></a>. Just as the United States and the Soviet Union were involved in an arms race centered around nuclear weapons during the Cold War, so are venomous snakes and their prey & predators involved in arms races centered around their primary weapon—venom.</div>
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A major difference is that, unlike nations or humans, animals cannot plan for the future and decide to invest more energy in research & development of novel or better weapons technology for future generations. Instead, co-evolutionary arms races happen through natural selection. What start out as tiny variations in toxin resistance can be magnified over many generations. </div>
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<tr><td class="tr-caption" style="text-align: center;">A kingsnake and a copperhead biting one another</td></tr>
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When vipers were first evolving their front fangs, defensive bites became a new option for them. At first, their predators were probably not very good at resisting the effects of the venom, especially if the predator’s physiology was similar to that of their prey, and venom would have made a very good defense mechanism. Vipers would sometimes be killed and eaten, but many predators later died from their bites. Kingsnake predators that were slightly better able to tolerate the effects of the venom were more likely to survive. Eventually, all the kingsnakes without these venom resistance traits had been killed by vipers that they tried to eat, and only the resistant ones remained. On the other side, vipers that had venom with toxins that were, for example, slightly more painful or fast-acting, might have been more likely to survive a predatory attack. Thus, the arms race escalates. Vipers also exhibit <a href="https://www.jstor.org/stable/pdf/3891592.pdf">flipping, jerking, “body bridging” and other escape behaviors as a defense against kingsnakes</a>—suggesting, since they do not try to bite kingsnakes in defense, that their venom is essentially useless as an anti-kingsnake defense mechanism by now and that kingsnakes have “won” this arms race.</div>
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<tr><td class="tr-caption" style="text-align: center;">A mongoose eating a boomslang</td></tr>
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This is why kingsnakes are immune to the venom of copperheads, cottonmouths, and North American rattlesnakes, but not to the venom of, for example, king cobras or black mambas. Because they live on different continents, there has never been an opportunity for kingsnakes and black mambas to enter into a co-evolutionary arms race (although both prey and predators of black mambas in Africa, such as <a href="https://s3.amazonaws.com/academia.edu.documents/43436458/Drabeck_et_al._2015.pdf">honey badgers</a>, and of king cobras in India, such as <a href="https://www.sciencedirect.com/science/article/pii/S0041010197000196">mongeese</a>, have probably accomplished much the same thing).</div>
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Kingsnakes also eat coralsnakes, but amazingly they are not immune to the venom of Eastern Coralsnakes (<i>Micrurus fulvius</i>)—kingsnakes injected with coralsnake venom die quickly, and kingsnake blood is <a href="https://www.jstor.org/stable/pdf/1565123.pdf">0% effective</a> at neutralizing venom proteins from coralsnakes. Presumably they are able to catch and consume coralsnakes without getting bitten. This could be because coralsnakes often eat other snakes, so perhaps their venom is more difficult for kingsnakes to evolve resistance against. Or, perhaps coralsnakes are rare or dangerous prey for kingsnakes, and it’s possible but not worth it for them to evolve resistance.</div>
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<tr><td class="tr-caption" style="text-align: center;">A milksnake constricting a Dekay's brownsnake</td></tr>
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Not every kingsnake species has been tested against every venom, but we do know that there is variation in which species are immune to which venoms. <a href="https://www.jstor.org/stable/pdf/1565123.pdf">The only study</a> to compare species in depth injected mice with mixtures of venom & snake blood and measured mouse symptoms and survival. They found that blood from Eastern Kingsnakes (<i>Lampropeltis getula</i>) had the widest spectrum of protection against the venoms tested and was the most effective at neutralizing many rattlesnake venoms, but the least effective against copperhead venom. Blood from kingsnakes from Florida & the Gulf Coast was the most effective at neutralizing the venom of copperheads & cottonmouths. <a href="https://www.jstor.org/stable/pdf/1565123.pdf">Mole Kingsnake</a> (<i>Lampropeltis calligaster</i>) blood is about 75% as effective at neutralizing Mojave Rattlesnake (<i>Crotalus scutulatus</i>) venom as the blood of Eastern Kingsnakes. Gray-banded Kingsnakes (<i>L. alterna</i>) have moderate neutralization potential against Western Diamondback (<i>C. atrox</i>) venom, but none against Eastern Diamondback (<i>C. adamanteus</i>) venom. Blood from milksnakes (formerly all called <i>L. triangulum</i>) from various locations had intermediate neutralization capacity, with blood from North American milksnakes being about 70% more effective against rattlesnake venom than blood from Central American milksnakes. <a href="https://www.jstor.org/stable/pdf/1438111.pdf">Another study</a> found that an eastern milksnake injected with copperhead venom died, and one injected with pigmy rattlesnake venom had "no noticeable ill effects", but a lack of replication prevents these results from being particularly meaningful. Somewhat surprisingly, blood from Long-nosed Snakes (<i>Rhinocheilus lecontei</i>), Cornsnakes (<i>Pantherophis guttatus</i>), <a href="https://www.sciencedirect.com/science/article/pii/0041010182900514">Mussuranas (<i>Clelia clelia</i>)</a>, and <a href="https://www.jstor.org/stable/pdf/20171334.pdf">Japanese Four-lined Ratsnakes (</a><a href="https://www.jstor.org/stable/pdf/20171334.pdf"><i>Elaphe quadrivirgata</i>)</a> was also effective at protecting mice from viper venoms, but blood from pinesnakes (<i>Pituophis</i>) and gartersnakes (<i>Thamnophis</i>) was not. Both <a href="https://www.jstor.org/stable/pdf/1435559.pdf">vipers </a>and <a href="https://www.sciencedirect.com/science/article/pii/004101019090145W">elapids</a> appear to have at least some level of resistance to their own venom, although <a href="https://books.google.ch/books?id=ai_6CAAAQBAJ&pg=PA522&lpg=PA522&dq=%22Antihemorrhagic+Factor+Present+in+the+Sera+of+Snakes%22&source=bl&ots=bYxDOz7sN0&sig=NgmusO-LjzybHndZhO04az5BJLQ&hl=en&sa=X&ved=2ahUKEwjRx6TQ3a_dAhXS_aQKHVEFB6UQ6AEwAHoECAMQAQ#v=onepage&q&f=false">detailed studies are lacking for most species</a>.</div>
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An 1850 folio from the Mahabharata</td></tr>
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Kingsnakes are just one of many species that have partial immunity or resistance to venom. <a href="https://www.sciencedirect.com/science/article/pii/0041010187902601">Hedgehogs</a>, skunks, <a href="http://www.dtic.mil/dtic/tr/fulltext/u2/a034590.pdf">opossums</a>, and possibly <a href="https://www.rufford.org/rsg/projects/sameh_darawshi">snake-eagles</a> also have resistance to viper venoms, and <a href="http://foodweb.uhh.hawaii.edu/MARE494_files/Heatwole%20&%20Powell%201997.pdf">eels are resistant to sea krait venom</a>. Unlike kingsnakes, we have actually <a href="https://www.sciencedirect.com/science/article/pii/S0304416500000222">figured out exactly which proteins</a> in opossum blood are responsible for its snake venom neutralization capacity. We also know that mongeese have followed a different route, <a href="http://www.pnas.org/content/89/16/7717.long">changing the shape of the toxin targets</a> in their cells rather than putting molecules into their blood to combat the toxins (<a href="https://www.sciencedirect.com/science/article/pii/0041010177901052">which means that their immunity cannot be transferred</a>). Other predators of venomous snakes, <a href="https://www.jstor.org/stable/pdf/1438255.pdf">such as indigo snakes (genus <i>Drymarchon</i>)</a>, appear to have gotten away with not evolving immunity, although I was unable to find any actual data on physiological responses of indigo snakes to venom, just statements saying they were not resistant, so it's possible that actual tests have not been carried out.</div>
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Opossum resistance to copperhead venom probably evolved in a similar way to kingsnake resistance, but vipers are also involved in co-evolutionary arms races with their prey. Many rodent prey of North American vipers are resistant, including <a href="https://www.sciencedirect.com/science/article/pii/0041010178900399">wood rats</a>, <a href="https://www.sciencedirect.com/science/article/pii/0041010182901192">prairie voles</a>, and <a href="https://www.sciencedirect.com/science/article/pii/0041010187901279">ground squirrels</a>. Think of how the U.S. during the Cold War had to balance foreign policy not just with the Soviet Union, but also with other nations. The emerging foreign policy is a compromise, just as the venom that evolves is a compromise of selection pressures from predators and prey. Resistant prey may select for venoms that are better at quickly incapacitating, whereas resistant predators may select for venoms that are less deadly and more painful; it’s difficult to predict exactly what will happen without knowing the exact mechanism of resistance. Sometimes s<span style="font-family: inherit;">election from predators and prey may act in the same direction, other times in opposite directions. The details of these processes are what evolutionary biologists study on a day-to-day basis.</span></div>
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<a href="https://www.blogger.com/null" name="1"><b>1 </b></a>Creating a vaccine against snake venom is harder than creating one against an infectious disease that is caused by a virus or a bacterium. There are <a href="http://www.redrockbiologics.com/rattlesnake_vaccine_for_dogs.php">pit viper venom vaccines available for dogs and horses</a>, made from the venom of Western Diamondback Rattlesnakes, but none are available for humans. Additionally, <a href="https://sciencebasedmedicine.org/are-rattlesnake-vaccines-for-dogs-effective/">the canine vaccines must be given twice per year, immediate veterinary care is still required, & protection against other species of venomous snakes is poor</a>, so the technology has a long way to go.<a href="#top1"><sup>↩</sup></a><br />
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<a href="https://www.blogger.com/null" name="2"><b>2 </b></a>The most famous co-evolutionary arms race is between <a href="http://snakesarelong.blogspot.com/2014/07/tetrodotoxin-resistant-snakes.html">toxin-resistant gartersnakes & tetrodotoxin-defended newts in the Pacific Northwest of the US & Canada</a>, although there are many others, such as that between most pathogens & the immune systems of their hosts, between brood parasites such as cuckoos & their hosts, and between bad-tasting plants and herbivores.<a href="#top2"><sup>↩</sup></a><br />
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ACKNOWLEDGMENTS
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If you want to know more, I'd suggest chapter 3 of <a href="http://christiewilcox.com/venomous.html">Christie Wilcox's book <i>Venomous</i></a>, from which I drew while researching & writing this article. Thanks to <a href="https://www.flickr.com/photos/35245294@N04/">Connie Wade</a>, <a href="https://www.flickr.com/photos/nclarkii">Pierson Hill</a>, <a href="https://www.flickr.com/people/alan_cressler/">Alan Cressler</a>, Joe McDonald, Elana Erasmus, and the <a href="https://commons.wikimedia.org/wiki/File:Fight_of_the_Mongoose_and_the_Serpent_Armies_(recto),_Babhruvahana_and_the_Mongoose_Fight_the_Serpents_(verso),_Scenes_from_the_Story_of_Babhruvahana,_Folio_from_a_Mahabharata_((War_of_the)_LACMA_M.71.1.31a-b_(1_of_2).jpg">Los Angeles County Museum of Art [public domain] via Wikimedia Commons</a> for providing their images for this article. Thanks to Laura Connelly for reading a draft of this article.</div>
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REFERENCES</div>
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
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<tr><td class="tr-caption" style="text-align: center;">A kingsnake eating a ringneck snake</td></tr>
</tbody></table>
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<span style="font-size: x-small;">de Wit, C. A. 1982. Resistance of the prairie vole (<i>Microtus ochrogaster</i>) and the woodrat (<i>Neotoma floridana</i>), in Kansas, to venom of the Osage copperhead (<i>Agkistrodon contortrix phaeogaster</i>). Toxicon 20:709-714 <<a href="https://www.sciencedirect.com/science/article/pii/0041010182901192">abstract</a>></span></div>
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<span style="font-size: x-small;">de Wit, C. A. and B. R. Weström. 1987. Venom resistance in the hedgehog, <i>Erinaceus europaeus</i>: purification and identification of macroglobulin inhibitors as plasma antihemorrhagic factors. Toxicon 25:315-323 <<a href="https://www.sciencedirect.com/science/article/pii/0041010187902601">abstract</a>></span></div>
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<span style="font-size: x-small;">Drabeck, D. H., A. M. Dean, and S. A. Jansa. 2015. Why the honey badger don't care: Convergent evolution of venom-targeted nicotinic acetylcholine receptors in mammals that survive venomous snake bites. Toxicon 99:68-72 <<a href="https://s3.amazonaws.com/academia.edu.documents/43436458/Drabeck_et_al._2015.pdf">academia.edu</a>></span></div>
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<span style="font-size: x-small;">Heatwole, H. and J. Powell. 1998. Resistance of eels (<i>Gymnothorax</i>) to the venom of sea kraits (<i>Laticauda</i> <i>colubrina</i>): a test of coevolution. Toxicon 36:619-625 <<a href="http://foodweb.uhh.hawaii.edu/MARE494_files/Heatwole%20&%20Powell%201997.pdf">PDF</a>></span><br />
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<span style="font-size: x-small;">Holding, M. L., D. H. Drabeck, S. A. Jansa, and H. L. Gibbs. 2016. Venom Resistance as a Model for Understanding the Molecular Basis of Complex Coevolutionary Adaptations. Integrative and Comparative Biology 10.1093/icb/icw082 <<a href="https://academic.oup.com/icb/article/56/5/1032/2420622">full-text</a>></span></div>
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<span style="font-size: x-small;">Jansa, S. A. and R. S. Voss. 2011. Adaptive evolution of the venom-targeted vWF protein in opossums that eat pitvipers. PLoS ONE 6:e20997 <<a href="http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0020997">full-text</a>></span></div>
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<span style="font-size: x-small;">Keegan, H. L. and T. F. Andrews. 1942. Effects of crotalid venom on North American snakes. Copeia 1942:251-254 <<a href="https://www.jstor.org/stable/pdf/1438015.pdf">PDF</a>></span></div>
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<span style="font-size: x-small;">Keegan, H. L. 1944. Indigo snakes feeding upon poisonous snakes. Copeia 1944:59 <<a href="https://www.jstor.org/stable/pdf/1438255.pdf">PDF</a>></span></div>
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<span style="font-size: x-small;">Lee, C.-Y., editor. 1979. Snake Venoms. Springer-Verlag, Berlin. <<a href="https://books.google.ch/books?id=ai_6CAAAQBAJ&printsec=frontcover&source=gbs_ge_summary_r&cad=0#v=onepage&q&f=false">full-text</a>></span><br />
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<span style="font-size: x-small;">Liu, Y.-B. and K. Xu. 1990. Lack of the blocking effect of cobrotoxin from <i>Naja naja atra</i> venom on neuromuscular transmission in isolated nerve muscle preparations from poisonous and non-poisonous snakes. Toxicon 28:1071-1076 <<a href="https://www.sciencedirect.com/science/article/pii/004101019090145W">abstract</a>></span><br />
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<span style="font-size: x-small;">Lomonte, B., L. Cerdas, J. Gené, and J. Gutierrez. 1982. Neutralization of local effects of the terciopelo (<i>Bothrops asper</i>) venom by blood serum of the colubrid snake <i>Clelia clelia</i>. Toxicon 20:571-579 <<a href="https://www.sciencedirect.com/science/article/pii/0041010182900514">abstract</a>></span><br />
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<span style="font-size: x-small;">Moussatché, H. and J. Perales. 1989. Factors underlying the natural resistance of animals against snake venoms. Memorias do Instituto Oswaldo Cruz 84:391-394 <<a href="http://www.scielo.br/pdf/mioc/v84s4/vol84%28fsup4%29_381-384.pdf">PDF</a>></span><br />
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<span style="font-size: x-small;">Neves-Ferreira, A. G., N. Cardinale, S. L. Rocha, J. Perales, and G. B. Domont. 2000. Isolation and characterization of DM40 and DM43, two snake venom metalloproteinase inhibitors from <i>Didelphis </i>marsupialis serum. Biochimica et Biophysica Acta (BBA)-General Subjects 1474:309-320 <<a href="https://www.sciencedirect.com/science/article/pii/S0304416500000222">abstract</a>></span><br />
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<span style="font-size: x-small;">Nichol, A. A., V. Douglas, and L. Peck. 1933. On the immunity of rattlesnakes to their venom. Copeia 1933:211-213 <<a href="https://www.jstor.org/stable/pdf/1435559.pdf">PDF</a>></span></div>
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<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Ovadia, M. and E. Kochva. 1977. Neutralization of Viperidae and Elapidae snake venoms by sera of different animals. Toxicon 15:541-547 <<a href="https://www.sciencedirect.com/science/article/pii/0041010177901052">abstract</a>></span><br />
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<span style="font-size: x-small;">Perez, J. C., W. C. Haws, V. E. Garcia, and B. M. Jennings III. 1978. Resistance of warm-blooded animals to snake venoms. Toxicon 16:375-383 <<a href="https://www.sciencedirect.com/science/article/pii/0041010178901587">abstract</a>></span></div>
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<span style="font-size: x-small;"><br /></span></div>
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<span style="font-size: x-small;">Perez, J. C., W. C. Haws, and C. H. Hatch. 1978. Resistance of woodrats (<i>Neotoma micropus</i>) to <i>Crotalus atrox</i> venom. Toxicon 16:198-200 <<a href="https://www.sciencedirect.com/science/article/pii/0041010178900399">abstract</a>></span></div>
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<span style="font-size: x-small;">Perez, J. C., S. Pichyangkul, and V. E. Garcia. 1979. The resistance of three species of warm-blooded animals to western diamondback rattlesnake (<i>Crotalus atrox</i>) venom. Toxicon 17:601-607 <<a href="https://www.sciencedirect.com/science/article/pii/0041010179902344">abstract</a>></span></div>
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<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Philpot, V. 1954. Neutralization of snake venom in vitro by serum from the nonvenomous Japanese snake <i>Elaphe quadrivirgata</i>. Herpetologica 10:158-160 <<a href="https://www.jstor.org/stable/pdf/20171334.pdf">PDF</a>></span><br />
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<span style="font-size: x-small;">Philpot, V. and R. G. Smith. 1950. Neutralization of pit viper venom by king snake serum. Experimental Biology and Medicine 74:521-523 <<a href="http://journals.sagepub.com/doi/abs/10.3181/00379727-74-17959">abstract</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Philpot, V. B., E. Ezekiel, Y. Laseter, R. G. Yaeger, and R. L. Stjernholm. 1978. Neutralization of crotalid venoms by fractions from snake sera. Toxicon 16:603-609 <<a href="https://www.sciencedirect.com/science/article/pii/0041010178901885">abstract</a>></span><br />
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<span style="font-size: x-small;">Poran, N. S., R. G. Coss, and E. Benjamini. 1987. Resistance of California ground squirrels (<i>Spermophilus beecheyi</i>) to the venom of the northern Pacific rattlesnake (<i>Crotalus viridis oreganus</i>): a study of adaptive variation. Toxicon 25:767-777 <<a href="https://www.sciencedirect.com/science/article/pii/0041010187901279">abstract</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Swanson, P. L. 1946. Effects of snake venoms on snakes. Copeia 1946:242-249 <<a href="https://www.jstor.org/stable/pdf/1438111.pdf">full-text</a>></span></div>
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<span style="font-size: x-small;">Voss, R. S. and S. A. Jansa. 2012. Snake-venom resistance as a mammalian trophic adaptation: lessons from didelphid marsupials. Biological Reviews 87:822-837 <<a href="https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1469-185X.2012.00222.x">PDF</a>></span></div>
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<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Weinstein, S. A., C. F. DeWitt, and L. A. Smith. 1992. Variability of venom-neutralizing properties of serum from snakes of the colubrid genus <i>Lampropeltis</i>. Journal of Herpetology 26:452-461 <<a href="https://www.jstor.org/stable/pdf/1565123.pdf">PDF</a>></span><br />
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<span style="font-size: x-small;">Weldon, P. J. 1982. Responses to ophiophagous snakes by snakes of the genus <i>Thamnophis</i>. Copeia 1982:788-794 <<a href="https://www.jstor.org/stable/pdf/1444088.pdf">PDF</a>></span></div>
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<span style="font-size: x-small;">Weldon, P. J. and G. M. Burghardt. 1979. The ophiophage defensive response in crotaline snakes: extension to new taxa. Journal of Chemical Ecology 5:141-151 <<a href="https://link.springer.com/content/pdf/10.1007%2FBF00987695.pdf">PDF</a>></span></div>
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<span style="font-size: x-small;">Weldon, P. J. and F. M. Schell. 1984. Responses by king snakes (<i>Lampropeltis getulus</i>) to chemicals from colubrid and crotaline snakes. Journal of Chemical Ecology 10:1509-1520 <<a href="https://www.researchgate.net/profile/Paul_Weldon3/publication/259248702_Responses_by_king_snakes_Lampropeltis_getulus_to_chemicals_from_colubrid_and_crotaline_snakes/links/00b7d52b05e630cfb2000000/Responses-by-king-snakes-Lampropeltis-getulus-to-chemicals-from-colubrid-and-crotaline-snakes.pdf">ResearchGate</a>></span></div>
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<span style="font-size: x-small;">Werner, R. M. and J. A. Vick. 1977. Resistance of the opossum (<i>Didelphis virginiana</i>) to envenomation by snakes of the family Crotalidae. Toxicon 15:29-32 <<a href="http://www.dtic.mil/dtic/tr/fulltext/u2/a034590.pdf">PDF</a>></span></div>
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<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Wilcox, C. 2016. Venomous: How Earth's Deadliest Creatures Mastered Biochemistry. Scientific American. <<a href="http://christiewilcox.com/venomous.html">official page</a>></span><br />
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<span style="font-size: x-small;">Witsil, A. J., R. J. Wells, C. Woods, and S. Rao. 2015. 272 cases of rattlesnake envenomation in dogs: Demographics and treatment including safety of F(ab')2 antivenom use in 236 patients. Toxicon 105:19-26 <<a href="https://www.sciencedirect.com/science/article/pii/S0041010115300581">abstract</a>></span></div>
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="https://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
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Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com11Jena, Germany50.927054 11.58923720000007150.7669025 11.26651370000007 51.087205499999996 11.911960700000071tag:blogger.com,1999:blog-7443075087825368900.post-58771399243910477152018-03-07T08:11:00.013-07:002023-12-13T13:52:13.654-07:00The House Snake Mess for Dummies<div style="text-align: center;">
<span style="font-size: x-small;">This article will soon be available in Spanish</span></div>
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Inspired by <a href="http://wsed.org/">Mike Van Valen's</a> <a href="https://herpunit.wordpress.com/2016/10/05/the-ratsnake-mess-for-dummies/">"The Ratsnake Mess for Dummies"</a><br />
Please note that the information in this article is current as of March 2018 (no later)<br />
Please <a href="mailto:amdurso@gmail.com">contact me</a> or leave a comment if you spot an error</div>
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Arguably House Snakes are much more of a mess than ratsnakes, which makes sense when you consider that they are they distributed over an area almost 7 times larger, including areas as diverse as the Sahara Desert, Congo Rainforest, Great Rift Valley, East African Savannah, Ethiopian Highlands, Okavango Delta, and Southern African Great Escarpment, and occur in a total of 46 countries, many of which have perennially turbulent political climates. It's no surprise that <a href="http://john.measey.com/media/c8242954-65ff-4830-835a-c820bf22d0d0/SO2nuA/PDFs/Measey_ed_2011_AJH.pdf">the number of herpetologists working in Africa is dwarfed by the number working in North America, and the vast majority of these people have not been of African descent (although that is beginning to slowly change)</a>.</div>
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEifz3ZZnz87LeZofBAqHetftiFG4xuCa0dhy8KyEFA7OIbRS6TAV6SfV1UHm6y_nM8YGH9Uy-_D1UGRm04FLv68mq05uWKxntPqzlXxz7Z51QeHfTCpfpbFyC1IcNXYWZLgSrkhaMZgq9LX/s1600/Boaedon+fuliginosus+Morocco+Budi+Rebollo+Fernandez.jpg" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="600" data-original-width="900" height="213" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEifz3ZZnz87LeZofBAqHetftiFG4xuCa0dhy8KyEFA7OIbRS6TAV6SfV1UHm6y_nM8YGH9Uy-_D1UGRm04FLv68mq05uWKxntPqzlXxz7Z51QeHfTCpfpbFyC1IcNXYWZLgSrkhaMZgq9LX/s320/Boaedon+fuliginosus+Morocco+Budi+Rebollo+Fernandez.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">African House Snake (<i>Boaedon fuliginosus</i>) from the<br />
northernmost part of the range in Morocco.<br />
Like everywhere in Africa, there are probably multiple<br />
undescribed cryptic species within this lineage</td></tr>
</tbody></table>
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What is surprising is that African House Snakes are <a href="https://www.researchgate.net/profile/Ian_Bride/publication/233688340_Dynamics_of_the_trade_in_reptiles_and_amphibians_within_the_United_Kingdom_over_a_ten-year_period/links/55225d330cf2a2d9e1453972/Dynamics-of-the-trade-in-reptiles-and-amphibians-within-the-United-Kingdom-over-a-ten-year-period.pdf">popular in the pet trade</a> and are important model organisms for studies of development, behavior, hormones and reproductive biology, yet we still know almost nothing about them in the wild, even though they are common and tolerant of anthropogenically-disturbed environments.</div>
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When most people think of African House Snakes, the scientific name that probably comes to mind is <i>Lamprophis fuliginosus</i>. In this article, I'll try to explain why this well-known species had to be moved into the genus <i>Boaedon </i>in 2011, and why it will probably be split up into multiple species sometime in the (hopefully-not-too-distant) future. The correct scientific name of many African House Snakes in captive breeding colonies may be difficult or impossible to determine, especially because most people don't know which part of Africa their House Snakes originally came from (and they may have since been bred with House Snakes from other parts of Africa).</div>
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<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEikQc_EiWwbnQpfg7XAlsr-SpYtSYDKVrnqu477Kn-kMuGkDgLGWq5GXBm1GY3yr2F0fuQ1PSpYpuz6ZhrOPQQLcfL0yakW_ULCa8UPLmuRS-wcf_gVnBzYVyncZtwPeLWj3HuflBkWW2ZH/s1600/lamprophiinae+cladogram.jpg" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1600" data-original-width="1144" height="400" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEikQc_EiWwbnQpfg7XAlsr-SpYtSYDKVrnqu477Kn-kMuGkDgLGWq5GXBm1GY3yr2F0fuQ1PSpYpuz6ZhrOPQQLcfL0yakW_ULCa8UPLmuRS-wcf_gVnBzYVyncZtwPeLWj3HuflBkWW2ZH/s400/lamprophiinae+cladogram.jpg" width="285" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><span style="font-size: x-small;">Simplified phylogenetic tree of Lamprophiinae, with</span><br />
<span style="font-size: x-small;">focus on "house snakes" (genera <i>Boaedon </i>& <i>Lamprophis</i>).<br />There's enough uncertainty about the structure within <i>Boaedon</i><br />that I didn't try to represent much of what's known.<br />For more detailed trees, see the <a href="https://s3.amazonaws.com/academia.edu.documents/3221432/2011_Kelly_et_al_Lamprohiidae_MPE.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1518528029&Signature=oG71252i9VyFmcOQhrjp2Vc2q5A%3D&response-content-disposition=inline%3B%20filename%3DKelly_C.M.R._Branch_W.R._Broadley_D.G._B.pdf" style="text-align: justify;">Kelly</a><span style="text-align: justify;">, </span><a href="https://journals.co.za/content/ajherp/64/1/EJC171369" style="text-align: justify;">Greenbaum</a><span style="text-align: justify;">, & </span><a href="http://www.documentation.ird.fr/hor/fdi:010069109" style="text-align: justify;">Trape</a><span style="text-align: justify;"> papers.<br />Green are species lacking genetic data that can't be placed yet.<br />Red stars are multiple cryptic species (there could be more).</span></span><br />
<span style="font-size: x-small;"><a href="https://drive.google.com/file/d/1aSRm-GbUZ6YcT16cALN3zfxOVfNjOCP-/view?usp=sharing">Click here</a> for a larger version</span></td></tr>
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To start, let's get a little taxonomic perspective. <a href="http://www.wienslab.com/Publications_files/Pyron_et_al_2011.pdf">Pyron et al.'s 2011</a> article firmly established <a href="http://reptile-database.reptarium.cz/advanced_search?taxon=Lamprophiidae&submit=Search">the family Lamprophiidae</a> for a large group of mostly African snakes (321 species) formerly classified as colubrids but actually more closely-related to elapids (<a href="https://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">more detail here</a> and <a href="http://www.hedgeslab.org/pubs/203.pdf">here</a>). They also found support for seven subfamilies of lamprophiids, of which only one, Lamprophiinae, concerns us today. <a href="http://reptile-database.reptarium.cz/advanced_search?taxon=Lamprophiinae&submit=Search">There are currently 78 species placed in Lamprophiinae</a><a href="#1" name="top1"><sup>1</sup></a>, of which 25 are or have been at some point commonly called "House Snakes" and/or <a href="http://reptile-database.reptarium.cz/advanced_search?common_name=Lamprophis&submit=Search">placed in the genus <i>Lamprophis</i></a>. Only one of these is <i>Boaedon </i>(formerly <i>Lamprophis</i>)<i> </i><i>fuliginosus</i>, but in order to understand it, we'll need to take a closer look at the others.</div>
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A great deal of clarity was gained from the taxonomic actions of <a href="https://s3.amazonaws.com/academia.edu.documents/3221432/2011_Kelly_et_al_Lamprohiidae_MPE.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1518528029&Signature=oG71252i9VyFmcOQhrjp2Vc2q5A%3D&response-content-disposition=inline%3B%20filename%3DKelly_C.M.R._Branch_W.R._Broadley_D.G._B.pdf">Chris Kelly & co-authors in 2011</a>, who split the species in the genus <i>Lamprophis </i>up into several genera, depending on their relationships to other genera of lamprophiines. Even this study was only able to include data on ~40% of the species of lamprophiine snakes, so it's probable that surprises and new discoveries still await us.<br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh1MHvEdqNTSZ-pIy74bUs-GlkK_XTvRypZGedMBWA4Dy66hgaHKiACVFjQEGIZEx3t9eW9xPOhhHmpru5tEYL6p2LchgsRxVJfizJWcreVfe05hU99G33kxZTCW9OWBzKuHJrewdQVclat/s1600/Inyoka+swazicus+Cliff+%2526+Suretha+Dorse_crop.png" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="246" data-original-width="410" height="192" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh1MHvEdqNTSZ-pIy74bUs-GlkK_XTvRypZGedMBWA4Dy66hgaHKiACVFjQEGIZEx3t9eW9xPOhhHmpru5tEYL6p2LchgsRxVJfizJWcreVfe05hU99G33kxZTCW9OWBzKuHJrewdQVclat/s320/Inyoka+swazicus+Cliff+%2526+Suretha+Dorse_crop.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Swazi Rock Snakes, <i>Inyoka swazicus</i>, are endemic to rocky<br />
outcrops in Swaziland and adjacent provinces of South Africa</td></tr>
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</div>
<a href="http://reptile-database.reptarium.cz/advanced_search?taxon=Lamprophiinae&submit=Search">There are currently 12 genera of lamprophiines</a>. Two of these, <i><a href="https://www.jstor.org/stable/1440964">Chamaelycus</a> </i>(4 species)<i> </i>and <i><a href="http://reptile-database.reptarium.cz/species?genus=Dendrolycus&species=elapoides&search_param=%28%28genus%3D%27Dendrolycus%27%29%29">Dendrolycus</a> </i>(1 species), have not been included in any molecular phylogenetic trees, so we're going to ignore them for now. The general relationships of the other 10 genera have been sketched out, and they're divided into two groups of roughly equal diversity. The first includes the African Wolf Snakes (<i>Lycophidion</i>; 20 species) and the African File Snakes (<i>Gonionotophis</i>, including the former genus <i>Mehelya</i>; 15 species), as well as two monotypic genera: <i>Hormonotus modestus </i>(Uganda House Snake or Yellow Forest Snake) and <i>Inyoka swazicus </i>(Swaziland House Snake or Swazi Rock Snake). Both of these were originally described as species of <i>Lamprophis</i>. <i>Hormonotus </i>left the genus in the 19th century, and <i>Inyoka </i>was created for <i>swazicus </i>by <a href="https://s3.amazonaws.com/academia.edu.documents/3221432/2011_Kelly_et_al_Lamprohiidae_MPE.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1518528029&Signature=oG71252i9VyFmcOQhrjp2Vc2q5A%3D&response-content-disposition=inline%3B%20filename%3DKelly_C.M.R._Branch_W.R._Broadley_D.G._B.pdf">Kelly et al. 2011</a> (it means ‘snake’ in the Nguni language group, the main language group in Swaziland). <a href="https://www.biodiversitylibrary.org/item/215769#page/289/mode/1up">When it was originally described in 1970</a>, <i>swazicus </i>was thought to be intermediate between <i>Lamprophis </i>and <i>Boaedon</i>, both of which were in use at the time, but it turns out that the resemblance is superficial and it's closely related to neither. That takes care of the first two of our 25 House Snake species, which aren't really House Snakes at all.</div>
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhVy29HfP8IVGVVN_NUG61rK-zsZMn1mch7aYM8-pjO1AzDfgMHyD0vG9yMrkSRrb-DFhNnSkYvzF-UYmK-SQVMkJlbY7mye-o9pIZwe5_gHE-KJckIaxEFF4x1ldXk3PEIpmnJwneWZuYa/s1600/Lycodonomorphus+inornatus+Cliff+%2526+Suretha+Dorse.png" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="522" data-original-width="788" height="211" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhVy29HfP8IVGVVN_NUG61rK-zsZMn1mch7aYM8-pjO1AzDfgMHyD0vG9yMrkSRrb-DFhNnSkYvzF-UYmK-SQVMkJlbY7mye-o9pIZwe5_gHE-KJckIaxEFF4x1ldXk3PEIpmnJwneWZuYa/s320/Lycodonomorphus+inornatus+Cliff+%2526+Suretha+Dorse.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">The Olive Water Snake, <i>Lycodonomorphus inornatus</i>,<br />
was formerly thought to be a <i>Lamprophis</i></td></tr>
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<div style="text-align: left;">
</div>
The second group of lamprophiines contains six genera. Three of these are rather small and pretty straightforward, if obscure: Ethiopian Mountain Snakes (<i><a href="http://reptile-database.reptarium.cz/advanced_search?genus=Pseudoboodon&submit=Search">Pseudoboodon</a></i>; 4 species), <a href="http://scienceblogs.com/tetrapodzoology/2010/09/13/everything-about-bothrolycus/">Günther's Black Snake (<i>Bothrolycus ater</i>)</a>, and Red-Black Striped Snakes (<i><a href="http://reptile-database.reptarium.cz/advanced_search?genus=Bothrophthalmus&submit=Search">Bothrophthalmus</a></i>; 2 species). None of these have ever been called House Snakes or placed in <i>Lamprophis</i><a href="#2" name="top2"><sup>2</sup></a>, and they are clearly morphologically distinct. A fourth genus, African Water Snakes (<i><a href="http://reptile-database.reptarium.cz/advanced_search?genus=Lycodonomorphus&submit=Search">Lycodonomorphus</a></i>; 9 species), includes two species that were formerly thought of as House Snakes: <i>Ly. inornatus </i>and <i>Ly. rufulus </i>(the second only briefly). <i>Ly. inornatus </i>is interesting because it's terrestrial, unlike the other species of <i>Lycodonomorphus</i>, which is part of why it was classified in <i>Lamprophis </i>for so long.</div>
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgTpILF32jSBZP3DZe2PGwi8X-7Zu7wrii-GXbXU_jZeOFovyjawJtXdiM298VvrzWd8hVTOPLA2YTJ-1yCQBdFZzzY7B9QvVw9T8qCAIVL1zG_zXQc9x3eeFfrNu7RnMzjaPQWlGXpdxE5/s1600/Lamprophis+fiskii+Cliff+%2526+Suretha+Dorse.png" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="534" data-original-width="797" height="214" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgTpILF32jSBZP3DZe2PGwi8X-7Zu7wrii-GXbXU_jZeOFovyjawJtXdiM298VvrzWd8hVTOPLA2YTJ-1yCQBdFZzzY7B9QvVw9T8qCAIVL1zG_zXQc9x3eeFfrNu7RnMzjaPQWlGXpdxE5/s320/Lamprophis+fiskii+Cliff+%2526+Suretha+Dorse.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Fisk's House Snake, <i>Lamprophis fiskii</i>, is found in<br />
rocky & sandy areas in the western part of South Africa</td></tr>
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</div>
The really important finding of <a href="https://s3.amazonaws.com/academia.edu.documents/3221432/2011_Kelly_et_al_Lamprohiidae_MPE.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1518528029&Signature=oG71252i9VyFmcOQhrjp2Vc2q5A%3D&response-content-disposition=inline%3B%20filename%3DKelly_C.M.R._Branch_W.R._Broadley_D.G._B.pdf">Kelly et al. 2011</a> was that <i>Lycodonomorphus </i>split up the remaining members of <i>Lamprophis </i>into two groups. The southern African group containing <i>Lamprophis aurora</i> got to keep the name <i>Lamprophis</i>, because <i>L. aurora </i>was the first species to be placed in <i>Lamprophis </i>(it is the "type species" of the genus). It got to bring along its close relatives <i>L. fiskii</i>, <i>L. fuscus</i>, and <i>L. guttatus</i>, all of which are small house snakes with attractive patterns, sometimes referred to as "dwarf house snakes", that are <a href="http://biodiversityadvisor.sanbi.org/wp-content/uploads/2015/11/Suricata_1_2014.pdf">popular in the pet trade despite being relatively poorly known in the wild</a>.</div>
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj-wds3ftg4Oc2gpY8G26jzwRS5bHWc0EfTmFADWNpwUsbGfel6RK_d_4J9W1RxAtWo1-tjO-Smq77qGVUh1T90Csk-O-GrZCq_ATSOlFMhyKXz4kRDTo5LO1cdJuwLXMAkwckS3ugyr-WQ/s1600/Boaedon+olivaceus+KM.jpg" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="960" data-original-width="951" height="200" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj-wds3ftg4Oc2gpY8G26jzwRS5bHWc0EfTmFADWNpwUsbGfel6RK_d_4J9W1RxAtWo1-tjO-Smq77qGVUh1T90Csk-O-GrZCq_ATSOlFMhyKXz4kRDTo5LO1cdJuwLXMAkwckS3ugyr-WQ/s200/Boaedon+olivaceus+KM.jpg" width="198" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Olive House Snakes, <i>Boaedon olivaceus</i><br />
are found in forests rather than savannah<br />
& grassland habitats</td></tr>
</tbody></table>
The other group needed a new name. Fortunately, <i><a href="http://reptile-database.reptarium.cz/advanced_search?genus=Boaedon&submit=Search">Boaedon</a> </i>had already been used to refer to this group for a long time, from the 1850s to the 1980s. Four species in Kelly's study got "new" names: <i>B. olivaceus</i>, <i>B. virgatus</i>, <i>B. lineatus</i>, and <i>B. fuliginosus</i>. Additionally, Kelly included <i>B. maculatus </i>in this group, because its morphology is similar to the other four species, but since we have no DNA evidence yet, this could change. These are sometimes informally called the "brown house snakes", in reference to their generally drabber patterns compared with the "dwarf house snakes". <a href="https://www.tandfonline.com/doi/abs/10.1080/04416651.1969.9650747?journalCode=ther19">Morphological differences</a> between these two genera include that <i>Boaedon </i>have enlarged anterior teeth on both the <a href="https://snakesarelong.blogspot.com/2018/02/basics-of-snake-skulls.html">upper & lower jaw</a>, and that the dorsal scales of <i>Boaedon </i>have apical pits, whereas those of <i>Lamprophis </i><a href="https://s3.amazonaws.com/academia.edu.documents/3221432/2011_Kelly_et_al_Lamprohiidae_MPE.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1518528029&Signature=oG71252i9VyFmcOQhrjp2Vc2q5A%3D&response-content-disposition=inline%3B%20filename%3DKelly_C.M.R._Branch_W.R._Broadley_D.G._B.pdf">do not</a>.</div>
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Three other species get to stick around in <i>Lamprophis</i> for now: <i>"L." abyssinicus </i>and <i>"L." erlangeri</i> from the Ethiopian highlands, and <i>"L." geometricus </i>from the Seychelles. Probably once we get genetic data from these they will be moved into another genus, possibly <i>Boaedon</i>.</div>
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhdY09_OQevCBS_ehT3GKE4SyQeptWy_jdh0EIj7x5khxZRLaO5rzmALg2rGmTEltjJNJO7iGWCfmF49pupdPxoTtI_lAVpBugd_d3Ci9syUZCiRPCOrdcgnnRS_pkPnGiOV2rIa7zepkx2/s1600/Boaedon+tree+Greenbaum+et+al+2015.png" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="579" data-original-width="461" height="400" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhdY09_OQevCBS_ehT3GKE4SyQeptWy_jdh0EIj7x5khxZRLaO5rzmALg2rGmTEltjJNJO7iGWCfmF49pupdPxoTtI_lAVpBugd_d3Ci9syUZCiRPCOrdcgnnRS_pkPnGiOV2rIa7zepkx2/s400/Boaedon+tree+Greenbaum+et+al+2015.png" width="317" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Most of the tree from <a href="https://journals.co.za/content/ajherp/64/1/EJC171369">Greenbaum et al. 2015</a>, showing<br />
the paraphyly of <i>B. fuliginosus </i>with respect to other<br />
<i>Boaedon </i>species, and the geographic diversity of the samples.</td></tr>
</tbody></table>
Now, the problems aren't over. The thing is that, in <a href="https://s3.amazonaws.com/academia.edu.documents/3221432/2011_Kelly_et_al_Lamprohiidae_MPE.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1518528029&Signature=oG71252i9VyFmcOQhrjp2Vc2q5A%3D&response-content-disposition=inline%3B%20filename%3DKelly_C.M.R._Branch_W.R._Broadley_D.G._B.pdf">Kelly's study</a>, <i>Boaedon "</i><i>fuliginosus</i><i>" </i>was split up by <i>B. olivaceus</i>, which is clearly a good species and it makes no sense to sink it into <i>fuliginosus</i>, as well as by <i>B. lineatus</i>, which has a more complex relationship with <i>B. "fuliginosus"</i><a href="#3" name="top3"><sup>3</sup></a>. There are <u>at least</u> seven lineages of <i>Boaedon "</i><i>fuliginosus</i><i>" </i>(probably more than 10), thus we can expect that at least 7-10 cryptic species are waiting to be described within this species complex. To quote <a href="https://s3.amazonaws.com/academia.edu.documents/3221432/2011_Kelly_et_al_Lamprohiidae_MPE.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1518528029&Signature=oG71252i9VyFmcOQhrjp2Vc2q5A%3D&response-content-disposition=inline%3B%20filename%3DKelly_C.M.R._Branch_W.R._Broadley_D.G._B.pdf">Kelly et al.</a>: "There have been several attempts to make sense of the intricate patterns of morphological variation in this complex, generally with only limited success."<a href="#4" name="top4"><sup>4</sup></a>. A handful of subspecies have been named based on morphology (e.g. <i><a href="http://www.lacerta.de/AS/Bibliografie/BIB_4908.pdf">mentalis</a> </i>in Namibia, <i><a href="http://www.lacerta.de/AS/Bibliografie/BIB_10537.pdf">angolensis</a> </i>from southeastern Angola to the southern DRC, <i><a href="https://www.biodiversitylibrary.org/item/128799#page/461/mode/1up">arabicus</a> </i>in Yemen, <i><a href="http://mapress.com/j/zt/article/view/zootaxa.4387.1.4">bedriagae </a></i>on the islands of São Tomé and Príncipe), some of which will probably eventually turn out to be used for full species.</div>
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<div style="text-align: justify;">
Which, if any, of these future species will get to keep the name <i>fuliginosus </i>is not clear, because these decisions are made based on the location of the original specimen, called the "type locality". The type locality for <i>L. fuliginosus</i> was originally and incorrectly reported in 1827 as "Java". <a href="https://snakesarelong.blogspot.de/2015/05/linnaean-snakes-part-i.html">People were more careless back then</a>. There is also no clear <a href="http://museum.wa.gov.au/explore/blogs/mark-harvey/what-type-specimen">type specimen</a>; at one point, one was designated, but it was lost by 1965. The type locality was subsequently corrected to the more accurate but still completely unhelpful "Africa" in 1962, and further restricted to either South Africa or Ghana, but which one isn't clear.<br />
<br />
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiZsqHqLud24SP0WK2CDAkhVlUxePtxqsKwjPyVwqE9ZwzjpRcQfECX3bUKW76oWD50tqHZ57zzJJjutaxBe-U4y2dAzYtcnqAyj7CEykJhLfy73KSa0JwSJKbHkgK1dz6QwIHU2_JCjU75/s1600/lamprophis.png" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1600" data-original-width="1397" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiZsqHqLud24SP0WK2CDAkhVlUxePtxqsKwjPyVwqE9ZwzjpRcQfECX3bUKW76oWD50tqHZ57zzJJjutaxBe-U4y2dAzYtcnqAyj7CEykJhLfy73KSa0JwSJKbHkgK1dz6QwIHU2_JCjU75/s320/lamprophis.png" width="279" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><div align="center" style="margin: 0in 0in 0.0001pt;">
<span style="font-size: x-small;">Map of the species currently in <i>Boaedon </i>& <i>Lamprophis</i><o:p></o:p></span></div>
<div align="center" style="margin: 0in 0in 0.0001pt;">
<span style="font-size: x-small;">Question marks indicate areas where the species range<o:p></o:p></span></div>
<div align="center" style="margin: 0in 0in 0.0001pt;">
<span style="font-size: x-small;">is uncertain (pink=<i>lineatus</i> complex, green=<i>olivaceus</i>,<o:p></o:p></span></div>
<div align="center" style="margin: 0in 0in 0.0001pt;">
<span style="font-size: x-small;">brown=<i>"fuliginosus"/"capensis" </i>complex)<o:p></o:p></span></div>
<div align="center" style="margin: 0in 0in 0.0001pt;">
<span style="font-size: x-small;"><a href="https://drive.google.com/file/d/173uOpphQhyat4PSbUgVGYGdoAYTLimis/view?usp=sharing">Click
here</a> for larger version</span><o:p></o:p></div>
</td></tr>
</tbody></table>
<div style="text-align: left;">
</div>
Finally, there is the issue of <i>Boaedon "capensis"</i>, a putative species <a href="http://www.tandfonline.com/doi/abs/10.1080/21564574.1997.9649980">described in 1997 by Hughes</a> and occurring east of a hazy and ill-defined zone angling northeast-southwest from the Gulf of Aden along the Great Rift Valley, then turning east and extending to the Atlantic Ocean possibly near the Angola-Namibia border, but potentially as far north as the mouth of the Congo River and thus also including three of the largest and most poorly-surveyed countries in Africa: Angola, the Democratic Republic of the Congo, and Sudan (including the still relatively new country of South Sudan). This name effectively replaces <i>fuliginosus </i>in eastern and southern Africa, but the exact boundaries are not remotely known, and it will probably turn out that both species are non-mutually-exclusive complexes of cryptic species. Because of the type locality confusion of <i>fuliginosus</i>, it could even turn out that both names (<i>fuliginosus </i>and <i>capensis</i>)<i> </i>are the same southern African species<a href="#5" name="top5"><sup>5</sup></a>, and that the western and central African species will need new names.</div>
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<br />
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjVUqFv-f2ot9wb8_SaEAuRfwrmf2U167QL4uU5vx0L1a0ko4JYzr-kSHu6yHgys8iKo1W1hoyvpPF84-g2EOTP7qSiPPqMLaigTMFWQRi8DjW5pIB9RJ1_nwN1SPrCSs0pfk8Z6uMfxFkV/s1600/Boaedon+radfordi+Greenbaum+et+al+2015+crop.png" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="315" data-original-width="442" height="228" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjVUqFv-f2ot9wb8_SaEAuRfwrmf2U167QL4uU5vx0L1a0ko4JYzr-kSHu6yHgys8iKo1W1hoyvpPF84-g2EOTP7qSiPPqMLaigTMFWQRi8DjW5pIB9RJ1_nwN1SPrCSs0pfk8Z6uMfxFkV/s320/Boaedon+radfordi+Greenbaum+et+al+2015+crop.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Boaedon radfordi</i>, a new species from the Uganda-DRC<br />
border region. From <a href="https://journals.co.za/content/ajherp/64/1/EJC171369">Greenbaum et al. 2015</a></td></tr>
</tbody></table>
Recent discoveries have begun the process of adding to the number of species of <i>Boaedon</i>: in 2015, <a href="https://journals.co.za/content/ajherp/64/1/EJC171369">Eli Greenbaum and colleagues</a> named a new species, <i>B. radfordi</i>, from the Albertine Rift in the eastern DRC and Uganda (which was formerly confused with <i>B. olivaceus</i>), and also unexpectedly found that a subspecies of <i>Lycodonomorphus subtaeniatus </i>was actually an undescribed species of <i>Boaedon </i>from a lake in south-central DRC, named <i>B.</i> <i>upembae</i>, that is most closely related to <i>B. virgatus</i>. They wisely refrained from making premature splits to the <i>fuliginosus</i>/<i>capensis </i>complex, stating that "Given the complicated taxonomic history and nebulous type locality for <i>B. fuliginosus</i>, substantial additional sampling and morphometric analyses are needed to assign...<i>B. fuliginosus</i> lineages to available names and to describe new species." They did, however, show that divergence among the various lineages currently referred to as <i>B. fuliginosus</i> could have happened as long as 21 million years ago.</div>
<div style="text-align: justify;">
<br />
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhv2kTSw8EsyFS5gGnlc8go0hKNSy_fTICIrddFtkT60-uftMHmIyaK9PD_mpr55j9GzUF1akKEBGL4GybJGDx5xGkfPZaDV7ybkncOeIeLW0XTPaEKOrfyh_AcMwBk9QTPf2tA1-4_k74c/s1600/Boaedon+longilineatus+Trape+%2526+Mediannikov+2016+fig27.png" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="526" data-original-width="865" height="193" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhv2kTSw8EsyFS5gGnlc8go0hKNSy_fTICIrddFtkT60-uftMHmIyaK9PD_mpr55j9GzUF1akKEBGL4GybJGDx5xGkfPZaDV7ybkncOeIeLW0XTPaEKOrfyh_AcMwBk9QTPf2tA1-4_k74c/s320/Boaedon+longilineatus+Trape+%2526+Mediannikov+2016+fig27.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Boaedon longilineatus</i>, a new species from Chad<br />
From <a href="http://www.documentation.ird.fr/hor/fdi:010069109">Trape & Mediannikov 2016</a></td></tr>
</tbody></table>
In 2016, <a href="http://www.documentation.ird.fr/hor/fdi:010069109">Trape & Mediannikov</a> examined 1,370 specimens from eight countries and described 5 new species of <i>Boaedon </i>from central Africa, <a href="http://reptile-database.reptarium.cz/advanced_search?genus=Boaedon&submit=Search">bringing the total number of species to 13 </a>(including <i>capensis </i>and the certainly paraphyletic "<i>fuliginosus</i>"). Together, two of these, <i>B. perisilvestris </i>and <i>B. subflavus</i>, seem to effectively separate <i>fuliginosus </i>(western Cameroon and west) and <i>capensis </i>(Angola-DRC-S.Sudan and east), having been split from the middle of the species complex's geographic range; but many sources still use <i>fuliginosus </i>for populations east of the distribution of <i>perisilvestris </i>and <i>subflavus</i>. Trape & Mediannikov seem comfortable with the idea of restricting <i>B. </i><i>fuliginosus </i>to West Africa, and <a href="http://reptile-database.reptarium.cz/species?genus=Boaedon&species=perisilvestris&search_param=%28%28genus%3D%27boaedon%27%29%29">suggest that a blackish color without clear lines on the head could distinguish the species there</a>, despite the absence of any consistent scale characteristics<a href="#6" name="top6"><sup>6</sup></a>. Right now, it's impossible to say how the 5 species described by Trape & Mediannikov fit with those described by Greenbaum or with the clades outlines in Kelly, because they used the 16S RNA gene, whereas the other two studies used three different genes (cyt-b, ND4, and c-mos).<br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjhuAE4SYzE5TY4eg5TRoMwslAhSLB578cMHvBBTjWnPjo3WNnEkpo7l6sYtvYxC8G8azHYpk6owhpQKNLJdXF-HKGZGnhCKyuc4ccY8pzS8JyHP3J3ZUzBJvQlnoBej23CuIxYpYqkF0DF/s1600/Boaedon+capensis+Cliff+%2526+Suretha+Dorse.png" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="532" data-original-width="798" height="213" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjhuAE4SYzE5TY4eg5TRoMwslAhSLB578cMHvBBTjWnPjo3WNnEkpo7l6sYtvYxC8G8azHYpk6owhpQKNLJdXF-HKGZGnhCKyuc4ccY8pzS8JyHP3J3ZUzBJvQlnoBej23CuIxYpYqkF0DF/s320/Boaedon+capensis+Cliff+%2526+Suretha+Dorse.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Boaedon capensis </i>from South Africa</td></tr>
</tbody></table>
So, we seem to be approaching stability, but the most problematic one remaining is the one everybody's heard of, knows and loves. Trape's latest definition notwithstanding, between <i>fuliginosus </i>and <i>capensis</i>, African House Snakes in the strictest sense occur in every country in Africa except for Algeria, Tunisia, Libya, Egypt, Sudan, and offshore countries like Madagascar, the Comoros, and the Seychelles<a href="#7" name="top7"><sup>7</sup></a>. At the moment, the <i>L. </i>"<i>fuliginosus</i>" complex is still one of the most widespread snake species in the world.<br />
<br />
<div style="text-align: justify;">
In case you lost count, a quick recap of species that are or have been in <i>Lamprophis</i>:</div>
<div style="text-align: justify;">
</div>
<ol style="text-align: start;">
<li><i>Hormonotus modestus </i>(Yellow Forest Snake or "Uganda House Snake"; moved in 1850s)</li>
<li><i>Inyoka swazicus </i>(Swazi Rock Snake or "Swaziland House Snake"; moved in 2011)</li>
<li><i>Pseudoboodon lemniscatus </i>(briefly in <i>Lamprophis </i>in 1904, barely counts, see footnote<a href="#2" name="top2"><sup>2</sup></a>)</li>
<li><i><i>Lycodonomorphus</i> inornatus </i>(originally described as a <i>Lamprophis </i>because it was terrestrial, but always a little weird; moved in 2011)</li>
<li><i><i>Lycodonomorphus </i>rufulus </i>(briefly in <i>Lamprophis </i>1840s-1860s, barely counts)</li>
<li><i>Lamprophis aurora </i>(type species for the genus, will always be a <i>Lamprophis </i>by definition)</li>
<li><i style="font-style: italic;">Lamprophis fiskii </i>(gets to stick with <i>aurora</i>)</li>
<li><i>Lamprophis fuscus </i>(gets to stick with <i>aurora</i>)</li>
<li><i>Lamprophis guttatus </i>(gets to stick with <i>aurora</i>)</li>
<li><i>"</i><i>Lamprophis" abyssinicus </i>(awaiting DNA data; Ethioipian highlands)</li>
<li><i>"</i><i>Lamprophis" erlangeri </i>(awaiting DNA data; Ethioipian highlands)</li>
<li><i>"</i><i>Lamprophis" geometricus </i>(awaiting DNA data; Seychelles)</li>
<li><i>Boaedon </i><i>lineatus </i>(type species for the genus, will always be a <i>Boaedon </i>by definition, although as defined it too is likely a cryptic species complex)</li>
<li><i style="font-style: italic;">Boaedon virgatus </i>(gets to stick with <i>lineatus</i>)</li>
<li><i>Boaedon olivaceus</i><i> </i>(gets to stick with <i>lineatus</i>)</li>
<li><i>Boaedon maculatus</i><i> </i>(awaiting DNA data; got to stick with the above 3 because of morphology; Horn of Africa)</li>
<li><i><i>Boaedon</i> radfordi</i><i> </i>(described by <a href="https://journals.co.za/content/ajherp/64/1/EJC171369">Greenbaum et al. 2015</a>, split from <i>olivaceus</i>)</li>
<li><i>Boaedon </i><i>upembae</i><i> </i>(formerly <i>Lycodonomorphus subtaeniatus </i><i>upembae</i>; moved by <a href="https://journals.co.za/content/ajherp/64/1/EJC171369">Greenbaum et al. 2015</a>; in the <i>B. </i><i>virgatus </i>group)</li>
<li><i>Boaedon littoralis</i><i> </i>(split from <i>B. </i><i>lineatus</i> by <a href="http://www.documentation.ird.fr/hor/fdi:010069109">Trape & Mediannikov 2016</a>, but lacks DNA data)</li>
<li><i>Boaedon longilineatus</i><i> </i>(split from <i>B. </i><i>lineatus</i> by Trape & Mediannikov 2016)</li>
<li><i>Boaedon paralineatus</i><i> </i>(split from <i>B. </i><i>lineatus</i> by Trape & Mediannikov 2016)</li>
<li><i>Boaedon perisilvestris</i><i> </i>(the first of many cryptic species to be split from <i>B. </i><i>fuliginosus</i>;<i> </i>by Trape & Mediannikov 2016)</li>
<li><i>Boaedon subflavus</i><i> </i>(the 2nd split from <i>B. </i><i>fuliginosus </i>by Trape & Mediannikov 2016)</li>
<li><i>Boaedon capensis</i><i> </i>(replaces <i>fuliginosus </i>in east Africa, could be multiple cryptic species)</li>
<li><i>Boaedon </i><i>fuliginosus </i>(definitely at least 7 cryptic species, probably many more, no guarantee that any will be called <i>fuliginosus</i>)</li>
</ol>
</div>
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</div>
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgJcuaQi9CegdiYhXE9LX8FlYjxmK29jincxXkiH1PpkdYFli1DZY2Ozn9g5hqJqW6wkpWmLUpPNm7uIG7uljogqjxF4tsU_7uyB7ivKyDply-K5Y1EtE-iCxOBSxiV-Eeu2A0VG0z3N8cD/s1600/Lamprophis+aurora+Cliff+%2526+Suretha+Dorse.png" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="496" data-original-width="757" height="209" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgJcuaQi9CegdiYhXE9LX8FlYjxmK29jincxXkiH1PpkdYFli1DZY2Ozn9g5hqJqW6wkpWmLUpPNm7uIG7uljogqjxF4tsU_7uyB7ivKyDply-K5Y1EtE-iCxOBSxiV-Eeu2A0VG0z3N8cD/s320/Lamprophis+aurora+Cliff+%2526+Suretha+Dorse.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">The Aurora House Snake, <i>Lamprophis aurora</i>, is the<br />
type species of the genus <i>Lamprophis</i>, meaning it will always<br />
be in <i>Lamprophis </i>unless that genus goes away completely</td></tr>
</tbody></table>
Whether <i>fuliginosus </i>goes away completely or remains, it won't be going back to <i>Lamprophis </i>unless <i>Lycodonomorphus </i>does too, or unless new <a href="https://snakesarelong.blogspot.com/2014/12/big-deal-snake-genomes.html">genomic data</a> overwhelm the signals found in the genes used by Kelly's, Greenbaum's, & Trape's studies. There's a recurring debate in taxonomy about whether we should attempt to preserve widely-used and well-known names like <i>fuliginosus</i>, since people are probably going to continue using them anyway, or do away with <a href="https://www.researchgate.net/profile/Frank_Burbrink/publication/228789496_Systematics_Of_The_Common_Kingsnake_Lampropeltis_Getula_Serpentes_Colubridae_And_The_Burden_Of_Heritage_In_Taxonomy/links/5523dcf60cf223eed3800086.pdf">"the burden of heritage"</a> and adhere strictly to a system that discards 150-year-old names if they prove inconvenient or impossible to keep, at the risk of creating confusion & resentment. Proponents of the second argue that eventually people won't remember the old names, and I think they're right: I was born in the 1980s and didn't realize that <i>Lamprophis </i><i>fuliginosus </i>was called <i>Boaedon </i>for 130 years beforehand; when I learned its name in ~1999, it was as <i>Lamprophis </i><i>fuliginosus </i>and that was that. These changes might seem radical, but whenever possible they reinstate older names, like <i>Boaedon</i>, the disuse of which might seem radical to an older generation.<br />
<br /></div>
<div style="text-align: justify;">
There's further debate about the utility of splitting up cryptic species complexes, especially if it makes it almost impossible to identify which species you're looking at by morphology alone. These same issues are recapitulated in the <a href="https://herpunit.wordpress.com/2016/10/05/the-ratsnake-mess-for-dummies/">North American ratsnake taxonomic "mess"</a>, North American <a href="https://www.ideals.illinois.edu/bitstream/handle/2142/25193/biochemicalevolu57high.pdf?sequ..">slimy salamanders</a>, <a href="https://snakesarelong.blogspot.com/2012/06/egg-eating-snakes.html">egg-eating snakes</a>, and in countless other species groups around the world. When I was writing this article, I thought more than once that I should just wait for a better time when it's all stabilized, but actually there's never a good time; we're always learning more. Ultimately, fleshing out and revising phylogenies and taxonomies will teach us a lot about biodiversity, evolution, and human nature. My advice is to try to be open-minded rather than bitter and ugly when discussing them. There is no "right" or "wrong", there are just rules we've (mostly) agreed to follow. It's an exciting time.<br />
<br />
If this group of snakes interests you, watch the labs of <a href="https://scholar.google.com/citations?user=hT9GyaEAAAAJ&hl=en">Christopher Kelly</a>, <a href="https://www.cenak.uni-hamburg.de/en/uebercenak/mitarbeiter/hallermann.html">Jakob Hallermann</a>, <a href="https://scholar.google.com/citations?user=_vVCy3AAAAAJ&hl=en">Aaron Bauer</a>, and <a href="https://www.researchgate.net/scientific-contributions/2082098204_Jean-Francois_Trape">Jean-François Trape</a> for future research that should make much of this article obsolete.</div>
<div style="text-align: justify;">
<br />
<i>Edit: In July 2020, writing in the in the African Journal of Herpetology, <a href="https://www.dropbox.com/s/isbjkisey04cjuw/Hallermann%20et%20al.%202020.pdf?dl=0">Hallermann et al.</a> provide the first DNA data for </i>geometricus<i>, proving as suspected that it belongs in </i>Boaedon<i>. They also revalidate </i>mentalis<i> and </i>angolensis<i> and describe three new species of the </i>fuliginosus<i> complex from Angola (named </i>bocagei<i>, </i>branchi<i>, and </i>fradei<i>). Their analysis continues to suggest numerous undescribed species elsewhere in Africa. They restrict </i>fuliginosus<i> to western Africa from Morocco to northern Angola, although it seems likely that it could be restricted further. They also provide a morphological key to the now nine species of </i>Boaedon<i> in Angola.</i>
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<i>Edit: In January 2021, again in the African Journal of Herpetology, <a href="https://www.researchgate.net/publication/348512728_Taxonomic_revision_of_the_Jita_snakes_Lamprophiidae_Boaedon_from_Sao_Tome_and_Principe_Gulf_of_Guinea_with_the_description_of_a_new_species">Ceríaco and colleagues</a> describe two new species from the islands of São Tomé (</i>Boaedon bedriagae<i>) and Príncipe (</i>Boaedon mendesi<i>) in the Gulf of Guinea off Africa's west coast, which were previously mostly considered a subspecies or synonym of either </i>fuliginosus<i> or </i>lineatus<i>. They also state that "Given the considerable number of names coined as subspecies of either </i>fuliginosus<i> or </i>lineatus<i>, the number of names currently considered as synonyms of these two species and the imprecise data about the type locality of the nominotypical forms, this revision may prove to be one of the most challenging taxonomic works of present-day African herpetology."</i></div><div style="text-align: justify;"><i><br /></i></div><div style="text-align: left;"><i>Edit: <a href="https://www.researchgate.net/profile/Arthur-Tiutenko/publication/364308879_Generic_affinities_of_African_house_snakes_revised_a_new_genus_for_Boodon_erlangeri_Serpentes_Elapoidea_Lamprophiidae_Lamprophiinae/links/6362296337878b3e877721d7/Generic-affinities-of-African-house-snakes-revised-a-new-genus-for-Boodon-erlangeri-Serpentes-Elapoidea-Lamprophiidae-Lamprophiinae.pdf" target="_blank">A new paper published in the journal Salamandra</a> on </i><span style="text-align: left;"><i>30 October 2022 led by <a href="https://www.researchgate.net/profile/Arthur-Tiutenko" target="_blank">Arthur Tiutenko</a> re-evaluated the </i></span><i>two Ethiopian highland endemics, </i>abyssinicus <i>and </i>erlangeri. <i>B</i><i>oth are removed from </i>Lamprophis<i> but neither belongs to </i>Boaedon. <i>Instead, </i>abyssinicus <i>is assigned to </i>Pseudoboodon <i>on the basis of morphology (still no DNA data are available). The first DNA data for </i>erlangeri<i> are published, from a specimen collected in 2016.</i> <i>Because it is more closely related to </i>Bothrophthalmus<i> and </i>Bothrolycus<i> than it is to other lamprophiines, </i>erlangeri<i> is assigned to a brand new genus, </i>Bofa<i> (an <a href="https://en.wikipedia.org/wiki/Oromo_language" target="_blank">Oromo</a> word for "snake")</i><i>. Color photos in life, CT scans of the skull, and detailed morphological comparisons with the other genera are graciously provided. They suggest "bofa(s)" or "Ethiopian forest snakes" as common names.</i> <i>In addition, </i>guttatus<i> is removed from </i>Lamprophis<i> and assigned to </i>Alopecion, <i>an "empty" genus originally used in the 1850s for </i>guttatus<i> and various other species, all of which have since ended up in other genera</i>. <i>I didn't see this last one coming, but it makes sense because </i>guttatus<i> </i><i>shares a common ancestor with</i><i> the other three </i>Lamprophis<i> (</i>aurora, fiskii,<i> and </i>fuscus<i>) less recently than those three share an ancestor with one another, and </i>guttatus <i>is a rocky habitat specialist that has morphological adaptations that allow it to </i><i>flatten the head to a great degree, in contrast to other </i>Lamprophis<i> that have relatively rounded heads. Admittedly, this is rather subjective, but the authors note that there might be cryptic species within </i>guttatus<i> that would perhaps benefit from being part of a genus that is morphologically, ecologically, and genetically distinct, however arbitrarily the lines between genera are often drawn.</i></div><div style="text-align: left;"><i><br /></i></div><div style="text-align: left;"><i>Edit: <a href="https://hal.science/hal-04048833/document" target="_blank">A new paper published 30 November 2022</a> in the bulletin of the French herpetological society described a new species of </i>Boaedon<i> from mountainous areas in Rwanda, Burundi, Uganda, Tanzania, and the DRC as </i>Boaedon montanus<i>, effectively splitting it from </i>fuliginosus/lineatus <i>in that area.</i></div><div><br /></div>
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<a href="https://www.blogger.com/null" name="1"><b>1 </b></a>Note the difference between the endings of the family ("-idae") and subfamily ("-inae") names.<a href="#top1"><sup>↩</sup></a><br />
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<a href="https://www.blogger.com/null" name="2"><b>2 </b></a>Except for <i>Pseudoboodon lemniscatus</i>, but that was only once, in 1904. It counts, but only in the same way as stuff you did once in college. This is complicated enough already.<a href="#top2"><sup>↩</sup></a><br />
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<a href="https://www.blogger.com/null" name="3"><b>3 </b></a>Sources differ on whether <i>B. lineatus </i>is distinct from <i>B. fuliginosis</i>, but it seems to be in western Africa (though both could be multiple cryptic species). Some resources use <i>B. lineatus </i>for house snakes with head stripes in e.g. Uganda, Ethiopia, and Sudan, but increasingly these are referred to as <i>B. capensis</i>. Characteristics used to distinguish <i>B. virgatus </i>& <i>B. olivaceus </i>from <i>B. fuliginosus/capensis/lineatus </i>include undivided subcaudal scales in <i>B. olivaceus</i> and only 23 dorsal scale rows in <i>B. virgatus</i>, as well as the fact that <i>B. virgatus </i>& <i>B. olivaceus </i>are found in forests whereas the others are savannah species.<a href="#top3"><sup>↩</sup></a><br />
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<a href="https://www.blogger.com/null" name="4"><b>4 </b></a>The presence or absence of head stripes has been used as a highly visible character, but ultimately this probably won't prove to be closely correlated with genetic variation (and it's complicated by the fact that some <i>Boaedon </i>populations have head stripes as juveniles but lose them as adults). This is also the case in North American ratsnakes, where former subspecies with radically different adult color patterns, like <i>E. o. rossalleni</i> and <i>E. o. quadrivittata </i><a href="https://herpunit.wordpress.com/2016/10/05/the-ratsnake-mess-for-dummies/">turned out to be so genetically similar to the more widespread black phenotype that they are now not recognized</a>. This is part of <a href="http://mapress.com/j/zt/article/download/zootaxa.4375.2.5/13578">a move away from the subspecies concept in general</a>, wherein many authors either synonymize subspecies with existing species as "mere variants" or elevate them to full species status using genetic data. I think we can expect this trend to continue with House Snakes.<a href="#top4"><sup>↩</sup></a><br />
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<a href="https://www.blogger.com/null" name="5"><b>5 </b></a>This could happen if South Africa is chosen as the type locality of <i>fuliginosus</i>, because the type locality of <i>capensis </i>is also in South Africa—if South Africa ultimately contains just one species from the <i>fuliginosus </i>complex, then it will get to keep the older name (<i>fuliginosus</i>), and other former members elsewhere should not use the name <i>capensis </i>in order to avoid further confusion. If the type locality of <i>fuliginosus </i>is chosen to be in Ghana instead, then the name will probably continue to be used in western Africa. Let us hope for the 2nd option.<a href="#top5"><sup>↩</sup></a><br />
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<a href="https://www.blogger.com/null" name="6"><b>6 </b></a>This isn't an identification guide, but if you want to see the scale characters for the different species, you can refer to the tables and descriptions in the <a href="https://s3.amazonaws.com/academia.edu.documents/3221432/2011_Kelly_et_al_Lamprohiidae_MPE.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1518528029&Signature=oG71252i9VyFmcOQhrjp2Vc2q5A%3D&response-content-disposition=inline%3B%20filename%3DKelly_C.M.R._Branch_W.R._Broadley_D.G._B.pdf">Kelly</a>, <a href="https://journals.co.za/content/ajherp/64/1/EJC171369">Greenbaum</a>, and <a href="http://www.documentation.ird.fr/hor/fdi:010069109">Trape</a> papers.<a href="#top6"><sup>↩</sup></a><br />
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<a href="https://www.blogger.com/null" name="7"><b>7 </b></a>"<i>B. </i><i>fuliginosus</i>" are <a href="https://www.biodiversitylibrary.org/item/128799#page/461/mode/1up">also found on the Arabian peninsula in Yemen</a>; this could be the most obvious future split if these are shown to be their own lineage, and several sources have already used the name <i>arabicus </i>for them, although just a few individuals are known and additional biological specimens from Yemen are hard to come by. <a href="http://mapress.com/j/zt/article/view/zootaxa.4387.1.4">A recent paper</a> used <i>bedriagae </i>as the name of a full species on the islands of São Tomé, with a new species being described from the neighboring island of Príncipe.<a href="#top7"><sup>↩</sup></a><br />
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ACKNOWLEDGMENTS</div>
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Thanks to <a href="http://reptile-database.reptarium.cz/advanced_search">Peter Uetz</a> for his advice on literature, and to <a href="https://www.flickr.com/photos/7929115@N04/sets/72157665775640148/">Konrad Mebert</a>, <a href="http://biodiversityfocused.co.za/about-us/">Cliff & Suretha Dorse</a>, and <a href="http://www.flickr.com/photos/cowyeow/">Dan Rosenberg</a> for the use of their photos.</div>
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REFERENCES</div>
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<span style="font-size: x-small;">For map references, see <a href="https://drive.google.com/file/d/173uOpphQhyat4PSbUgVGYGdoAYTLimis/view?usp=sharing">map inset</a></span><br />
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<span style="font-size: x-small;">Bates, M. F., W. Branch, A. Bauer, M. Burger, J. Marais, G. Alexander, and M. De Villiers. 2014. Atlas and red list of the reptiles of South Africa, Lesotho and Swaziland. South African National Biodiversity Institute <<a href="http://biodiversityadvisor.sanbi.org/wp-content/uploads/2015/11/Suricata_1_2014.pdf">full-text</a>></span><br />
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<span style="font-size: x-small;">Bogert, C. M. 1940. Herpetological results of the Vernay Angola Expedition. Part 1. Snakes, including an arrangement of African Colubridae. Bulletin of the American Museum of Natural History 77:1-107 <<a href="http://digitallibrary.amnh.org/handle/2246/335">link</a>></span><br />
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<span style="font-size: x-small;">Branch, W. R. 1984. The House Snakes of southern Africa (genus <i>Lamprophis</i>). Litteratura Serpentium 204:106-120 <<a href="http://www.snakesociety.nl/jaargangen/1984e/Litteratura%20Serpentium%204-3-4%20106-120%20Branch,%20The%20House%20snakes%20of%20Southern%20Africa,%20genus%20Lamprophis.pdf">link</a>></span><br />
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<span style="font-size: x-small;">Brassine, M. C., C. M. R. Kelly, N. P. Barker, and M. H. Villet. 2008. The phylogenetics of the <i>Lamprophis fuliginosus</i>/<i>capensis </i>species complex in southern Africa. Page 13 Proceedings of the 9th Conference of the Herpetological Association of Africa, Sterkfontein Dam, South Africa.</span><br />
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<span style="font-size: x-small;">Broadley, D. G. 1969. The African house snakes—How many genera? The Journal of the Herpetological Association of Africa 5:6-8 <<a href="https://www.tandfonline.com/doi/abs/10.1080/04416651.1969.9650747?journalCode=ther19">preview</a>></span><br />
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<span style="font-size: x-small;">Ceríaco, L. M., M. P. Marques, and A. M. Bauer. 2018. Miscellanea Herpetologica Sanctithomae, with a provisional checklist of the terrestrial herpetofauna of São Tomé, Príncipe and Annobon islands. Zootaxa 4387:91-108.</span><br />
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<span style="font-size: x-small;">Conradie, W., R. Bills, and W. Branch. 2016. The herpetofauna of the Cubango, Cuito, and lower Cuando river catchments of south-eastern Angola. Amphibian and Reptile Conservation 10:6-36 <<a href="http://www.lacerta.de/AS/Bibliografie/BIB_10537.pdf">full-text</a>></span><br />
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<span style="font-size: x-small;">de Witte, G. F. 1963. The colubrid snake genera <i>Chamaelycus </i>Boulenger and <i>Oophilositum </i>Parker. Copeia 1963:634-636 <<a href="https://www.jstor.org/stable/1440964">full-text</a>></span></div>
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<span style="font-size: x-small;">Greenbaum, E., F. Portillo, K. Jackson, and C. Kusamba. 2015. A phylogeny of Central African <i>Boaedon </i>(Serpentes: Lamprophiidae), with the description of a new cryptic species from the Albertine Rift. African Journal of Herpetology 64:18-38 <<a href="https://journals.co.za/content/ajherp/64/1/EJC171369">abstract</a>></span></div>
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<span style="font-size: x-small;">Hallermann, J. and A. Schmitz. 2007. First results on the taxonomy of the <i>Lamprophis fuliginosus </i>complex in Africa. 14th European Congress of Herpetology and SEH Ordinary General Meeting <<a href="http://webpages.icav.up.pt/pessoas/herpmeeting/Abstract%20book%2014ECH.pdf">abstract book</a>></span><br />
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<span style="font-size: x-small;">Hughes, B. 1997. <i>Dasypeltis scabra</i> and <i>Lamprophis fuliginosus</i> - two pan-African snakes in the Horn of Africa: a tribute to Don Broadley. African Journal of Herpetology 46:68-77 <<a href="http://www.tandfonline.com/doi/abs/10.1080/21564574.1997.9649980">abstract</a>></span></div>
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<span style="font-size: x-small;">Kelly, C. M. R., W. R. Branch, D. G. Broadley, N. P. Barker, and M. H. Villet. 2011. Molecular systematics of the African snake family Lamprophiidae Fitzinger, 1843 (Serpentes: Elapoidea), with particular focus on the genera <i>Lamprophis </i>Fitzinger 1843 and <i>Mehelya </i>Csiki 1903. Molecular Phylogenetics and Evolution 58:415-426 <<a href="https://s3.amazonaws.com/academia.edu.documents/3221432/2011_Kelly_et_al_Lamprohiidae_MPE.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1518528029&Signature=oG71252i9VyFmcOQhrjp2Vc2q5A%3D&response-content-disposition=inline%3B%20filename%3DKelly_C.M.R._Branch_W.R._Broadley_D.G._B.pdf">academia.edu</a>></span></div>
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<span style="font-size: x-small;">Pyron, R. A., F. T. Burbrink, G. R. Colli, A. N. M. de Oca, L. J. Vitt, C. A. Kuczynski, and J. J. Wiens. 2011. The phylogeny of advanced snakes (Colubroidea), with discovery of a new subfamily and comparison of support methods for likelihood trees. Molecular Phylogenetics and Evolution 58:329-342 <</span><a href="http://www.wienslab.com/Publications_files/Pyron_et_al_2011.pdf" style="font-size: small;">full-text</a><span style="font-size: x-small;">></span></div>
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<span style="font-size: x-small;">Roux-Estève, R. and J. Guibé. 1965. Contribution a l’étude du genre <i>Boaedon </i>(Serpentes, Colubridae). Bulletin du Muséum National d’Histoire Naturelle, Paris 36:761-774.</span></div>
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<span style="font-size: x-small;">Roux-Estève, R. and J. Guibé. 1965. Étude comparée de <i>Boaedon fuliginosus </i>(Boié) et <i>B. lineatus</i> D. et B. (Ophidiens). Bulletin de l’Institut Fondamental d’Afrique Noire, Dakar 27A:397-409.</span></div>
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<span style="font-size: x-small;">Schaefer, N. 1970. A new species of house snake from Swaziland, with notes on the status of the two genera <i>Lamprophis </i>and <i>Boaedon</i>. Annals of the Cape provincial Museums 8:205-208 <<a href="https://www.biodiversitylibrary.org/item/215769#page/289/mode/thumb">full-text from BHL</a>></span><br />
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<span style="font-size: x-small;">Schätti, B. 1989. Amphibians and reptiles from the Yemen Arab Republic and Djibouti. Revue Suisse de Zoologie 96:905-937 <<a href="https://www.biodiversitylibrary.org/item/128799#page/437/mode/1up">full-text from BHL</a>></span><br />
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<span style="font-size: x-small;">Thorpe, R. and C. McCarthy. 1978. A preliminary study, using multivariate analysis, of a species complex of African house snakes (<i>Boaedon fuliginosus</i>). Journal of Zoology 184:489-506 <<a href="http://onlinelibrary.wiley.com/doi/10.1111/j.1469-7998.1978.tb03303.x/full">abstract</a>></span></div>
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<span style="font-size: x-small;">Trape, J.-F. o. and O. Mediannikov. 2016. Cinq serpents nouveaux du genre <i>Boaedon </i>Duméril, Bibron & Duméril, 1854 (Serpentes : Lamprophiidae) en Afrique centrale. Bulletin de la Societe Herpetologique de France 159:61-111 <<a href="http://www.documentation.ird.fr/hor/fdi:010069109">abstract</a>></span></div>
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<span style="font-size: x-small;">Vidal, N., W. R. Branch, O. S. G. Pauwels, S. B. Hedges, D. Broadley, M. Wink, C. Cruaud, U. Joger, and Z. Nagy. 2008. Dissecting the major African snake radiation: a molecular phylogeny of the Lamprophiidae Fitzinger (Serpentes, Caenophidia). Zootaxa 1945:51-66 <<a href="http://www.hedgeslab.org/pubs/203.pdf">full-text</a>></span><br />
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<span style="font-size: x-small;">Visser, J. 1979. Notes on two rare house snakes – Part 1. <i>Lamprophis fiskii </i>Boulenger (1887) and <i>L.</i> <i>swazicus </i>Schaefer (1970). Journal of the Herpetological Association of Africa 19:10-13.</span><br />
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<span style="font-size: x-small;">Visser, J. 1979. Notes on two rare house snakes – Part 2: The generic status of <i>Lamprophis</i> <i>fiskii </i>Boulenger (1887) and <i>Lamprophis swazicus </i>Schaefer (1970). Journal of the Herpetological Association of Africa 21:31-37.</span><br />
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="https://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
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<br />Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com9Jena, Germany50.927054 11.58923720000007150.7669025 11.26651370000007 51.087205499999996 11.911960700000071tag:blogger.com,1999:blog-7443075087825368900.post-59405183816991083802018-02-12T13:54:00.000-07:002018-04-12T03:54:44.817-06:00Basics of snake skulls<div class="separator" style="clear: both; text-align: center;">
<span style="font-size: x-small;">This article will soon be available in Spanish!</span></div>
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Snakes have a lot more bones than we do, but they have a lot fewer types of bones. Aside from <a href="http://carlgans.org/bor-view/?borv=15&borpage=100">a few boas, pythons, pipesnakes, and blindsnakes with vestigial femurs</a>, most snakes just have a few hundred vertebrae with one pair of ribs each (except in the neck & tail), and a skull.</div>
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The snake skull is a remarkable structure. Snake skulls are highly kinetic, with a lot more moving parts than our skulls. Human skulls have just one movable part: the temporomandibular joint, which opens and closes your mouth. Snake skulls have many joints and moving parts; they can move the left and right sides of their jaws independently, as well as the outer (maxilla) and inner (palatine+pterygoid) parts of their upper jaws. Many bones that are tightly knit together in the skulls of most animals are loosely connected by stretchy ligaments in snakes, allowing them to <a href="https://www.youtube.com/watch?v=LLk4rsCNFFU">stretch their jaws over huge prey</a> (pardon the goofy music in the linked video). Contrary to the popular phrase, snakes cannot actually "unhinge" their jaws (Harry Greene explains this very well in <a href="https://www.youtube.com/watch?v=Mm9h6KE-ZOk">this video</a>).</div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgsF3kOrQUqTflt4KMQJyhWz0hC6bwvNq99Q3r9nJv8VwKF23XheBd7siahKu7Ol6heIjIjiO5RzlYKzNu04zdoznzSZGC8ALCCL2SbwKpdYcXxGfsawD0rAnf4sylZShosYi1FykeF5k1N/s1600/skull+side+general+alethinophidian+BiolReptil+Cundall+%2526+Irish+v20+Fig2-3a+p357.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="210" data-original-width="544" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgsF3kOrQUqTflt4KMQJyhWz0hC6bwvNq99Q3r9nJv8VwKF23XheBd7siahKu7Ol6heIjIjiO5RzlYKzNu04zdoznzSZGC8ALCCL2SbwKpdYcXxGfsawD0rAnf4sylZShosYi1FykeF5k1N/s1600/skull+side+general+alethinophidian+BiolReptil+Cundall+%2526+Irish+v20+Fig2-3a+p357.png" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">The right side of the skull of an <a href="https://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">alethinophidian</a> snake (nose pointing to the right).<br />
<span style="font-size: 12.8px;">Bones with teeth are the maxilla (mx), palatine (pal), pterygoid (pt), and dentary (d).</span><br />
<span style="font-size: 12.8px;">From </span><a href="http://carlgans.org/bor-view/?borv=20&borpage=355&borp=365" style="font-size: 12.8px;">Cundall & Irish 2008</a>. For a key to all abbreviations, click <a href="http://carlgans.org/bor-view/?borv=20&borpage=351&borp=358">here</a>.<br />
The bones or parts of bones that are shaded are not present in all snake species.</td></tr>
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Most snakes have teeth on four pairs of bones, two of which are the same as pairs of bones where humans do: the maxilla (most of our upper jaw) and the dentary (our lower jaw). In addition, almost all snakes have teeth on two bones that in humans form part of the roof of the mouth: the palatine and the pterygoid<a href="#1" name="top1"><sup>1</sup></a>, which are connected one in front of the other. This means that snakes have two upper jaws on each side: an outer one (the maxilla) and an inner one (the palatine+pterygoid). <a href="https://snakesarelong.blogspot.com/2013/09/basics-of-snake-fangs.html">If a snake has fangs</a>, they are always on the maxilla<a href="#2" name="top2"><sup>2</sup></a>. Some snakes, such as pythons, also have teeth on the premaxilla, where we humans have our incisors, although in most snakes the premaxilla is a part of the snout, has no teeth, and does not act as part of the jaws.</div>
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<tr><td><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjqYvy9F0Ic5Sv1nWj6STdcIuCBctL7JdKu0Xx4lp3YovO3C1k6zsoY6VPQgj6mN_OJhhzJYk46LAjpyS7h59JR1L1HsbG-1c6zyR6SLJz1lzwp0M_yjvrQ-MXNJgotZKCszHYLk8ba5K3S/s1600/skull+ventral+general+alethinophidian+BiolReptil+Cundall+%2526+Irish+v20+Fig2-3b+p357.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="210" data-original-width="540" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjqYvy9F0Ic5Sv1nWj6STdcIuCBctL7JdKu0Xx4lp3YovO3C1k6zsoY6VPQgj6mN_OJhhzJYk46LAjpyS7h59JR1L1HsbG-1c6zyR6SLJz1lzwp0M_yjvrQ-MXNJgotZKCszHYLk8ba5K3S/s1600/skull+ventral+general+alethinophidian+BiolReptil+Cundall+%2526+Irish+v20+Fig2-3b+p357.png" /></a></td></tr>
<tr><td class="tr-caption" style="font-size: 12.8px;">The right half of the skull of a snake, looking up from the bottom (nose pointing to the right).<br />
<span style="font-size: 12.8px;">Bones with teeth are the maxilla (mx), palatine (pal), pterygoid (pt), and dentary (d). The premaxilla (pmx) has no teeth.</span><br />
<span style="font-size: 12.8px;">From </span><a href="http://carlgans.org/bor-view/?borv=20&borpage=355&borp=365" style="font-size: 12.8px;">Cundall & Irish 2008</a>. For a key to all abbreviations, click <a href="http://carlgans.org/bor-view/?borv=20&borpage=351&borp=358" style="font-size: 12.8px;">here</a><span style="font-size: 12.8px;">.</span>The bones or parts of bones that are shaded are not present in all snake species.</td></tr>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgaJnpCiJQFGnC4zlhlhExpipqq9BoNUH72xMLKf6lcEhkBpjA5eNenjPAohX0wX0PQXDvARkJB0MBOI99VRnWYvmSJp5RPYXhu1EYj2_aXCN9OBNmGksLvHcmeL5Lyg0FN13eLQBiOvapr/s1600/Python+molurus+bite+Gibby_crop.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="253" data-original-width="170" height="200" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgaJnpCiJQFGnC4zlhlhExpipqq9BoNUH72xMLKf6lcEhkBpjA5eNenjPAohX0wX0PQXDvARkJB0MBOI99VRnWYvmSJp5RPYXhu1EYj2_aXCN9OBNmGksLvHcmeL5Lyg0FN13eLQBiOvapr/s200/Python+molurus+bite+Gibby_crop.jpg" width="134" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Tooth marks left by a<br />
python bite (upper jaw<br />
above, lower jaw below).</td></tr>
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You can sometimes see this pattern of tooth marks left behind when a non-venomous snake lets go after biting something, and in fact <a href="https://www.quora.com/What-can-be-done-in-case-of-snake-bites-before-going-to-hospital-How-does-one-recognize-venomous-snakes-and-stop-the-venom-from-spreading-How-does-one-prevent-pain">many resources</a> suggest that you can use the tooth pattern to determine<a href="#3" name="top3"><sup>3</sup></a> whether or not a bite has come from a venomous snake (a viper at least, <a href="https://www.aafp.org/afp/2002/0401/p1367.html">which are responsible for >99% of venomous snakebites in the USA</a>), since most dangerously venomous snakes have different tooth patterns on account of their fangs, and <a href="https://eaglemountainpublishing.s3.amazonaws.com/PDF/Biology%20of%20the%20Vipers/CH%2013_hayes_.pdf">most of their non-fang teeth don't usually come into contact with the target</a>. I mentioned above that fangs are always on the maxilla, and that's because the maxilla is the primary prey-catching bone in the snake skull. As far as we know, <a href="http://people.whitman.edu/~jacksok/nature.pdf">fangs evolved only once</a>, as enlarged teeth at the back of the maxilla in the ancestor of all living colubroid snakes about 75 million years ago. In many living species of snakes, this is still the situation, and <a href="https://snakesarelong.blogspot.com/2017/08/how-many-snakes-are-venomous-constrictors.html">the vast majority of these are not dangerous to humans</a> (although some can inflict painful bites if allowed to chew for a few minutes, and <a href="http://www.jstor.org/stable/3892025?seq=1#page_scan_tab_contents">a few can be deadly</a>). In at least three cases (vipers, elapids, and <a href="https://snakesarelong.blogspot.de/2012/08/stiletto-snakes.html">atractaspidids</a>), the fangs have moved up to the front of the maxilla, through the developmental suppression of the front part of the maxilla (and its teeth) in the snake embryo. I covered this and the <a href="http://people.whitman.edu/~jacksok/JMorph.pdf">evolution of grooved and hollow fangs</a> in more detail in <a href="https://snakesarelong.blogspot.com/2013/09/basics-of-snake-fangs.html">my article about snake fangs</a>.</div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiyOKDUa4HB1Nl2t6fTfI0n8-hzJbXBNC6xTWXoBJgLDssUR4fIlMU_KBp4uUTLr7ZgZBODcy3uHYO_MTbLIjq-_UZYaO-3cvCmZxGFU4PNgsjf_-Tyd9nB2w3i37d3-K0QdUDAKujwGP_u/s1600/skull+dorsal+general+alethinophidian+BiolReptil+Cundall+%2526+Irish+v20+Fig2-3c+p357.png" imageanchor="1" style="margin-left: auto; margin-right: auto; text-align: center;"><img border="0" data-original-height="196" data-original-width="542" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiyOKDUa4HB1Nl2t6fTfI0n8-hzJbXBNC6xTWXoBJgLDssUR4fIlMU_KBp4uUTLr7ZgZBODcy3uHYO_MTbLIjq-_UZYaO-3cvCmZxGFU4PNgsjf_-Tyd9nB2w3i37d3-K0QdUDAKujwGP_u/s1600/skull+dorsal+general+alethinophidian+BiolReptil+Cundall+%2526+Irish+v20+Fig2-3c+p357.png" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">The right half of the skull of a snake, looking down from the top (nose pointing to the right).<br />
<span style="font-size: 12.8px;">No teeth are visible. </span><span style="font-size: 12.8px;">From </span><a href="http://carlgans.org/bor-view/?borv=20&borpage=355&borp=365" style="font-size: 12.8px;">Cundall & Irish 2008</a>. For a key to all abbreviations, click <a href="http://carlgans.org/bor-view/?borv=20&borpage=351&borp=358" style="font-size: 12.8px;">here</a><span style="font-size: 12.8px;">.</span>The bones or parts of bones that are shaded are not present in all snake species.</td></tr>
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Although most people are most interested in the teeth and fangs, the rest of the snake skull is no less fascinating. The outer and inner upper jaw are connected by a toothless upper jaw bone called the ectopterygoid, <a href="https://watermark.silverchair.com/j.1095-8312.1986.tb01730.x.pdf?token=AQECAHi208BE49Ooan9kkhW_Ercy7Dm3ZL_9Cf3qfKAc485ysgAAAdEwggHNBgkqhkiG9w0BBwagggG-MIIBugIBADCCAbMGCSqGSIb3DQEHATAeBglghkgBZQMEAS4wEQQMQFDGCH1RbvO3cKoKAgEQgIIBhLbK_DYu7EqZzwUJ-QmgqG9MEoKGs0v2wfrwhk0nQkKgkisAEbBLXl_2CxKdWZVMpoH1q1dcIQZz4F-kLZKpHyApgw0T03YCEXOuQBKqvkfril2GOlwSO_9zsFBlmOxUubqG0RQEtbIWbgNmcxa_ABKL6OFeWT05JZw8qcqpRJYvmLODJwkzPlGIMWMZKGWdv9tHuuhQIqP_MhqOnOKe0EsIQl6K8sPXWeToIFKziVA0xl5VWxzIaK5tFIUC0oxxzxK-RFlOD4S2WL-bo8Kmybc1LB1C4Nbs9mAOuNkyEY5Jezcq8Yu50DNcMqZ6rKuNP07juTaeB0gEg2oqBbk--NvrtQlGCWdb6crnyXu_F0tU2Mmyr-hznMjFcbIKFiSH12Y5BRrRRQOxA1d7c1hvvXucPqWHsXb1V32o_8q92uBviHiOshhzIT9O7gFU-5NiHu4Nhaq4xeTJwk82GWXi8JXelQbTDe3TNL2EsWM76aX1TknUa9qa2P0ZlIKP3qJgpC8c07s">which works like a lever to transfer muscular power from the muscles attached to the pterygoid out to the maxilla, which has no muscles of its own</a>. When a snake is eating, the entire upper jaw (inner and outer parts) is raised and moved slightly backward, alternating the left and right sides and pulling the prey into the mouth: <a href="https://www.youtube.com/watch?v=Mm9h6KE-ZOk">the characteristic "jaw-walking" or "pterygoid walk" motion of feeding snakes</a>. So, the front of the pterygoid is attached to the back of the palatine, the ectopterygoid hangs off the outside of the pterygoid, and the maxilla hangs off of the other end of the ectopterygoid. In vipers, whose fangs fold, the maxilla and its fang are pushed forward by the ectopterygoid and pterygoid.<br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjbyJoeeQi60-iR1B6TK7PTN0Db3_o563qhccfz4Kf1BJn-u5Icm4Nd8GhePz-H3H3bvZ4Oe9GHFEAVDjHLxoCta53yL5PWjv_6zVWLm0t24jXnkkyPF0J4C6l1E98IsF3kF16SCg1bxYFa/s1600/Kardong+1977+viper+fang+erection+apparatus+wo+caption.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="292" data-original-width="868" height="134" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjbyJoeeQi60-iR1B6TK7PTN0Db3_o563qhccfz4Kf1BJn-u5Icm4Nd8GhePz-H3H3bvZ4Oe9GHFEAVDjHLxoCta53yL5PWjv_6zVWLm0t24jXnkkyPF0J4C6l1E98IsF3kF16SCg1bxYFa/s400/Kardong+1977+viper+fang+erection+apparatus+wo+caption.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><div style="text-align: center;">
<span style="font-size: x-small;">Roughly the same fang movements are made during striking and swallowing. Supratemporal (st), quadrate (q), mandible (ma), pterygoid (pt), ectopterygoid (ec), palatine (pa), prefrontal (pf), maxilla (mx). From <a href="http://www.jstor.org/stable/1443913?casa_token=o12zxT2o9hgAAAAA:pSyTOZiTHZVeMisGWf5ms_3Q0n5sABlpieuOH76POFRsyQftfSPpG2BckbNiBTpf_9AKOxpHmn4ivdSyTbgNtC7-khgLdNiuwrG9kqe5HjOBRuTgxv0&seq=1#page_scan_tab_contents">Kardong 1977</a></span></div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhv052jOdrF01BGbs7xIuNS4JU4_T1rdC0D6YnJIAq6hCpe5VIdN1QE0oIUL6MGvD0kFe9xrQ2pLQlISEGJnW7YZbWHSbtQZ0AEdbdL2y-9_9nmDqHXX6e8icIvaPc58b2WCwN2T4ZFRt-X/s1600/Kardong+1977+viper+pterygoid+walk+Fig4.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1012" data-original-width="576" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhv052jOdrF01BGbs7xIuNS4JU4_T1rdC0D6YnJIAq6hCpe5VIdN1QE0oIUL6MGvD0kFe9xrQ2pLQlISEGJnW7YZbWHSbtQZ0AEdbdL2y-9_9nmDqHXX6e8icIvaPc58b2WCwN2T4ZFRt-X/s320/Kardong+1977+viper+pterygoid+walk+Fig4.png" width="182" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">The independent left and right movement<br />
of the upper jaws of a viper.<br />
Abbreviations as above. From <a href="http://www.jstor.org/stable/1443913?casa_token=o12zxT2o9hgAAAAA:pSyTOZiTHZVeMisGWf5ms_3Q0n5sABlpieuOH76POFRsyQftfSPpG2BckbNiBTpf_9AKOxpHmn4ivdSyTbgNtC7-khgLdNiuwrG9kqe5HjOBRuTgxv0&seq=1#page_scan_tab_contents">Kardong 1977</a>.</td></tr>
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Amazingly, in most snakes there is no direct connection between the upper jaws and the braincase<a href="#4" name="top4"><sup>4</sup></a>. Instead, the palatine and maxilla are connected to the braincase by long ligaments, which give them great freedom of motion. The front end of the palatine is connected more firmly to the snout, albeit still with some freedom to move. The rear end of the maxilla is also connected by a ligament to the lower jaw. It's really the movements of the palatine and pterygoid that swallow the prey. The lower jaws mainly press the prey against the upper jaws, and the teeth on the dentary and maxilla rarely contact the prey and play little active role in swallowing.<br />
<br />
The lower jaws or mandible participate in the process of feeding as well, and unlike in humans they have a loose attachment of the lower jaws to each other at the front of the dentary bones. The dentary bones are connected firmly at the back to the articular bones, which are connected to the quadrate bones at a flexible joint, which are connected to the back of the braincase by the supratemporal bones, also at a somewhat moveable joint. Together with the flexible palato-maxillary apparatus ("upper jaws"), this three-part lower jaw allows snakes to open their mouths very wide, walk their heads over, and consume things that are as big as they are without breaking them into smaller pieces or using their non-existent hands. The quadrate also attaches to the <a href="https://snakesarelong.blogspot.com/2015/09/can-snakes-hear.html">columella, which is the sole inner ear bone in reptiles; thus, the lower jaw also conducts sound to the ear</a>.<br />
<br />
So there you have it. The snake skull is divided into four functional units: the braincase, the snout, the palato-maxillary apparatus ("upper jaws") and the mandibular apparatus ("lower jaws"), each of which can move independently (well, except for the braincase, which is relatively stationary). The upper jaws are divided into two partially separated structural-functional units, a medial swallowing unit and a lateral prey capture unit, both of which work with the lower jaws to accomplish their tasks.<br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhOscAEpgkPwI1zzf2ktfGfcPHIRGrOz9QblX9-iX6MlYdjHwc3AmLmNCqr0Qh7jv6tqRiiRPgU74ktzZg8p1XyILZnCwCsAD6OhgDYncf2UCXzjOhS8NrJDpnJ-StqIWcyI37x96dq46iu/s1600/Casarea+skull+Frazzetta+1970.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="284" data-original-width="444" height="204" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhOscAEpgkPwI1zzf2ktfGfcPHIRGrOz9QblX9-iX6MlYdjHwc3AmLmNCqr0Qh7jv6tqRiiRPgU74ktzZg8p1XyILZnCwCsAD6OhgDYncf2UCXzjOhS8NrJDpnJ-StqIWcyI37x96dq46iu/s320/Casarea+skull+Frazzetta+1970.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">From <a href="https://www.jstor.org/stable/2459073">Frazzetta 1970</a>. <a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg3PKaeumzxEofrqUqfu0juZTQQCnEcTHevQwmG7BcELuNWQneE5lSIcxi0UVdTis8L4BLaAcDG4_WBwRNfcLdMzRL67X4qo_Vxi-xgtHKabAhxv0dj1FzONWpUxELcqipb7804PfEf9dRg/s1600/Fullscreen+capture+4292012+95849+AM.bmp.jpg">Click for larger size</a>.</td></tr>
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A quick note about a special case: one of my favorite snakes, <a href="https://snakesarelong.blogspot.de/2012/04/round-island-splitjaw-snakes.html">and one of the first I wrote about on this blog</a>, <i>Casarea dussumeri</i>, are often called Round Island boas, although I chose to use the more apt "splitjaw snakes" in my article. As if the usual kinesis of the snake skull isn't enough, <a href="http://digimorph.org/specimens/Casarea_dussumieri/">these snakes have a maxilla that is uniquely subdivided into two movable parts</a>, called the anterior and posterior maxilla. The anterior maxilla has 10 teeth and the posterior maxilla has 12. It is thought that the divided maxilla evolved through incomplete development, because the maxilla of other snakes forms in two parts before fusing together in the embryo, and the function is thought to be to help <i>Casarea </i>encircle hard, cylindrical prey such as skinks.</div>
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We still have a lot left to learn about snake skulls. We didn't even cover half of the bones in this article. You don't actually so much <a href="http://insider.si.edu/2014/05/intern-helped-discover-fossil-giant-snake-titanoboa/">find snake skulls</a> as you do carefully prepare them. The individual bones are so small and light and fragile that they tend not to fossilize well, nor can they easily be found among the other bones of a snake's skeleton. Even normal cleaning and preparation methods can damage the fragile bones of tiny snake skulls. Thus, there is much left to discover about how they work!<br />
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<tr><td class="tr-caption" style="text-align: center;"><i style="font-size: 12.8px;">Skull of Natrix natrix </i><span style="font-size: 12.8px;">from </span><a href="https://zslpublications.onlinelibrary.wiley.com/doi/full/10.1111/jzo.12477" style="font-size: 12.8px;">Andjelković et al. 2017</a><span style="font-size: 12.8px;">. Mobile connections marked with red dashed arrows and circles.</span><span style="font-size: 12.8px;">Paired bones are shown in yellow (pa – palatine, pt – pterygoid, ec – ectopterygoid, mx – maxilla, st </span><span style="font-size: 12.8px;">– supratemporal,</span><span style="font-size: 12.8px;">q – quadrate, cp – compound bone, d – dentary, pf – prefrontal), unpaired bones are shown in green or grey (pmx – premaxilla, na – nasal, b – braincase, smx – septomaxillae & vomers).</span></td></tr>
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<a href="https://www.blogger.com/null" name="1"><b>1 </b></a>Although the pterygoids are stand-alone bones in the roof of the mouth of many vertebrates, in humans they are called the pterygoid processes of the sphenoid bone because they are fused to the sphenoid bone.<a href="#top1"><sup>↩</sup></a><br />
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<a href="https://www.blogger.com/null" name="2"><b>2 </b></a>There is one very strange snake, <i>Pythonodipsas carinata</i> from Africa, that has <a href="http://www.jstor.org/tc/accept?origin=/stable/pdf/1444655.pdf">an ungrooved fang on the palatine bone</a>. They aren't any studies of their functional morphology so we don't really know exactly how they use their palatine fangs, but they <a href="https://snakesarelong.blogspot.com/2017/08/how-many-snakes-are-venomous-constrictors.html">use constriction to subdue their prey.</a><a href="#top2"><sup>↩</sup></a><br />
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<a href="https://www.blogger.com/null" name="3"><b>3 </b></a>I don't necessarily recommend this, partly because if you've been bitten then it's too late, and partly because it's better just to learn the few venomous snake species that live in your area than it is to try to follow some "rule" that inevitably has exceptions.<a href="#top3"><sup>↩</sup></a><br />
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<a href="https://www.blogger.com/null" name="4"><b>4 </b></a><a href="https://snakesarelong.blogspot.com/2012/08/stiletto-snakes.html">Atractaspidids</a> have a ball-and-socket joint between the prefrontal (part of the braincase) and the maxilla, which along with a gap, bridged by a ligament, between the pterygoid and palatine, allows them to <a href="http://www.jstor.org/stable/pdf/1445406.pdf">"strike" with their fang backwards</a>, with a closed mouth, using just the fang on one side, a useful if terrifying adaptation for envenomating prey in underground burrows. A hook-like ridge on the fang increases the size of the wound, presumably enhancing the absorption of venom.<a href="#top4"><sup>↩</sup></a><br />
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ACKNOWLEDGMENTS<br />
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Thanks to <a href="http://www.reptileforums.co.uk/forums/snakes/326960-most-painful-snake-bite-comparing-11.html">gibby</a> for the use of their photograph.</div>
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REFERENCES</div>
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<span style="font-size: x-small;">Albright, R. G. and E. M. Nelson. 1959. Cranial kinetics of the generalized colubrid snake <i>Elaphe obsoleta quadrivittata</i>. I. Descriptive morphology. Journal of Morphology 105:193-239.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Albright, R. G. and E. M. Nelson. 1959. Cranial kinetics of the generalized colubrid snake <i>Elaphe obsoleta quadrivittata</i>. II. Functional morphology. Journal of Morphology 105:241-291.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Andjelković, M., Tomović, L., & Ivanović, A. 2017. Morphological integration of the kinetic skull in <i>Natrix </i>snakes. Journal of Zoology, 303:188-198 <<a href="https://zslpublications.onlinelibrary.wiley.com/doi/full/10.1111/jzo.12477">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Cundall, D. 1983. Activity of head muscles during feeding by snakes: a comparative study. American Zoologist 23:383-396.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Cundall, D. and H. W. Greene. 2000. Feeding in snakes. Pages 293–333 in K. Schwenk, editor. Feeding: Form, Function, and Evolution in Tetrapod Vertebrates. Academic Press, San Diego, CA.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Cundall, D. and F. Irish. 2008. The snake skull. Pages 349-692 in C. Gans, A. S. Gaunt, and K. Adler, editors. Biology of the Reptilia. Volume 20, Morphology H. The Skull of Lepidosauria. The University Of Chicago Press, Chicago, Illinois, USA <<a href="http://carlgans.org/bor-view/?borv=20&borpage=349">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Frazzetta. T. 1970. From hopeful monsters to bolyerine snakes? The American Naturalist 104:55-72 <<a href="https://www.jstor.org/stable/2459073">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Frazzetta, T. 1971. Notes upon the jaw musculature of the Bolyerine snake, <i>Casarea dussumieri</i>. Journal of Herpetology 5:61-63</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Irish, F. and P. Alberch. 1989. Heterochrony in the evolution of bolyeriid snakes. Fortschritte der Zoolologie 35:205.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Juckett, G. and J. G. Hancox. 2002. Venomous snakebites in the United States: management review and update. American Family Physician 65:1367-1375 <<a href="https://www.aafp.org/afp/2002/0401/p1367.html">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Kardong, K. 1974. Kinesis of the jaw apparatus during the strike in the cottonmouth snake, <i>Agkistrodon piscivorus</i>. Forma et functio 7:327-354.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Kardong, K. V. 1977. Kinesis of the jaw apparatus during swallowing in the cottonmouth snake, <i>Agkistrodon piscivorus</i>. Copeia 1977:338-348 <<a href="http://www.jstor.org/stable/1443913?casa_token=o12zxT2o9hgAAAAA:pSyTOZiTHZVeMisGWf5ms_3Q0n5sABlpieuOH76POFRsyQftfSPpG2BckbNiBTpf_9AKOxpHmn4ivdSyTbgNtC7-khgLdNiuwrG9kqe5HjOBRuTgxv0&seq=1#page_scan_tab_contents">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Lombard, R. E., H. Marx, and G. B. Rabb. 1986. Morphometrics of the ectopterygoid in advanced snakes (Colubroidea): a concordance of shape and ph</span><span style="font-size: x-small;">ylogeny. Biological Journal of the Linnean Society 27:133-164 <</span><a href="https://watermark.silverchair.com/j.1095-8312.1986.tb01730.x.pdf?token=AQECAHi208BE49Ooan9kkhW_Ercy7Dm3ZL_9Cf3qfKAc485ysgAAAdEwggHNBgkqhkiG9w0BBwagggG-MIIBugIBADCCAbMGCSqGSIb3DQEHATAeBglghkgBZQMEAS4wEQQMQFDGCH1RbvO3cKoKAgEQgIIBhLbK_DYu7EqZzwUJ-QmgqG9MEoKGs0v2wfrwhk0nQkKgkisAEbBLXl_2CxKdWZVMpoH1q1dcIQZz4F-kLZKpHyApgw0T03YCEXOuQBKqvkfril2GOlwSO_9zsFBlmOxUubqG0RQEtbIWbgNmcxa_ABKL6OFeWT05JZw8qcqpRJYvmLODJwkzPlGIMWMZKGWdv9tHuuhQIqP_MhqOnOKe0EsIQl6K8sPXWeToIFKziVA0xl5VWxzIaK5tFIUC0oxxzxK-RFlOD4S2WL-bo8Kmybc1LB1C4Nbs9mAOuNkyEY5Jezcq8Yu50DNcMqZ6rKuNP07juTaeB0gEg2oqBbk--NvrtQlGCWdb6crnyXu_F0tU2Mmyr-hznMjFcbIKFiSH12Y5BRrRRQOxA1d7c1hvvXucPqWHsXb1V32o_8q92uBviHiOshhzIT9O7gFU-5NiHu4Nhaq4xeTJwk82GWXi8JXelQbTDe3TNL2EsWM76aX1TknUa9qa2P0ZlIKP3qJgpC8c07s" style="font-size: small;">link</a><span style="font-size: x-small;">></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Maisano, J. A. and O. Rieppel. 2007. The skull of the Round Island boa, <i>Casarea dussumieri </i>Schlegel, based on high-resolution X-ray computed tomography. Journal of Morphology 268:371-384 <<a href="https://www.ncbi.nlm.nih.gov/pubmed/17309080">abstract</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Raynaud, A. 1985. Development of Limbs and Embryonic Limb Reduction. Pages 59-148 in C. Gans and F. Billett, editors. Biology of the Reptilia. Volume 15. Development B. John Wiley & Sons, New York <<a href="http://carlgans.org/bor-view/?borv=15&borp=8">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Rieppel, O. 2012. “Regressed” Macrostomatan Snakes. Fieldiana Life and Earth Sciences 5:99-103 <<a href="http://www.bioone.org/doi/abs/10.3158/2158-5520-5.1.99">link</a>></span><br />
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="https://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com6Jena, Germany50.927054 11.58923720000007150.7669025 11.26651370000007 51.087205499999996 11.911960700000071tag:blogger.com,1999:blog-7443075087825368900.post-74512040454400444562017-12-25T20:46:00.001-07:002019-01-26T06:37:05.506-07:00Life is Short but Snakes are Long 2017 Year in Review<div style="text-align: justify;">
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<a href="https://drive.google.com/file/d/1wXSAR-yFSroX6zJCGEA38TQVJghnwgeV/view?usp=sharing"><span style="font-size: x-small;">Haga clic aquí para leer en español</span></a></div>
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Dear reader,</div>
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<i>Life is Short but Snakes are Long</i> turned 5 years old in April, and reached 1 million views on June 26th, 2017. I celebrated with a month-long recapitulation of the "best-of" on Twitter in the spring, and by taking a much-needed break from writing new content during the second half of the year. I didn't make a formal announcement of this break because I honestly wasn't sure how long it would last (although I knew it wasn't permanent). During the past six months I've focused on settling in to my new job in Germany, traveling around Europe, publishing a key article from my dissertation, and applying to postdocs and "real" jobs. I'm currently waiting to hear whether I'll be starting something new in 2018—if I am, I plan to continue to write <i>Life is Short but Snakes are Long </i>as I have been. If I'm not, I plan to begin revising certain past articles, and writing new ones, for a book. In either case, I'm looking forward to a lot of new content in 2018.</div>
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As always, thank you for reading.</div>
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Andrew Durso</div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhRrpmUPnRTywBoCTVBn15vpbuWwqtURqVPdtfGwHwKAUR3O8EeJXscIE_caGu28BFo_mY0PMwV30PuPnNlFqBQIzvPPmyutnl-ammQw4sJsPnBg6ARbWTeUQA0aSWFgJJRqtiAchHm2qS-/s1600/20170409_IMG_2743.JPG" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="769" data-original-width="1024" height="300" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhRrpmUPnRTywBoCTVBn15vpbuWwqtURqVPdtfGwHwKAUR3O8EeJXscIE_caGu28BFo_mY0PMwV30PuPnNlFqBQIzvPPmyutnl-ammQw4sJsPnBg6ARbWTeUQA0aSWFgJJRqtiAchHm2qS-/s400/20170409_IMG_2743.JPG" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Vipera berus </i>from Germany</td></tr>
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Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com1tag:blogger.com,1999:blog-7443075087825368900.post-43103030202639934322017-08-31T14:02:00.000-06:002018-06-12T09:29:30.735-06:00How many snakes are venomous and how many are constrictors?<div style="text-align: center;">
<span style="font-size: x-small;"><a href="https://drive.google.com/file/d/146YUU5rdbz_mdn0eqjUJjA-fws5qLYa1/view?usp=sharing">Click here to read this post in Spanish</a></span>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiz0ELDgHvLdZpiK0gVSoW6dszsrZF_l8FqEvkN8enj-KD-O2gy4ZcPWbMGMYl46IuY2KEpoeoMQXVEwrnHfYKbdkHzYPY0CxkLgF9dS9gq8Ks1QU1i8mdxc2SOvayuLCnYzWyGscwNoevH/s1600/piechart_venom_const.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="362" data-original-width="360" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiz0ELDgHvLdZpiK0gVSoW6dszsrZF_l8FqEvkN8enj-KD-O2gy4ZcPWbMGMYl46IuY2KEpoeoMQXVEwrnHfYKbdkHzYPY0CxkLgF9dS9gq8Ks1QU1i8mdxc2SOvayuLCnYzWyGscwNoevH/s320/piechart_venom_const.png" width="318" /></a></td></tr>
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<span style="font-size: x-small;">Data from <a href="http://reptile-database.reptarium.cz/advanced_search?taxon=Serpentes&submit=Search">all 3,631 living species of snakes in The Reptile Database</a></span></div>
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<span style="font-size: x-small;">as of April 2017. I made the assumption that prey-killing behavior</span></div>
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<span style="font-size: x-small;">didn't vary within genera, so if I found data for one species in a genus</span></div>
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<span style="font-size: x-small;">I applied it to all others in the absence of specific data for those species.</span></div>
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Many people are aware that some snakes constrict their prey, and others use venom to kill their prey. Recently, somebody asked me what the breakdown was, and I had to admit that I didn't know exactly. My initial estimate was that 20% were venomous in a way that is medically-significant to humans, and that probably a similar number of species are <a href="https://snakesarelong.blogspot.com/2013/09/basics-of-snake-fangs.html">opisthoglyphs</a> that use venom that is not life-threatening to humans to subdue their prey (with a decent number of these pending discovery, confirmation, or further investigation). Estimating the percentage of constrictors was more difficult, but I suspected that it was no more than the percentage of snake species that use venom, and probably somewhat less. A lot of people don't realize that there is a huge third category of snakes that just <span style="text-align: start;">seize their prey and swallow it alive, sometimes subduing it first by crushing it with strong jaws or pinning it to the ground with a coil (which hardly counts as constriction but could be an evolutionary precursor).</span></div>
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This inspired me to do some literature searching, and as I suspected nobody has ever attempted to estimate the exact percentages of snake species that use each kind of prey-killing behavior. As such, I have prepared a preliminary analysis, the full contents of which I intend to make publicly available after peer review. I hope that doing so will stimulate others to publish their observations of feeding behavior in poorly-known snakes (of which there are many), and add to the long history of discussion about the evolution of snake feeding modes, most of which took place before we had a solid grasp on <a href="http://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">the evolutionary relationships of extant snake families</a>.</div>
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I found that the answer to this question is not as simple as it may seem. Many snakes unambiguously use venom or constriction, but many use neither, and some use both! Of course the data are not as detailed or abundant as we would like. What follows is a break-down of the categories I used, and some interesting exceptions that I uncovered.<br />
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<b><span style="font-size: large;">Constrictors</span></b><br />
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Unambiguous constrictors make up just 11% of snake species, but include several well-known groups that are common in the popular consciousness, in zoos, and in the pet trade, including:</div>
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<li style="text-align: justify;"><b>Boas:</b> <a href="http://reptile-database.reptarium.cz/advanced_search?taxon=Boidae&submit=Search">61 species</a>, including the eponymous Neotropical <i>Boa constrictor</i>, anacondas (<i>Eunectes</i>), and smaller tree and rainbow boas (<i>Corallus</i>, <i>Epicrates</i>, <i>Chilabothrus</i>) as well as several (sub)families of booid snakes from various and sundry locations around the world—<i>Candoia</i> from New Guinea and Melanesia, sand boas (<i>Eryx</i>) from northeast Africa, the Middle East, and southwestern Asia, <i>Charina</i> and <i>Lichanura</i> from North America, <i>Ungaliophis</i> and <i>Exiliboa</i> from Central America, <i>Acrantophis</i> and <i>Sanzinia</i> from Madagascar, and <i>Calabaria</i> from tropical west-central Africa.</li>
<li><b>Pythons: </b><a href="http://reptile-database.reptarium.cz/advanced_search?taxon=Pythonidae&submit=Search">40 species</a> from Africa, Asia, and Australia</li>
<li style="text-align: justify;"><b>Ratsnakes, kingsnakes, and close relatives: </b>43 species of New World colubrine colubrids in the clade Lampropeltini and their Old World counterparts, including:</li>
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<li style="text-align: justify;"><i>Lampropeltis </i>kingsnakes, the consummate constrictors, <a href="http://b3.ifrm.com/30233/130/0/p3002545/The_king_of_snakes__performance_and_morphology_of_intraguild_predators__Lampropeltis__and_their_prey__Pantherophis_.pdf">which prey on other constricting snakes and are rarely, if ever, out-constricted because they are capable of exerting 20 kilopascals of pressure, twice as much as a ratsnake (average 10 kPa)</a></li>
<li style="text-align: justify;">southwestern North American species in the genera <i>Pseudelaphe</i>, <i>Arizona</i>, <i>Rhinocheilus</i>, <i>Bogertophis</i>, and <i>Senticolis</i></li>
<li style="text-align: justify;"><i>Pantherophis </i>ratsnakes, as well as their sister taxon <i>Pituophis</i> (pine, bull, and gophersnakes), which often press their prey against rocks or other solid objects</li>
<li style="text-align: justify;">2 species of live-bearing Eurasian <i>Coronella </i>and their close relative, the Frog-eating Rat Snake<i> Oocatochus rufodorsatus</i> from eastern Russia, Korea, Taiwan, and northeastern China</li>
<li style="text-align: justify;">26 species of Old World <i>Elaphe </i>and their relatives in the genera <i>Zamenis</i>, <i>Orthriophis</i>, <i>Oreocryptophis</i>, <i>Euprepiophis</i>, and <i>Archelaphe</i>.</li>
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as well as some more obscure groups:</div>
<div style="text-align: left;">
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi7jcD8CsLH9lahFh4C_DyQ3k66fqDbTtPkBFqdwjiQuNO1M0wQtj0ftQfJsBX9nNNypxzqKchjOfrVbhtR5LTUEggVq_WgB0AWw9a_O1gL_FWctt2IJy8YK4Dx5VLjqHU_7Hdhf5RcKDit/s1600/Anilius+scytale+constricting+Marques+%2526+Sazima+1998.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="295" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi7jcD8CsLH9lahFh4C_DyQ3k66fqDbTtPkBFqdwjiQuNO1M0wQtj0ftQfJsBX9nNNypxzqKchjOfrVbhtR5LTUEggVq_WgB0AWw9a_O1gL_FWctt2IJy8YK4Dx5VLjqHU_7Hdhf5RcKDit/s400/Anilius+scytale+constricting+Marques+%2526+Sazima+1998.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Anilius scytale</i> constricting an amphisbaenian<br />
From <a href="https://www.researchgate.net/profile/Ivan_Sazima2/publication/275961566_Winding_to_and_fro_constriction_in_the_snake_Anilius_scytale/links/554bb1550cf21ed2135af5ca/Winding-to-and-fro-constriction-in-the-snake-Anilius-scytale.pdf">Marques & Sazima 1998</a></td></tr>
</tbody></table>
<ul>
<li style="text-align: justify;"><b><a href="https://snakesarelong.blogspot.com/2015/01/dwarf-boas.html">Tropidophiids</a> </b>or "dwarf boas", which are not closely related to boids and certainly evolved constriction independently (34 species)</li>
<li style="text-align: justify;">Their close relative <i><a href="https://snakesarelong.blogspot.com/2015/02/anilius-pipesnake-that-wasnt.html"><b>Anilius scytale</b></a></i> (sort of; this snake <a href="https://www.researchgate.net/profile/Ivan_Sazima2/publication/275961566_Winding_to_and_fro_constriction_in_the_snake_Anilius_scytale/links/554bb1550cf21ed2135af5ca/Winding-to-and-fro-constriction-in-the-snake-Anilius-scytale.pdf">has been observed to constrict large prey</a> such as amphisbaenians)</li>
<li style="text-align: justify;"><i style="font-weight: bold;">Loxocemus bicolor</i>, the Mexican burrowing snake, a close relative of pythons</li>
<li style="text-align: justify;">Two speceis of Asian sunbeam snakes (genus <i>Xenopeltis</i>), which are also closely related to pythons</li>
<li>At least some (maybe all) Asian pipesnakes (family Cylindrophiidae)</li>
<li>Filesnakes (genus <a href="https://snakesarelong.blogspot.com/2014/08/filesnakes-wartsnakes-or-elephant.html" style="font-style: italic;">Acrochordus</a>), which don't necessarily kill fish by constricting them but use their coils to hold them while they swallow</li>
<li>some lamprophiine colubrids (especially the well-known African house snakes <i>Lamprophis</i> and <i>Boaedon</i>)</li>
<li>the colubrine colubrid tribe Lycodontini (mostly wolf snakes, genus <i>Lycodon</i>)</li>
<li>some snail-eating snakes (<i>Dipsas</i>) <a href="http://www.jstor.org/stable/1564072">coil around snails as they pry them out of their shells</a></li>
<li><a href="https://www.jstor.org/stable/pdf/1565963.pdf">even Wandering Gartersnakes (<i>Thamnophis elegans</i>)—sometimes!</a> (more below)</li>
</ul>
<div style="text-align: justify;">
These groups of snakes vary considerably in how often they employ constriction to kill their prey. Some probably use it almost all the time (although even ratsnakes eat prey that they don't constrict, such as bird eggs), whereas others use constriction only rarely, when encountering an unusually large or dangerous prey item relative to their size and strength (for example, <a href="https://link.springer.com/content/pdf/10.3758%2FBF03328698.pdf">one study</a> showed that species of <i>Python</i>, <i>Boa</i>, <i>Pantherophis</i>, and <i>Lampropeltis</i> always constricted mice if they were at least 90% the diameter of the snake's head). Some, such as <i><a href="https://www.jstor.org/stable/pdf/3891636.pdf?refreqid=excelsior%3A195ea68b0d2eed6e998c266b0df1fb9c">Regina alleni</a></i> and <i>Acrochordus</i> filesnakes, may use constriction more so to immobilize the prey than to kill it/it probably doesn’t work that well under water (although Wandering Gartersnakes <a href="https://www.jstor.org/stable/pdf/1565963.pdf">usually killed mice before eating them</a>).<br />
<br />
It seems that mammal-eating is a driver of the evolution of constriction in many cases: species that eat mammals are the only members of their genera/families that use constriction (<i>Thamnophis elegans</i>, <i>Boiga irregularis</i>, <i>Lamprophis</i>/<i>Boaedon</i>, some members of the <i><a href="http://www.scielo.br/pdf/rbzool/v13n1/v13n1a14.pdf">Oxyrhopus</a></i>/<i>Clelia</i>/<i>Pseudoboa </i>clade) and both these and species that are nested within mammal-eating clades but have shifted to other prey (<i>Lampropeltis extenuatum, Elaphe quadrivirgata, Cemophora coccinea</i><a href="#1" name="top1"><sup>1</sup></a>) tend to have more variable, less efficient constricting behavior that is generally only used to immobilize rather than to kill prey, if it is used at all. As <a href="https://www.jstor.org/stable/pdf/1565963.pdf">Alan de Queiroz and Rebecca Groen put it</a>: “<i>Thamnophis elegans</i> are not finely tuned constricting machines” and “Numerous trials in which a garter snake, holding a mouse in its jaws, was chaotically thrown about by the prey's movements support our interpretation that long constriction latencies do not reflect adaptive plasticity in <i>T. elegans</i>.”. Constriction probably functions to reduce the cost of feeding in terms of time, energy, and/or the probability that the prey will harm the snake.<br />
<br />
Conspicuously not in this category, we have the poorly-named and misleading North American Racer, <i>Coluber constrictor</i>, which is not a constrictor (<a href="https://snakesarelong.blogspot.com/2015/06/the-linnaean-snakes-part-ii.html">thanks for nothing, Linnaeus</a>).</div>
<br /></div>
</div>
<ul>
</ul>
<b style="text-align: justify;"><span style="font-size: large;">Venom</span></b><br />
<br />
<div style="text-align: justify;">
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhOeQwGGHwOi7jmE1dyvhpbAudY7b14BxxeFzq_WnjqwcjJhh9mZZfCWSbph87CUU4u0CNqVVYLh6zy9QnKTGQKMlY16PMXRQS3F6TZyKFuYWLjxZtuYJ1DrqDi90wLy5mZPVBXrc9XECPW/s1600/Dendroaspis+polylepis+eating+a+bird+Danny+Davies.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1200" data-original-width="1600" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhOeQwGGHwOi7jmE1dyvhpbAudY7b14BxxeFzq_WnjqwcjJhh9mZZfCWSbph87CUU4u0CNqVVYLh6zy9QnKTGQKMlY16PMXRQS3F6TZyKFuYWLjxZtuYJ1DrqDi90wLy5mZPVBXrc9XECPW/s320/Dendroaspis+polylepis+eating+a+bird+Danny+Davies.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Black Mamba (<i>Dendroaspis polylepis</i>) eating a bird</td></tr>
</tbody></table>
It's pretty clear which snakes use strong venom to subdue their prey; most of these are dangerously venomous to humans and so we're well aware of them. There are five major groups:</div>
<ul>
<li><b><a href="http://reptile-database.reptarium.cz/advanced_search?taxon=Viperidae&submit=Search">Viperids</a> </b>(341 species), including well-known pit vipers such as rattlesnakes, copperheads, and cottonmouths</li>
<li><b><a href="http://reptile-database.reptarium.cz/advanced_search?taxon=Elapidae&submit=Search">Elapids</a> </b>(359 species), including coralsnakes, cobras, mambas, kraits, sea snakes, and diverse terrestrial Australian snakes ranging from death adders (genus <i>Acanthophis</i>) to bandy-bandys (genus <i>Vermicella</i>)</li>
<li>Genus <a href="http://reptile-database.reptarium.cz/advanced_search?genus=atractaspis&submit=Search" style="font-style: italic; font-weight: bold;">Atractaspis</a><b style="font-style: italic;"> </b>(21 species), the <a href="https://snakesarelong.blogspot.de/2012/08/stiletto-snakes.html">stiletto snakes</a><span style="font-weight: normal;"> now known to be <a href="http://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">lamprophiids</a>, which stab backwards with their fangs, mouth closed, to envenomate prey in subterranean burrows</span></li>
<li>Non-front-fanged <b>colubrine colubrids</b>, most notably <b>boomslangs </b>(<i>Dispholidus typus</i>), <b>twigsnakes </b>(genus <i>Thelotornis</i>), and probably their close relatives in the genus <i>Thrasops</i>, all of which have many functional characteristics of front-fanged snakes while their elongated teeth remain at the rear of the (albeit rather short)
maxilla</li>
<li>some Asian <b>natricine colubrids</b> in the genera <i>Rhabdophis</i>,<i> Macropisthodon</i>,<i> </i>and <i>Balanophis</i>, which in addition to being (<a href="https://www.jstor.org/stable/pdf/1563518.pdf">in a few cases lethally</a>) venomous, also have the distinction of being among <a href="https://blogs.scientificamerican.com/guest-blog/poisonous-snakes-cant-resist-toxic-toad-tucker-or-can-they/">the only known poisonous snakes</a></li>
</ul>
<div>
<div style="text-align: justify;">
Also, many snakes use venom to subdue their prey but are not dangerous to humans, either because they have fangs in the back of their mouth, have venom that is not adapted for causing physiological damage to mammals, or both. These include:</div>
<ul>
<li>numerous <b>dipsadine </b>colubrids from the Caribbean and Central and South America, such as <i>Xenodon</i>, <i>Thamnodynastes</i>, <i>Hydrodynastes</i>, <i>Coniophanes</i>, <i>Erythrolamprus</i>, <i>Rhadinaea</i>, <i>Leptoderia</i>, and <i>Apostolepis</i> (and a few from North America, such as <i>Heterodon </i>and <i>Hypsiglena</i>)</li>
<li>some <b>colubrine </b>colubrids (genera such as <i>Boiga, Leptophis</i>, <i>Tantilla</i>, <i>Toxicodryas</i>, <i>Platyceps</i>, <i>Oxybelis</i>, <i>Hierophis</i>, <i>Crotaphopeltis</i>, <i>Drymobius</i>, <i>Chilomeniscus</i>, <i>Ficimia</i>, and <i>Gyalopion</i>) as well as the Asian genera <i>Ahaetulla </i>and <i>Chrysopelea</i>, sometimes split into a different subfamily (Ahaetullinae)</li>
<li>at least some <b>natricine</b><b style="font-style: italic;"> </b>colubrids, such as <i>Paratapinophis praemaxillaris</i> and some North American gartersnakes (<i>Thamnophis</i>)</li>
<li>many species in the family <b>Homalopsidae</b>,<b> </b>53 species of southeast Asian semi-aquatic snakes, some of which are also well-known for <a href="https://snakesarelong.blogspot.com/2012/06/snakes-that-chew-their-food.html">pulling apart large crabs and eating pieces of them</a></li>
<li>some (maybe most) <b>lamprophiids</b>, including aparallactines (<i>Amblyodipsas</i>, <i>Aparallactus</i>, <i>Micrelaps</i>, <i>Polemon</i>, <i>Xenocalamus</i>), lamprophiines (<i>Gonionotophis</i>), psamophiines (<i>Mimophis</i>, <i>Psammophis</i>), and the weird genus <i>Psammodynastes </i>("mock viper")</li>
</ul>
<div style="text-align: justify;">
and probably many more. It's actually possible that this is the largest group, because some of the "unknown" and "neither" species probably actually belong here. An interesting exception are <a href="https://snakesarelong.blogspot.com/2012/05/snakes-that-eat-caviar.html">Turtle-headed Seasnakes (<i>Emydocephalus annulatus</i>)</a> and Beaded Seasnakes (<i>Aipysurus eydouxii</i>), which eat fish eggs and have mostly lost their venom, fangs, and venom glands. Another example of a reduction in fangs are <a href="http://arizona.openrepository.com/arizona/handle/10150/196086">some fossorial species of <i>Tantilla</i></a>, which have only slightly enlarged and faintly grooved rear maxillary teeth, in contrast to the more well-developed rear fangs of most other members of this large genus. These snakes appear to specialize on beetle larvae rather than on centipedes, although no one has looked to see if their venom is any different as a result.<br />
<br /></div>
<span style="font-size: large;"><b>Neither</b></span><br />
<br />
<div style="text-align: justify;">
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh2Ad6ccxh9dOv5jz0bGKUmksALn1OPRFfVADXKyx3VQ3Utx6XLrt0NxUZhojie30RmcSjl45dQ4veqmWAq32eoAQRLjZPlkBRGw38qM_okQR5PqH3GluUm1tl6jdi1sHVdyfY3bvVFO5_5/s1600/Dipsas+indica+constricting+snail+Sazima+1989.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="575" data-original-width="815" height="225" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh2Ad6ccxh9dOv5jz0bGKUmksALn1OPRFfVADXKyx3VQ3Utx6XLrt0NxUZhojie30RmcSjl45dQ4veqmWAq32eoAQRLjZPlkBRGw38qM_okQR5PqH3GluUm1tl6jdi1sHVdyfY3bvVFO5_5/s320/Dipsas+indica+constricting+snail+Sazima+1989.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Dipsas indica </i>coiling around a snail, from <a href="https://www.jstor.org/stable/1564072?seq=1#page_scan_tab_contents">Sazima 1989</a></td></tr>
</tbody></table>
Most snakes (38% of species) seize their prey and swallow them alive. Generally these snakes are eating prey that are much smaller than they are, which lack serious physical defenses (although many of them may have chemical defenses that the snakes circumvent in other ways, such as through <a href="https://snakesarelong.blogspot.com/2014/07/tetrodotoxin-resistant-snakes.html">toxin resistance</a>). These include:</div>
<ul>
<li><b>Scolecophidians: </b>Almost 450 species of blindsnakes that <a href="https://medicine.stonybrookmedicine.edu/anatomy/people/facultypage/kley/videos">eat ant and termite larvae and pupae</a></li>
<li><a href="http://snakesarelong.blogspot.com/2017/02/shield-tailed-snakes-uropeltidae.html"><b>Uropeltids </b>or </a><b><a href="http://snakesarelong.blogspot.com/2017/02/shield-tailed-snakes-uropeltidae.html">Shield-tailed Snakes</a>: </b>55 species that eat almost exclusively earthworms</li>
<li><a href="https://www.researchgate.net/profile/Matt_Goetz/publication/233697572_The_feeding_behavior_of_the_snail-eating_snake_Pareas_carinatus_Wagler_1830_Squamata_Colubridae/links/57bfed1908aed246b0f7de31.pdf"><b>Snail-eating snakes</b> in the family <b>Pareidae</b></a><b>: </b>20 southeast Asian species <a href="https://snakesarelong.blogspot.de/2012/04/asymmetrical-snakes.html">with asymmetrical jaws</a></li>
<li>About 300 species of mostly South American <b>dipsadine colubrids </b><a href="https://snakesarelong.blogspot.com/2012/08/goo-eating-snakes-and-eggs-that-evade.html">that eat soft, gooey things</a> like slugs, snails, earthworms, or frog eggs, including:</li>
<ul>
<li>the most speciose genus of snakes, <i><a href="http://reptile-database.reptarium.cz/advanced_search?genus=Atractus&submit=Search">Atractus</a></i> (currently with 140 species)</li>
<li>the genus <i><a href="http://reptile-database.reptarium.cz/advanced_search?genus=Geophis&submit=Search">Geophis</a></i>, currently in a four-way tie for the 5th-most speciose snake genus with 50 species</li>
<li>some slender arboreal snakes in the genus <i>Sibon</i>, which crawl backward through crevices to <a href="https://oup.silverchair-cdn.com/oup/backfile/Content_public/Journal/az/23/2/10.1093/icb/23.2.455/2/23-2-455.pdf">wedge snails into them, providing an anchor against which they use their body muscles to pull out the soft parts</a></li>
<li>Asian <i><a href="http://snakesarelong.blogspot.com/2013/05/hot-spring-snakes.html">Thermophis</a></i></li>
<li>North American <i>Carphophis</i> and <i>Contia</i></li>
</ul>
<li>Most Old and New World <b>natricine colubrids</b></li>
<ul>
<li>most gartersnakes (<i>Thamnophis</i>)</li>
<li>watersnakes (<i>Nerodia</i>) and their relatives (<i>Regina</i>,<i> Seminatrix</i>,<i> Clonophis,</i> <i>Tropidoclonion</i>)</li>
<li>slug-eating <i>Storeria</i></li>
<li>Asian ecological analogues (<i>e.g.,</i> <i>Amphiesma</i>, <i>Hebius</i>, <i>Opisthotropis</i>,<i> Xenochrophis</i>)</li>
</ul>
<li>Numerous <b>colubrine colubrids</b>, such as:</li>
<ul>
<li>New World racers and coachwhips (<i>Coluber</i>)</li>
<li><a href="https://snakesarelong.blogspot.com/2012/09/recent-conservation-successes-with.html">Indigo snakes</a> (<i>Drymarchon</i>), which are well-known for crushing their prey in their strong jaws</li>
<li>Indian ratsnakes (<i>Ptyas</i>)</li>
<li>the Mole Snake (<i>Pseudaspis cana</i>)</li>
<li><a href="http://snakesarelong.blogspot.com/2012/06/egg-eating-snakes.html">Egg-eating snakes</a> (<i>Dasypeltis</i>), which have no need to kill the bird eggs that they eat or prevent them from escaping</li>
</ul>
<li><b><a href="http://reptile-database.reptarium.cz/advanced_search?taxon=Calamariinae&submit=Search">Calamariinae</a></b>, an obscure subfamily of 89 species of colubrids from Asia that are thought to eat mostly earthworms</li>
</ul>
<div>
<div>
<div style="text-align: justify;">
Some of the aforementioned goo-eaters do use their coils to support the shells of snails while they pry out the soft innards. <a href="http://www.jstor.org/stable/1564072"><i>Dipsas</i> coils around the snail’s shell</a> and <i>Sibynomorphus </i>use as s-shaped loop of their body to support the shell, whereas some <i>Sibon</i> crawl backward through crevices to <a href="https://oup.silverchair-cdn.com/oup/backfile/Content_public/Journal/az/23/2/10.1093/icb/23.2.455/2/23-2-455.pdf">wedge snails into them, providing an anchor against which they use their body muscles to pull out the soft parts</a>.</div>
</div>
</div>
<div>
<br /></div>
<span style="font-size: large;"><b>Both</b></span><br />
<br />
<div style="text-align: justify;">
Finally, there are some really interesting examples of snakes that use both venom and constriction to subdue their prey, although not always at the same time. Perhaps most impressive but <a href="https://www.reddit.com/r/snakes/comments/5m4pqz/are_there_any_snakes_that_are_constrictors_and/">least well-documented in the scientific literature</a> are two viper species that <a href="https://www.youtube.com/watch?v=EYrL75mBNwI">sometimes use constriction</a> in conjunction with venom: <i>Ovophis</i> <i>monticola </i>and <i>O.</i><i> okinavensis</i><a href="#2" name="top2"><sup>2</sup></a><i>.</i><br />
<br />
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhjXvx7nSVX4HEpys6hZ5Rr1s5n-7VFZfW27DWAv39fqbXAhn_U2jxO1hDWHUvOlDZGERZgG3eLPM5ZQtWuVBHJLBf5XrmpQwDTR8xFmv0s3sCRAzVqYjsVFA5urKhyDhyphenhyphendP4iot73cffqs/s1600/Pseudonaja-textilis-using-constriction-to-subdue-prey+Mirtschin+et+al+2006.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="381" data-original-width="407" height="299" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhjXvx7nSVX4HEpys6hZ5Rr1s5n-7VFZfW27DWAv39fqbXAhn_U2jxO1hDWHUvOlDZGERZgG3eLPM5ZQtWuVBHJLBf5XrmpQwDTR8xFmv0s3sCRAzVqYjsVFA5urKhyDhyphenhyphendP4iot73cffqs/s320/Pseudonaja-textilis-using-constriction-to-subdue-prey+Mirtschin+et+al+2006.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Pseudonaja textilis </i>constricting a mouse<br />
From <a href="http://www.venomsupplies.com/assets/published-paper.pdf">Mirtschin et al. 2006</a></td></tr>
</tbody></table>
<a href="https://www.jstor.org/stable/pdf/1445266.pdf">A review by Rick Shine & Terry Schwaner</a> brought together data on numerous Australian elapids that, although they clearly have and use venom, also use their coils to subdue and hold prey while envenoming it. In many of these species, including tiger snakes (<i>Notechis</i>), brown snakes (<i>Pseudonaja</i>), curl/myall snakes (<i>Suta</i>), whip snakes (<i>Demansia</i>), Australian coral snakes (<i>Simoselaps</i>), crowned snakes (<i>Cacophis</i>), and olive seasnakes (<i>Aipysurus laevis</i>), the coils are not used alone as the primary method of prey subjugation, and <a href="https://www.researchgate.net/profile/Bryan_Fry/publication/309457895_Rapid_Radiations_and_the_Race_to_Redundancy_An_Investigation_of_the_Evolution_of_Australian_Elapid_Snake_Venoms/links/5811198508aee15d491504ce/Rapid-Radiations-and-the-Race-to-Redundancy-An-Investigation-of-the-Evolution-of-Australian-Elapid-Snake-Venoms.pdf?origin=publication_detail&ev=pub_int_prw_xdl&msrp=QURjYUCNtT3UmRp-xq58Fwk_3qLgBeDuaP13LjZITmXgFVfRj18gd38w7x1AdNGp_EeIIj3O0c2wHC5QOw2IfLfQyKFsl4zP05E-p15PxPA.5isSzruTCNngnNYoVkf0JkQzVjOqxGnKkbJ1k-bIKukyyLArhs9IQo3pzxhFevbLbocq568be3VWLGpb4HaDhw._KK41mFzc_4z5Ej0D67aj5fqBAWAB5LvLMz7I06W4qWLJftcWocWYPs1f5202ck-44e5hU_gUF5A8-MMpV1vwg.7rGDXslXS_mbt_HA1lTprdVoXqtUxi-u_2CZ64ZQXgoWUBg7RwKXcq3DS_6m29-j6u7DD1s0z8WEg2g8KSi0UQ">one recent paper</a> suggested that we think of them as <i>"part of a 'combined arsenal' of prey subjugation strategies"</i>.<br />
<br />
To explain the "apparent paradox of why a species should use both venom and constriction to subdue its prey", Shine & Schwaner offered three possible non-mutually-exclusive explanations:<br />
<ol>
<li>The venom may be of low toxicity and thus slow to act, so holding onto the prey with either jaws or coils might allow more venom to be injected</li>
<li>Species with short fangs, such as <i>Pseudonaja</i>, and/or that feed on on heavily armored prey , such as skinks, may use constriction to give themselves additional time to find a "chink in the armor" and envenomate their prey</li>
<li>Using constriction in addition to venom may prevent snakes from losing track of bitten and envenomated prey that escape, or from being harmed by retaliating prey that are held onto</li>
</ol>
The Australian elapids recorded to use constriction feed mainly on lizards and frogs, although <a href="http://www.nrcresearchpress.com/doi/abs/10.1139/z77-144">mammals are common </a><a href="http://www.nrcresearchpress.com/doi/abs/10.1139/z77-144">prey items of <i>Pseudonaja</i> and <i>Notechis</i></a>. Puff Adders (<i>Bitis arietans</i>) <a href="https://snakesarelong.blogspot.de/2017/03/snakebite-antivenom-research-and-basic.html">choose to release large rodents and rabbits, but hold onto smaller prey</a>, although they have not been reported to use constriction (and given their specialized body shape, they probably do not, nor do they need to since they are equipped with long fangs, strong venom, and <a href="https://snakesarelong.blogspot.com/2013/03/non-toxic-venoms.html">strike-induced chemosensory searching</a>). However, immobilizing prey with coils probably plays a larger role in prey subjugation for many rear-fanged species with slower-acting venom, such as:</div>
</div>
<div>
<ul>
<li><span style="text-align: justify;">colubrine colubrids </span><i style="text-align: justify;">Boiga irregularis</i><span style="text-align: justify;">, </span><i>Macroprotodon</i>, <i>Platyceps gracilis</i>, <i>Stegonotus</i>, <i>Telescopus</i>, <i>Trimorphodon</i></li>
<li><span style="text-align: justify;">dipsadine colubrids from the Caribbean (</span><i>Alsophis</i>, <i>Cubophis</i>), Central & South America (<i>Clelia</i>, <i>Helicops, </i><i>Imantodes</i>, <i><i>Oxyrhopus</i><span style="font-style: normal;">, <i>Philodryas</i>, </span>Tropidodryas</i>, <i>Siphlophis</i>, <i>Phimophis</i>, and <i>Pseudoboa</i>), and North America (<i>Diadophis</i>, <i>Farancia</i>)</li>
<li>the sibynophiine colubrid <i>Sibynophis collaris</i></li>
<li>some homalopsids, like <i>Fordonia</i>, <i>Hypsiscopus</i>, and <i>Myron</i></li>
<li>a few lamprophiine lamprophiids, such as <i>Lycophidion</i></li>
<li>pseudaspine lamprophiids <i>Pseudaspis </i>and <i>Pythonodipsas</i></li>
<li>some pseudoxyrhophiine lamprophiids <i>Leioheterodon </i>and <i>Madagascarophis</i></li>
<li>some psammophiine lamprophiids (<i>e.g., </i>the Montpellier Snake and its relatives in the genus <i>Malpolon,</i> <i>Hemirhagerrhis, Psammophis, </i>and<i> Rhamphiophis</i>)</li>
<li>even Wandering Gartersnakes (<i>Thamnophis elegans</i>)—sometimes!</li>
</ul>
<div>
</div>
</div>
<div style="text-align: justify;">
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgGAJ6e88elv8JmOCRYB7GjfZZxYTlrw9yoO-tkaYjMFvCMvYIU3-2hu3XS1aA2aApeG6O4pv4B8DJ2e89jqXlISP8ASjBCBsRNh3POI0DltM0zFY8mcL09I4LFgtrELu7pFlK9DbhtxMLY/s1600/Elaphe_quadrivirgata_capturing_Rana_ornativentris.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1280" data-original-width="1600" height="256" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgGAJ6e88elv8JmOCRYB7GjfZZxYTlrw9yoO-tkaYjMFvCMvYIU3-2hu3XS1aA2aApeG6O4pv4B8DJ2e89jqXlISP8ASjBCBsRNh3POI0DltM0zFY8mcL09I4LFgtrELu7pFlK9DbhtxMLY/s320/Elaphe_quadrivirgata_capturing_Rana_ornativentris.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Elaphe quadrivirgata</i> not constricting a frog (<i>Rana ornativentris</i>)<br />
<a href="https://www.jstor.org/stable/pdf/1564643.pdf">Mori (1991)</a> showed that these snakes constrict large mice,<br />
pin small mice with a single coil, and swallow frogs alive</td></tr>
</tbody></table>
In many cases, only large endothermic prey (usually mammals) are constricted, whereas snakes will swallow small, easily subdued prey alive. <a href="http://research.pbsci.ucsc.edu/eeb/mehta/PDF/Mehta%26Burghardt2008.pdf">Even some specialized constrictors will consume small prey whole</a>, suggesting that almost all snakes can change strategies depending on what type of prey they are subduing. The bottom line is that, <span style="text-align: start;">if you're a snake that's eating mammals, you need to have either constriction or venom, and maybe both, because:</span><br />
<div style="text-align: start;">
</div>
<ol>
<li>Mammals are big, or at least a lot of snakes like to eat mammals that are relatively large compared to their body size</li>
<li>They are endotherms with the metabolic capacity for sustained struggling</li>
<li>They can fight back with sharp teeth and strong jaws capable of seriously injuring or killing a snake, in a way that a frog or a lizard cannot</li>
</ol>
</div>
<div style="text-align: justify;">
This generalization is supported by <a href="http://www.jstor.org/stable/1564643?seq=1#page_scan_tab_contents">observations</a> showing that <a href="http://www.jstor.org/stable/3891860">mammals tend to be killed by constriction prior to being swallowed more often than prey such as frogs</a>, and that larger prey tend to be killed by constriction first, then swallowed. Evidently <a href="https://www.researchgate.net/profile/Matthew_Bealor/publication/274083592_The_evolutionc_of_the_stimulus_control_of_constricting_behaviour_Inferences_from_North_American_gartersnakes_Thamnophis/links/5515c5b70cf2d70ee272d1ed.pdf">the amount of struggling</a> is one cue used by <i>Thamnophis elegans </i>to decide whether or not to constrict prey. Experiments carried out by <a href="http://www.sci.kyoto-u.ac.jp/en/research/researcher/akiramori-bi.html">Akira <span style="text-align: start;">Mori</span></a> and others have shown that "<a href="https://www.jstage.jst.go.jp/article/hsj2000/20/1/20_1_39/_pdf">the degree of such behavioral flexibility is, to some extent, species-specific, and it has been suggested that dietary specialists change their behavior more efficiently than dietary generalists, especially when they are young"</a>.</div>
<br />
<span style="font-size: large;"><b>Unknown</b></span><br />
<br />
<div style="text-align: justify;">
After my initial pass at collecting these data (during which I made several sweeping assumptions, some of which later turned out to be oversimplifications), I was left with 36% of species unknown. Following a more thorough literature search, I managed to get this down to 10%, which is still 363 species of snakes. In many cases I made assumptions based on generalizations about the biology of groups of snakes—for instance, I assumed that all scolecophidians use neither constriction nor venom, that all vipers use venom, and so forth. But many dipsadine and colubrine colubrids, and many lamprophiids have not been directly studied, and I could find no reports in the literature about their feeding habits. In some cases we don't even know what they eat, and ecological diversity in these groups is very high, such that there are few consistent patterns that I could use to infer prey subjugation mode for these 370 species. <a href="https://snakesarelong.blogspot.de/2014/10/how-to-teach-yourself-about-obscure.html">Teach yourself about obscure snakes</a> and help fill in the blanks!</div>
<br />
A few examples:<br />
<ul>
<li><a href="https://snakesarelong.blogspot.de/2012/04/dwarf-pipesnakes.html">anomochilids (dwarf pipesnakes)</a>—we don't even know what they eat</li>
<li>many Malagasy pseudoyrhophiine lamprophiids (<i>e.g., Ithycyphus</i>, <i>Alluaudina</i>)</li>
<li>lamprophiids from mainland Africa, such as <i><a href="http://scienceblogs.com/tetrapodzoology/2010/09/13/everything-about-bothrolycus/">Bothrolyc<span style="font-family: inherit;">us</span></a></i><span style="font-family: inherit;">, <i>Chamaelycus</i>, and <i>Dendrolycus</i></span></li>
<li>obscure dipsadines—<i>e.g., Cryophis</i>, <i>Tantalophis</i></li>
<li>obscure colubrines—<i>e.g., Bamanophis, Hemerophis</i></li>
<li>most of the <a href="https://snakesarelong.blogspot.com/2016/02/dragonsnakes-and-filesnakes-revisited.html">xenodermids (<i>Fimbrios</i>, <i>Parafimbrios</i>, <i>Stoliczkia</i>, <i>Xylophis</i>)</a></li>
<li>species known only from a few specimens, like <i><a href="https://snakesarelong.blogspot.com/2016/09/xenophidion-snake-with-mystery-penis.html">Xenophidion</a></i></li>
</ul>
<span style="font-size: large;"><b>Evolution of prey subjugation strategies in snakes</b></span><br />
<br />
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhyWb2E1Q0bQ4hZx-g83HWZYQ8kR2Ad3mLjcVMIjdiQwzAofGGJGPO6b7PewJMS6Gz1t_sMdaoXtfYNaFWbRr7i7bNmtVi3jdZHwzy3z6jH0T5Vl_hx8vMyEqZ0PZbZSWlJSqXZQddL9Ijh/s1600/Greene+1994+tree.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" data-original-height="429" data-original-width="551" height="311" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhyWb2E1Q0bQ4hZx-g83HWZYQ8kR2Ad3mLjcVMIjdiQwzAofGGJGPO6b7PewJMS6Gz1t_sMdaoXtfYNaFWbRr7i7bNmtVi3jdZHwzy3z6jH0T5Vl_hx8vMyEqZ0PZbZSWlJSqXZQddL9Ijh/s400/Greene+1994+tree.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Phylogenetic tree from <a href="https://books.google.de/books?id=8SGvMlyMXHUC&lpg=PP1&pg=PA369#v=onepage&q&f=false">Greene 1994</a><br />
For an overview of some of the updates, click <a href="https://snakesarelong.blogspot.de/2013/05/basics-of-snake-taxonomy.html">here</a></td></tr>
</tbody></table>
<div style="text-align: justify;">
<a href="https://books.google.de/books?id=8SGvMlyMXHUC&lpg=PP1&pg=PA369#v=onepage&q&f=false">The most recent similar review</a> was done by <a href="http://www.eeb.cornell.edu/greene/greene.html">Harry Greene</a> in 1994, in which he revised <a href="http://science.sciencemag.org/content/200/4337/74">earlier hypotheses</a> he put forth with <a href="http://web.utk.edu/~gburghar/">Gordon Burghardt</a> in the journal <i>Science </i>16 years before. We now know a lot more about the snake family tree than we did in 1994, particularly the fine details of relationships within the <a href="https://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">Caenophidia</a>. Overall, the basic pattern has held up rather well—constriction evolved first in basal alethinophidians during the late Cretaceous, accompanying or preceding most other evolutionary innovations that permit snakes to consume large prey, such as kinetic skulls. Greene pointed out that this was before the origin of rodents, often mentioned as potentially relevant to the evolution of snake prey-killing behaviors. Constriction was then lost at least twice—once in uropeltids (which feed underground on earthworms, although I'm not actually aware of any detailed observations of uropeltid feeding behavior) and at least once in basal colubroids, where it might have been at first replaced by venom. Venom was then subsequently lost in numerous caenophidian lineages, replaced by re-evolution of constriction in some or by other specializations (<a href="http://people.whitman.edu/~jacksok/lycodon.pdf">tooth diastemata for holding skinks</a>, <a href="http://snakesarelong.blogspot.com/2012/06/egg-eating-snakes.html">egg-eating</a>) in others, and in some caenophidian lineages snakes use both as appropriate, sometimes together (or they may elect to use neither even if both are available).</div>
<br />
<div style="text-align: justify;">
Both constriction and venom reduce the cost of feeding in terms of time, energy, and/or the probability of the prey harming the snake, but in constricting snakes, everyday locomotion and large prey neutralization are coupled, whereas in venomous snakes they are independent (snakes don't use their fangs to get around). This could be one reason why venom as an evolutionary innovation led to a more speciose radiation of snakes; it's also more susceptible to evolutionary arms races, because prey can evolve resistance to certain venom compounds, but not to constriction. Specialization for constriction is more than just behavior—<a href="https://link.springer.com/article/10.1007/s10682-010-9432-z">constricting species also have more vertebrae per unit length than non-constricting species</a>. And there are costs to both, which must be outweighed by the benefits of that defining snake trait: <a href="https://snakesarelong.blogspot.com/2012/07/snake-eating-snakes.html">being able to consume prey almost as large, and sometimes much larger, than yourself</a>!</div>
<br />
<hr width="80%" />
<span class="Apple-style-span" style="font-size: x-small;"><br />
<a href="https://www.blogger.com/null" name="1"><b>1 </b></a>An interesting exception are Scarletsnakes, <i>Cemophora coccinea</i>, the closest relatives of kingsnakes, which feed mostly on reptile eggs but also use their coils to hold lizard prey in the rare instances when they eat them. It is certain that Scarletsnakes evolved from constricting ancestors but because they almost never eat prey that need to be killed beforehand, evidently they rarely constrict.<a href="#top1"><sup>↩</sup></a><br />
</span>
<br />
<hr width="80%" />
<span class="Apple-style-span" style="font-size: x-small;"><br />
<a href="https://www.blogger.com/null" name="2"><b>2 </b></a><i>okinavensis</i> has been shown <a href="https://bmcevolbiol.biomedcentral.com/articles/10.1186/1471-2148-13-93">not to be closely related to other <i>Ovophis</i></a>, but no new genus has yet been created for it because more data are needed.<a href="#top2"><sup>↩</sup></a><br />
</span>
<br />
<br />
<div style="text-align: center;">
ACKNOWLEDGMENTS</div>
<div style="text-align: center;">
<br /></div>
<div style="text-align: justify;">
Thanks to Karen Morris for asking me this question, and to <a href="about:invalid#zClosurez">Alpsdake</a> and <a href="https://www.flickr.com/photos/101514565@N02/12724661493/in/photolist-kordxp-hNthqm-4ibSWS-5CG1Lo-T2VNqj-gfk3FU-DfNYw-9tobuN-ij7ZtY-869WfY-4CrstQ-5DcVNZ-8DZTUS-f77BCv-f77FhR-f77FfP-8ykej3-axM68Q-4r7efh-ECaMq-f77BAt-oV8KGv-dRD1pS-7pEq6D-7W8PWq-9mXP1x-4pnn5M-7pJiDb-f7mToU-2T2d6s-j987RJ-fCAGfr-7JM4Ri-dRD6Nw-7W5vZt-7W8Lw5-cDnwkh-7W8NZu-no3uL-93unEJ-f7mTkS-efTjVK-dHixp9-9nK9oZ-dHd5tv-aAGmaK-yQPK4R-4CVXZf-bkpxU7-9mXPhc">Danny Davies</a> for the use of their photos.</div>
<div style="text-align: center;">
<br /></div>
<div style="text-align: center;">
SELECTED REFERENCES</div>
<br />
<div style="text-align: justify;">
<div style="text-align: center;">
<span style="font-size: x-small;"><i>For a full list of all the references I consulted in preparing this post, click <a href="https://drive.google.com/file/d/0B2jxDXUJh_X7VWhLZ1BnVlA4dnM/view?usp=sharing">here</a></i></span></div>
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Andrade, R. d. O. and R. A. M. Silvano. 1996. Comportamento alimentar e dieta da "Falsa-coral" <i>Oxyrhopus guibei</i> Hoge & Romano (Serpentes, Colubridae). Revista Brasileira de Zoologia 13:143-150 <<a href="http://www.scielo.br/pdf/rbzool/v13n1/v13n1a14.pdf">full-text</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Auffenberg, W. 1961. Additional remarks on the evolution of trunk musculature in snakes. The American Midland Naturalist 65:1-16 <<a href="https://www.jstor.org/stable/pdf/2422998.pdf">full-text</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Bealor, M. T. and A. J. Saviola. 2007. Behavioural complexity and prey-handling ability in snakes: gauging the benefits of constriction. Behaviour 144:907-929 <<a href="https://www.researchgate.net/profile/Anthony_Saviola/publication/233600319_Behavioural_complexity_and_prey-handling_ability_in_snakes_Gauging_the_benefits_of_constriction/links/0046353bc12f2362ef000000/Behavioural-complexity-and-prey-handling-ability-in-snakes-Gauging-the-benefits-of-constriction.pdf">ResearchGate</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<br />
<div style="text-align: justify;">
<span style="font-size: x-small;"><span style="text-align: center;">Bealor, M. T., J. L. Miller, A. de Queiroz, and David A. Chiszar. 2013. </span><span style="text-align: center;">The evolution of the stimulus control of constricting behaviour: inferences from North American gartersnakes (</span><i style="text-align: center;">Thamnophis</i><span style="text-align: center;">). Behaviour 150:225-253 <<a href="https://www.researchgate.net/profile/Matthew_Bealor/publication/274083592_The_evolutionc_of_the_stimulus_control_of_constricting_behaviour_Inferences_from_North_American_gartersnakes_Thamnophis/links/5515c5b70cf2d70ee272d1ed.pdf">full-text</a>></span></span></div>
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">de Queiroz, A. and R. R. Groen. 2001. The inconsistent and inefficient constricting behavior of Colorado western terrestrial garter snakes, <i>Thamnophis elegans</i>. Journal of Herpetology 35:450-460 <<a href="https://www.jstor.org/stable/pdf/1565963.pdf">full-text</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Franz, R. 1977. Observations on the food, feeding behavior, and parasites of the striped swamp snake, <i>Regina alleni</i>. Herpetologica 33:91-94 <<a href="https://www.jstor.org/stable/pdf/3891636.pdf?refreqid=excelsior%3A195ea68b0d2eed6e998c266b0df1fb9c">full-text</a>></span></div>
<div style="text-align: justify;">
<br /></div>
<div style="text-align: justify;">
<span style="font-size: x-small;">Gans, C. 1976. Aspects of the biology of uropeltid snakes. Pages 191-204 in A. d. A. Bellairs and C. B. Cox, editors. Morphology and Biology of Reptiles. Linnean Society Symposium Series No.3. Academic Press, London.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Götz, M. 2002. The feeding behavior of the snail-eating snake <i>Pareas carinatus</i> Wagler 1830 (Squamata: Colubridae). Amphibia-Reptilia 23:487-493 <<a href="https://www.researchgate.net/publication/233697572_The_feeding_behavior_of_the_snail-eating_snake_Pareas_carinatus_Wagler_1830_Squamata_Colubridae">ResearchGate</a>></span></div>
<div style="text-align: justify;">
<span style="font-size: x-small;"><br /></span></div>
<div style="text-align: justify;">
<span style="font-size: x-small;">Greene, H. W. 1994. Homology and behavioral repertoires. Pages 369-391 in B. Hall, editor. Homology: The Heirarchical Basis of Comparative Biology. Academic Press, San Diego <<a href="https://books.google.de/books?id=8SGvMlyMXHUC&lpg=PP1&pg=PA369#v=onepage&q&f=false">Google book</a>></span></div>
<div style="text-align: justify;">
<span style="font-size: x-small;"><br /></span></div>
<div style="text-align: justify;">
<span style="font-size: x-small;">Greene, H. W. and G. M. Burghardt. 1978. Behavior and phylogeny: constriction in ancient and modern snakes. Science 200:74-77 <</span><a href="http://science.sciencemag.org/content/200/4337/74" style="font-size: x-small;">abstract</a><span style="font-size: x-small;">></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Hampton, P. M. 2011. Ventral and sub-caudal scale counts are associated with macrohabitat use and tail specialization in viperid snakes. Evolutionary Ecology 25:531-546 <<a href="https://link.springer.com/article/10.1007/s10682-010-9432-z">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Holm, P. A. 2008. Phylogenetic biology of the burrowing snake tribe Sonorini (Colubridae). PhD dissertation. University of Arizona <<a href="http://arizona.openrepository.com/arizona/handle/10150/196086">full-text</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Jackson, K. and T. H. Fritts. 2004. Dentitional specialisations for durophagy in the Common Wolf snake, <i>Lycodon aulicus </i>capucinus. Amphibia-Reptilia 25:247-254 <<a href="http://people.whitman.edu/~jacksok/lycodon.pdf">full-text</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Loop, M. S. and L. G. Bailey. 1972. The effect of relative prey size on the ingestion behavior of rodent-eating snakes. Psychonomic Science 28:167-169 <<a href="https://link.springer.com/content/pdf/10.3758%2FBF03328698.pdf">full-text</a>></span></div>
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<span style="font-size: x-small;">Marques, O. A. V. and I. Sazima. 2008. Winding to and fro: constriction in the snake </span><i style="font-size: small;">Anilius scytale</i><span style="font-size: x-small;">. Herpetological Bulletin 103:29-31 <<a href="https://www.researchgate.net/profile/Ivan_Sazima2/publication/275961566_Winding_to_and_fro_constriction_in_the_snake_Anilius_scytale/links/554bb1550cf21ed2135af5ca/Winding-to-and-fro-constriction-in-the-snake-Anilius-scytale.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Martins Teixeria, D., M. Luci Lorini, V. G. Persson, and M. Porto. 1991. <i>Clelia clelia</i> (Mussurana). Feeding behavior. Herpetological Review 22:131-132 <<a href="https://www.dropbox.com/s/v82fk176wpb21xd/HR%201991.22.4R.pdf?dl=1">link</a>></span></div>
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<span style="font-size: x-small;">Mehta, R. S. and G. M. Burghardt. 2008. Contextual flexibility: reassessing the effects of prey size and status on prey restraint behaviour of macrostomate snakes. Ethology 114:133-145 <<a href="http://research.pbsci.ucsc.edu/eeb/mehta/PDF/Mehta%26Burghardt2008.pdf">full-text</a>></span><br />
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<span style="font-size: x-small;">Mirtschin, P. J., N. Dunstan, B. Hough, E. Hamilton, S. Klein, J. Lucas, D. Millar, F. Madaras, and T. Nias. 2006. Venom yields from Australian and some other species of snakes. Ecotoxicology 15:531-538 <<a href="http://www.venomsupplies.com/assets/published-paper.pdf">full-text</a>></span><br />
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<span style="font-size: x-small;">Mori, A. 1991. Effects of prey size and type on prey-handling behavior in <i>Elaphe quadrivirgata</i>. Journal of Herpetology 24:160-166 <<a href="https://www.jstor.org/stable/pdf/1564643.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Mori, A. and K. Tanaka. 2001. Preliminary observations on chemical preference, antipredator responses, and prey-handling behavior of juvenile <i>Leioheterodon madagascariensis</i> (Colubridae). Current Herpetology 20:39-49 <<a href="https://www.jstage.jst.go.jp/article/hsj2000/20/1/20_1_39/_pdf">full-text</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Mushinsky, H. R. 1984. Observations of the feeding habits of the short-tailed snake, <i>Stilosoma extenuatum</i> in captivity. Herpetological Review 15:67-68 <<a href="https://www.dropbox.com/s/ryi201r8xcar0a4/HR%201984.15.3R.pdf?dl=1">link</a>></span></div>
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<span style="font-size: x-small;">Penning, D. A. and B. R. Moon. 2017. The king of snakes: performance and morphology of intraguild predators (<i>Lampropeltis</i>) and their prey (<i>Pantherophis</i>). The Journal of Experimental Biology 220:1154 <<a href="http://b3.ifrm.com/30233/130/0/p3002545/The_king_of_snakes__performance_and_morphology_of_intraguild_predators__Lampropeltis__and_their_prey__Pantherophis_.pdf">link</a>></span></div>
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<span style="font-size: x-small;">Rossi, J. V. and R. Rossi. 1993. Notes on the captive maintenance and feeding behavior of a juvenile short-tailed snake (Stilosoma extenuatum). Herpetological Review 24:100-101 <<a href="https://www.dropbox.com/s/pr3l437v7idasef/HR%201993.24.3R.pdf?dl=1">link</a>>
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<span style="font-size: x-small;"><span style="font-size: x-small;">Savitzky, A. H. 1980. The role of venom delivery strategies in snake evolution. Evolution 34:1194-1204 <<a href="http://onlinelibrary.wiley.com/doi/10.1111/j.1558-5646.1980.tb04065.x/full">link</a>></span></span><br />
<span style="font-size: x-small;"><span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Sazima, I. 1989. Feeding behavior of the snail-eating snake, <i>Dipsas indica</i>. Journal of Herpetology 23:464-468 <<a href="http://www.jstor.org/stable/1564072">link</a>></span></span></div>
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<span style="font-size: x-small;"><span style="font-size: x-small;">Shine, R. 1977. Habitats, diets, and sympatry in snakes: a study from Australia. Canadian Journal of Zoology 55:1118-1128 <</span><a href="http://www.nrcresearchpress.com/doi/abs/10.1139/z77-144" style="font-size: x-small;">abstract</a><span style="font-size: x-small;">></span></span></div>
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<span style="font-size: x-small;"><span style="font-size: x-small;">Shine, R. and T. Schwaner. 1985. Prey constriction by venomous snakes: a review, and new data on Australian species. Copeia 1985:1067-1071 <<a href="https://www.jstor.org/stable/pdf/1445266.pdf">link</a>></span></span><br />
<span style="font-size: x-small;"><span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Stettler, P. H. 1959. Zur Lebensweise von <i>Dipsas turgidus </i>(Cope), einer schneckenfressenden Schlange. Aquarien und Terrarien 8:238-241.</span></span></div>
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<span style="font-size: x-small;"><span style="font-size: x-small;">Vidal, N. and S. B. Hedges. 2002. Higher-level relationships of snakes inferred from four nuclear and mitochondrial genes. Comptes Rendus-Biologies 325:977-985 <<a href="https://www.academia.edu/462748/Higher-level_relationships_of_snakes_inferred_from_four_nuclear_and_mitochondrial_genes">link</a>></span></span></div>
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<span style="font-size: x-small;"><span style="font-size: x-small;">Willard, D. E. 1977. Constricting methods of snakes. Copeia 1977:379-382 <<a href="https://www.jstor.org/stable/1443922">link</a>></span></span></div>
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<span style="font-size: x-small;"><span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="https://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></span></div>
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</div>Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com3Jena, Germany50.927054 11.58923720000007150.7669025 11.26651370000007 51.087205499999996 11.911960700000071tag:blogger.com,1999:blog-7443075087825368900.post-15057093524783091462017-05-31T09:35:00.000-06:002017-06-09T07:10:44.875-06:00Snakes of Morocco<div style="text-align: justify;">
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<span style="font-size: x-small;"><a href="https://drive.google.com/file/d/0B2jxDXUJh_X7TTV5eV95dHpoLVU/view?usp=sharing">Click here to read this post in Spanish!</a></span><a href="https://drive.google.com/file/d/0B2jxDXUJh_X7TTV5eV95dHpoLVU/view?usp=sharing" style="font-size: small;">¡Haga clic aquí para leer este post en español!</a></div>
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Earlier this month I went to Morocco to attend <a href="https://sites.google.com/site/5thbiologyofthevipers/home">the 5th Biology of the Vipers conference</a>. The conference was organized by <a href="https://cibio.up.pt/people/details/fmfreiria">Fernando Martínez-Freiría</a> and <a href="https://www.researchgate.net/profile/Soumia_Fahd2">Soumia Fahd</a> and featured a fantastic <a href="https://sites.google.com/site/5thbiologyofthevipers/programme/conference">three-day scientific program</a> in Chefchaouen followed by a <a href="https://sites.google.com/site/5thbiologyofthevipers/programme/post-meeting-excursion">six-day field excursion</a> to southern Morocco to look for snakes. I learned a lot and got lots of great feedback on <i>Life is Short but Snakes are Long</i>, but unfortunately I didn't have time to finish writing May's article, which I aim to put up next week. On the way back, I also stopped by Jerez de la Frontera and finally met Alvaro Pemartin and Estefania Carrillo, whose dedicated translations have brought <i>Life is Short but Snakes are Long </i>to Spanish-speaking readers around the world!</div>
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<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj4k47lh_FpbTrZmRaBLkoqRox3Fv93HXS4OVgfYG9RozBYWgmIuXf-NuyTgtf5ZP1QH3D1a73WBOZevkKYhBTL03OoTDejVqHKPSvy0FFEU_W-pXT7u73PBtaejm1UqaEn93OFp5yBLjZy/s1600/IMG_3208.JPG" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1201" data-original-width="1600" height="300" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj4k47lh_FpbTrZmRaBLkoqRox3Fv93HXS4OVgfYG9RozBYWgmIuXf-NuyTgtf5ZP1QH3D1a73WBOZevkKYhBTL03OoTDejVqHKPSvy0FFEU_W-pXT7u73PBtaejm1UqaEn93OFp5yBLjZy/s400/IMG_3208.JPG" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Montpellier snake (<i>Malpolon monspessulanus</i>)</td></tr>
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<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjJNXOFrQ1KjvITfhs-uU5xcX_sAOC_p1OHfR5kTEsduQ98O6t25ApJ9zdWSJEjfv7ywOcDSehuTuDTzsEI9xowIblIoaJ3e_oEhRGFr1bhEddEoG4_jQsJC1cOxJfmTj8W-sVrN7sVwUfH/s1600/IMG_3391.JPG" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1201" data-original-width="1600" height="300" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjJNXOFrQ1KjvITfhs-uU5xcX_sAOC_p1OHfR5kTEsduQ98O6t25ApJ9zdWSJEjfv7ywOcDSehuTuDTzsEI9xowIblIoaJ3e_oEhRGFr1bhEddEoG4_jQsJC1cOxJfmTj8W-sVrN7sVwUfH/s400/IMG_3391.JPG" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Saharan horned viper (<i>Cerastes cerastes</i>)</td></tr>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiybFOu13dDAkIVBi3Xv63scyDrvvIf1MqH9n_pRKSc0WDCNs-GyensuXlx0estymRGzQCz0dLP5W6mrOmwZPtStEsYdHuJt_Qr0RcQBUNvIOSebLtvsZXzB26Im_lXbJm7ToObW-zZ0dzn/s1600/IMG_3400.JPG" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1200" data-original-width="1600" height="300" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiybFOu13dDAkIVBi3Xv63scyDrvvIf1MqH9n_pRKSc0WDCNs-GyensuXlx0estymRGzQCz0dLP5W6mrOmwZPtStEsYdHuJt_Qr0RcQBUNvIOSebLtvsZXzB26Im_lXbJm7ToObW-zZ0dzn/s400/IMG_3400.JPG" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Puff Adder (<i>Bitis arietans</i>)</td></tr>
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<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgyDFyMDyuToqldxO8Qsc6T6uUqSJa-ulgpUTt5xwwlp_fUdXj5GemhFG5c2DKtIh79LaunHtMbAIr10NofdgLeLJmkQrdFoTjrDIdQaesfa5Ubm2wAIU8JacESsaUvZC7gj_IkdpVf3rVN/s1600/IMG_3476.JPG" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1200" data-original-width="1600" height="300" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgyDFyMDyuToqldxO8Qsc6T6uUqSJa-ulgpUTt5xwwlp_fUdXj5GemhFG5c2DKtIh79LaunHtMbAIr10NofdgLeLJmkQrdFoTjrDIdQaesfa5Ubm2wAIU8JacESsaUvZC7gj_IkdpVf3rVN/s400/IMG_3476.JPG" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Egyptian Cobra (<i>Naja haje</i>)</td></tr>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgmK5sYna8-9IUhegxuAlRKoXKEjb6qhxYe8X1jFa04wsUcgRMLgeiW-Ml7YSe3wNmXWPh5bshjp2q155WEzr_y8s9nMl57nX4Ixt1mNPAv82z1WJ5pxVsc-YBz24vMzGbUvLkSXAdTXQxk/s1600/_DSC3249.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1600" data-original-width="1069" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgmK5sYna8-9IUhegxuAlRKoXKEjb6qhxYe8X1jFa04wsUcgRMLgeiW-Ml7YSe3wNmXWPh5bshjp2q155WEzr_y8s9nMl57nX4Ixt1mNPAv82z1WJ5pxVsc-YBz24vMzGbUvLkSXAdTXQxk/s320/_DSC3249.jpg" width="213" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Me on the streets of Chefchaouen</td></tr>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgzADcBowVqFiIjipkEp3LpUgfJrlhfIRQIca5alOIJ1lzeAGg3RhBEluXWEUHCVY2rKt8obKjr7fK5tiL8Cw5kD-I4WIlZbQXeMAgwqaOQG_6DFhe_c6M4xvUYdQT3upo9h7wtw4JMbUEJ/s1600/P1390851.JPG" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1600" data-original-width="1200" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgzADcBowVqFiIjipkEp3LpUgfJrlhfIRQIca5alOIJ1lzeAGg3RhBEluXWEUHCVY2rKt8obKjr7fK5tiL8Cw5kD-I4WIlZbQXeMAgwqaOQG_6DFhe_c6M4xvUYdQT3upo9h7wtw4JMbUEJ/s320/P1390851.JPG" width="240" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Me looking for vipers in Toubkal National Park</td></tr>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjIsNUHJRe_tCs_rGcK4PRRExk1Mi8fpm2e72xVmoXh8isfhPvEHJGfM9-NtBOEO09fRda8GR1pgpkV80maLVLpf3swYjrl9cYcVWc-0i2DSa-DQtdu1SzI_JnT6aZXkg9atvfdZhPpOtzi/s1600/IMG-20170524-WA0002.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" data-original-height="1200" data-original-width="1600" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjIsNUHJRe_tCs_rGcK4PRRExk1Mi8fpm2e72xVmoXh8isfhPvEHJGfM9-NtBOEO09fRda8GR1pgpkV80maLVLpf3swYjrl9cYcVWc-0i2DSa-DQtdu1SzI_JnT6aZXkg9atvfdZhPpOtzi/s320/IMG-20170524-WA0002.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Estefania and I in Jerez</td></tr>
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We also met with an Aisaoua, a member of the traditional brotherhood of snake hunters in Morocco, who collect the snakes that are used by the <a href="https://jleetingle.wordpress.com/2013/11/25/moroccan-serpent-charming/">snake charmers</a> who put on shows. This tradition is at least 800 years old, and possibly as old as 2,000 years, and is an example of a human-reptile interaction with both positive and negative aspects. It was really interesting to see their method for finding snakes—they are very effective! More on this in a future article.</div>
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ACKNOWLEDGMENTS</div>
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Thanks to Konrad Mebert and Alvaro Pemartin III for allowing me to use their photographs, </div>
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REFERENCES</div>
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<span style="font-size: x-small;">Bons, J., P. Geniez, A. Montori, V. Roca, and E. Asociación Herpetológica. 1996. Amphibiens et reptiles du Maroc (Sahara Occidental compris) : atlas biogéographique = Anfibios y reptiles de Marruecos (incluido Sáhara Occidental) : atlas biogeográfico = Amphibians & reptiles of Morocco (including Western Sahara) : biogeographical atlas.</span></div>
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<span style="font-size: x-small;">Pleguezuelos, J. M., M. Feriche, J. C. Brito, and S. Fahd. 2016. Snake charming and the exploitation of snakes in Morocco. Oryx :1-8 <<a href="https://www.researchgate.net/publication/311851291_Snake_charming_and_the_exploitation_of_snakes_in_Morocco">link</a>></span></div>
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<span style="font-size: x-small;">Tingle, J. L. and T. Slimani. 2017. Snake charming in Morocco. The Journal of North African Studies :1-18 <<a href="https://www.researchgate.net/profile/Tahar_Slimani/publication/315335770_The_Journal_of_North_African_Studies_Snake_charming_in_Morocco_Snake_charming_in_Morocco/links/58ccf2154585157b6dac20aa/The-Journal-of-North-African-Studies-Snake-charming-in-Morocco-Snake-charming-in-Morocco.pdf">link</a>></span><br />
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="https://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
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Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com2tag:blogger.com,1999:blog-7443075087825368900.post-84340577064732265122017-04-30T12:48:00.002-06:002017-09-02T01:48:51.619-06:00The 21st century blindsnake revolution<div style="text-align: center;">
<span style="font-size: x-small;"><a href="https://drive.google.com/file/d/0B2jxDXUJh_X7dmlhcXVUSHBMU2M/view?usp=sharing">Click here to read this post in Spanish</a></span><br />
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<tr><td class="tr-caption" style="text-align: center;">Brongersma's Wormsnake (<i><a href="http://reptile-database.reptarium.cz/species?genus=Amerotyphlops&species=brongersmianus">Amerotyphlops brongersmianus</a></i>),<br />
a widespread species from South America</td></tr>
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Blindsnakes (Scolecophidia) don't get enough attention. They include <a href="https://snakesarelong.blogspot.com/2014/04/the-most-widespread-snake-in-world.html">the world's most widespread snake species</a>, the world's <a href="http://reptile-database.reptarium.cz/species?genus=Tetracheilostoma&species=carlae">smallest living snake species</a>, and <a href="https://medicine.stonybrookmedicine.edu/anatomy/people/facultypage/kley/videos">a diversity of jaw-raking feeding mechanisms unrivaled in bizarreness among land vertebrates</a>. I recently noticed, much to my surprise, the the number of described species of blindsnakes has doubled in the last 13 years, from 305 in 2004 to 599 today; that's 16.5% of all snakes! <i>June 2017 EDIT: This was a big mistake on my part. As of 2017 there are 442 species described instead of 599. I made this mistake because I was confused about the search terms being used on my go-to reference for reptile taxonomy, <a href="http://reptile-database.reptarium.cz/advanced_search">The Reptile Database</a>. I was assuming that Leptotyphlopidae + Anomalepidae + Typhlopoidea = Scolecophidia, a search term that is no longer available in The Reptile Database, because of several phylogenies that show it to be paraphyletic. If you search for "Typhlopoidea" on The Reptile Database, you get a list of all 442 blindsnakes 442, including Leptotyphlopidae and Anomalepididae, and not only the three families of Typhlopoidea according to Vidal et al. 2010 (Typhlopidae, Xenotyphlopidae, Gerrhopilidae). I thought that Typhlopoidea only returned the latter three families and I added the 139 species of Leptotyphlopidae and 18 species of Anomalepididae to get an incorrect total of 599. Thanks to <a href="https://www.zfmk.de/en/zfmk/claudia-koch">Claudia Koch</a> of the Alexander Koenig Zoological Research Museum in Bonn for pointing this out to me.</i> There are certainly many undiscovered species of blindsnakes, so it's likely that their numbers will continue to grow (as one recent study put it, <a href="https://www.researchgate.net/profile/Rafe_Brown/publication/297599848_Two_New_Species_of_Malayotyphlops_from_the_Northern_Philippines_with_Redescriptions_of_Malayotyphlops_luzonensis_Taylor_and_Malayotyphlops_ruber_Boettger/links/575496d508ae6807fb04d068.pdf">"...even our most liberal estimates of species numbers will likely prove to be an underestimate of the true diversity...of secretive blind snakes"</a>).</div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh4td-_PHBK600h2ZSa88SvhcKS0tL77OELxeKdusMUaUyadidJuELRS-DVRhOhvc9IfIBjGi80aMIK79AzmIfLYmdXWMWoQKp3e9iyLkOsyQW-jfqHkSqGNryD80Ed-P9bcLW386w5Rtud/s1600/Blindsnake+evolutionary+tree+Vidal+et+al+2010.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="400" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh4td-_PHBK600h2ZSa88SvhcKS0tL77OELxeKdusMUaUyadidJuELRS-DVRhOhvc9IfIBjGi80aMIK79AzmIfLYmdXWMWoQKp3e9iyLkOsyQW-jfqHkSqGNryD80Ed-P9bcLW386w5Rtud/s400/Blindsnake+evolutionary+tree+Vidal+et+al+2010.jpg" width="272" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Blindsnake evolutionary tree.<br />
Extinction of the dinosaurs (K-T boundary) was<br />
between the green and pink-shaded areas.<br />
From <a href="http://pubmedcentralcanada.ca/pmcc/articles/PMC2936224/pdf/rsbl20100220.pdf">Vidal et al. 2010</a></td></tr>
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One of the biggest phylogenetic rearrangements within the Scolecophidia was <a href="http://pubmedcentralcanada.ca/pmcc/articles/PMC2936224/pdf/rsbl20100220.pdf">the recognition of two new families in 2010</a>. The new families <a href="http://reptile-database.reptarium.cz/advanced_search?taxon=Gerrhopilidae&exact%5B0%5D=taxon&submit=Search">Gerrhopilidae</a> and <a href="http://reptile-database.reptarium.cz/species?genus=Xenotyphlops&species=grandidieri&search_param=%28%28taxon%3D%27Xenotyphlopidae%27%29%29">Xenotyphlopidae</a> were formerly part of <a href="http://reptile-database.reptarium.cz/advanced_search?taxon=typhlopidae&submit=Search">Typhlopidae</a>, but were discovered to be distantly related to other typhlopids and were separated, although these three families are grouped together in the superfamily <a href="http://reptile-database.reptarium.cz/advanced_search?taxon=Typhlopoidea&submit=Search">Typhlopoidea</a> to emphasize their closer relationship to one another than to the other two families of scolecophidians (<a href="http://reptile-database.reptarium.cz/advanced_search?taxon=Leptotyphlopidae&submit=Search">Leptotyphlopidae</a> and <a href="http://reptile-database.reptarium.cz/advanced_search?taxon=Anomalepididae&submit=Search">Anomalepididae</a>). The original diversification of blindsnakes is thought to have been caused by the breakup of Gondwana, whereas the later diversification of Typhlopoidea is associated with the breakup of East Gondwana into Antarctica, Madagascar, India, and Australia (with subsequent colonization by typhlopids from West Gondwana [Africa/South America]). Subsequent diversification within the Typhlopidae coincides with the early Paleozoic Era, just after the extinction of the dinosaurs, and includes four major groups: a Eurasian-Australasian one, an African one, a Malagasy one, and a South American-West Indian one. Because sea levels were low at this time, dispersal among continents and islands was relatively easy, at least for a small vertebrate with low metabolism and most likely travelling along with their invertebrate prey. The relationships of blindsnakes track plate tectonics better than those of any other vertebrate group, perhaps because of their tendency to stay put.</div>
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<tr><td class="tr-caption" style="text-align: center;"><i>Gerrhopilus mirus</i> from Sri Lanka</td></tr>
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The two "new" families probably originated on the ancient landmass "Indigascar" (modern India and Madagascar, which were physically connected long after their isolation from other continents and India's subsequent unification with Asia). One family, <a href="http://reptile-database.reptarium.cz/advanced_search?taxon=Gerrhopilidae&exact%5B0%5D=taxon&submit=Search">Gerrhopilidae</a> ("Indo-Malayan blindsnakes"), were formerly known as the <i>Typhlops ater</i> species group. They differ from other blindsnakes in having gland-like structures ‘peppered’ over the head scales. Many species also have a divided preocular and/or ocular scale, and the second supralabialal scale overlaps the preocular in all species but one (<i>G. tindalli</i>). The family contains at least 16 species in the genus <i>Gerrhopilus</i>, and possibly others (the most-recently described species are from <a href="https://www.researchgate.net/profile/Van_Wallach/publication/277038209_Two_New_Blind_Snakes_of_the_Typhlops_ater_Species_Group_from_Papua_New_Guinea_Serpentes_Typhlopidae/links/570826dc08aed09e916d24a8.pdf">1996</a> and <a href="https://www.jstor.org/stable/4092848">2005</a>). This is where it starts to get really weird.</div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhLhBvtYkXFv7RzrWqQzSgfB7B2QY_pLokMimE7qq5V8qiMuwr1UrFNNN5F7NIlr7QmV2ieIl7oi5VrJ_ovv-EiPfreiUY_hGVzv8Q3Us92uqQzQIGFCNmPMKISnoqdsPTER5kOTzNFA_l6/s1600/1811_Freycinet_Map.jpg" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="212" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhLhBvtYkXFv7RzrWqQzSgfB7B2QY_pLokMimE7qq5V8qiMuwr1UrFNNN5F7NIlr7QmV2ieIl7oi5VrJ_ovv-EiPfreiUY_hGVzv8Q3Us92uqQzQIGFCNmPMKISnoqdsPTER5kOTzNFA_l6/s320/1811_Freycinet_Map.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><a href="https://en.wikipedia.org/wiki/Freycinet_Map_of_1811">The 1811 Freycinet map of Australia</a>, where<br />
<i>Cathetorhinus melanocephalus</i> was not found</td></tr>
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There is another candidate member of the family Gerrhopilidae. The genus <i>Cathetorhinus </i>contains a single species, known from only a single specimen (<a href="https://science.mnhn.fr/institution/mnhn/collection/ra/item/0.138">Natural History Museum, Paris RA-0.138</a>, an adult male). It was collected by French zoologists François Péron and Charles-Alexandre Lesueur on <a href="https://en.wikipedia.org/wiki/Baudin_expedition_to_Australia">a scientific expedition to Australia led by Nicolas Baudin</a> between 1801 and 1803, and scientifically described (along with an unprecedented and unqeualed number of other new snake species) <a href="http://www.biodiversitylibrary.org/item/101228#page/286/mode/1up">in the 1844 volume of Duméril & Bibron's opus </a><i><a href="http://www.biodiversitylibrary.org/item/101228#page/286/mode/1up">Erpetologie Générale</a> </i>(the series is also the provenance of the mudsnake plate that I use as a logo for this blog). <i>Cathetorhinus melanocephalus </i>was the only blindsnake they collected, despite visiting the Canary Islands, Mauritius, Timor, and South Africa in addition to Australia (of which members of the expedition later produced <a href="https://en.wikipedia.org/wiki/Freycinet_Map_of_1811">the first complete map</a>). Unfortunately, for reasons lost to history and despite their general habits as conscientious collectors<a href="#1" name="top1"><sup>1</sup></a>, the location where they found <i>Cathetorhinus melanocephalus</i> was not recorded (I'm speculating here, but it may have been because they were distracted by fearing for their lives—of a total of 24 scientists who went on the expedition, <a href="https://en.wikipedia.org/wiki/Baudin_expedition_to_Australia#Scientists_and_artists">5 died and 10 disembarked at Mauritius due to illness</a>).</div>
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<tr><td class="tr-caption" style="text-align: center;"><i>Cathetorhinus melanocephalus</i><br />
From <a href="https://www.researchgate.net/profile/Olivier_Pauwels/publication/236983437_The_systematic_status_of_Cathetorhinus_melanocephalus_Dumeril_Bibron_1844_Serpentes_Typhlopidae/links/02e7e51ac7d54d25cd000000/The-systematic-status-of-Cathetorhinus-melanocephalus-Dumeril-Bibron-1844-Serpentes-Typhlopidae.pdf?origin=publication_detail&ev=pub_int_prw_xdl&msrp=cGRCmIMcir9sD1AtjG4BAu6Up2PBlq_Nm_D00LAF6Q0MAsqtVkae9C1QrL7fIPuJqLn73Cl1A1zfiqbDXLN3jhZiZGPCsZbESKGrtIiZlxs.XHJwWSAnXm9BaKBbMjVJthuW7gX4XryX0G4MasxcxF0CmLt7wNXNTc7oCvQuzbr7M5ol-U26bGaNbQGHEmkLzQ.I47bax12KYTyDPgzn-jhcYc2gj4iDJ_PKY_JVXzIecWc_y80R0LC2wbK8X-EhN_2NNI2n2UfSOSusrURCBne9A.zNRpW3j16R7Vwr5lgFbDJWVqjfa6mSr5V3wW4XXTuEXBrC0PKwiO92j97URY3yFPKIlrm1ik2nxZnNdUNEGgng">Wallach & Pauwels 2008</a></td></tr>
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This wouldn't be such a problem (lots of type specimens have vague or missing type localities; <a href="https://snakesarelong.blogspot.com/2015/05/linnaean-snakes-part-i.html">Linnaeus correctly attributed fewer than half of his snakes to the right continent</a> "Indiis") except that no other specimens have ever been found. It is taxonomically unique <a href="https://www.researchgate.net/profile/Olivier_Pauwels/publication/236983437_The_systematic_status_of_Cathetorhinus_melanocephalus_Dumeril_Bibron_1844_Serpentes_Typhlopidae/links/02e7e51ac7d54d25cd000000/The-systematic-status-of-Cathetorhinus-melanocephalus-Dumeril-Bibron-1844-Serpentes-Typhlopidae.pdf?origin=publication_detail&ev=pub_int_prw_xdl&msrp=cGRCmIMcir9sD1AtjG4BAu6Up2PBlq_Nm_D00LAF6Q0MAsqtVkae9C1QrL7fIPuJqLn73Cl1A1zfiqbDXLN3jhZiZGPCsZbESKGrtIiZlxs.XHJwWSAnXm9BaKBbMjVJthuW7gX4XryX0G4MasxcxF0CmLt7wNXNTc7oCvQuzbr7M5ol-U26bGaNbQGHEmkLzQ.I47bax12KYTyDPgzn-jhcYc2gj4iDJ_PKY_JVXzIecWc_y80R0LC2wbK8X-EhN_2NNI2n2UfSOSusrURCBne9A.zNRpW3j16R7Vwr5lgFbDJWVqjfa6mSr5V3wW4XXTuEXBrC0PKwiO92j97URY3yFPKIlrm1ik2nxZnNdUNEGgng">based on its morphology</a>, descriptions of which have been rather inconsistent over the decades, partially because blindsnakes are really small and their scales are really hard to count, especially given the crummy optics of the 19th century. Except for the head glands, <i>Cathetorhinus </i>shares more anatomical characteristics with <i>Gerrhopilus</i> than with any other blindsnakes. A 2008 study reviewed the history of the Baudin expedition and concluded that <a href="https://www.researchgate.net/profile/Olivier_Pauwels/publication/236983437_The_systematic_status_of_Cathetorhinus_melanocephalus_Dumeril_Bibron_1844_Serpentes_Typhlopidae/links/02e7e51ac7d54d25cd000000/The-systematic-status-of-Cathetorhinus-melanocephalus-Dumeril-Bibron-1844-Serpentes-Typhlopidae.pdf?origin=publication_detail&ev=pub_int_prw_xdl&msrp=cGRCmIMcir9sD1AtjG4BAu6Up2PBlq_Nm_D00LAF6Q0MAsqtVkae9C1QrL7fIPuJqLn73Cl1A1zfiqbDXLN3jhZiZGPCsZbESKGrtIiZlxs.XHJwWSAnXm9BaKBbMjVJthuW7gX4XryX0G4MasxcxF0CmLt7wNXNTc7oCvQuzbr7M5ol-U26bGaNbQGHEmkLzQ.I47bax12KYTyDPgzn-jhcYc2gj4iDJ_PKY_JVXzIecWc_y80R0LC2wbK8X-EhN_2NNI2n2UfSOSusrURCBne9A.zNRpW3j16R7Vwr5lgFbDJWVqjfa6mSr5V3wW4XXTuEXBrC0PKwiO92j97URY3yFPKIlrm1ik2nxZnNdUNEGgng">“the provenance of this species remains unknown: it is certainly Old World, and may be from (in order of probability) Timor, Australia, Mauritius or Tenerife”</a>. And so it would have remained, if not for some really excellent bibliographical sleuthing by biologist and scholar <a href="http://dodobooks.com/anthony-chekes-publications/">Anthony Cheke</a>, an expert on Mascarene fauna. Cheke reviewed the unpublished original notes made by Lesueur on the voyage, and found a reference to "a very small [snake] species 4–5 inches maximum...the only one found during our stay [on Mauritius in 1803]...found amongst stones while clearing some land...about 8 inches be-low the soil surface". This tantalizing description suggests a blindsnake in size, habitat, and behavior, and although Cheke himself had assumed that it referred to the <a href="https://snakesarelong.blogspot.com/2014/04/the-most-widespread-snake-in-world.html">Brahminy Blindsnake (<i>Indotyphlops braminus</i>)</a>, he later realized that the first records of introduction of this widespread species were from 1869, 66 years later.<a href="#2" name="top2"><sup>2</sup></a> Although this isn't concrete proof, it's highly suggestive that Lesueur's blindsnake was <i>Cathetorhinus melanocephalus</i>, since it was the only blindsnake collected on the entire journey.<a href="#3" name="top3"><sup>3</sup></a> <span style="text-align: start;">Fossils of an endemic Mauritian typhlopid were discovered around 1900 and described as </span><i style="text-align: start;">Typhlops cariei</i><span style="text-align: start;">, but direct comparison of the bones with those of </span><i>Cathetorhinus </i>has not been made. Could <i>Cathetorhinus </i>still survive in the wild? Many non-native blindsnake predators were already introduced to Mauritius when Lesueur and Péron visited, including rats, shrews, and tenrecs, and others have since become established, such as mongeese. Only time, and further field work on Mauritius, will tell.<br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEghm_Rs4lti4jslM5Gyspd5cvQoua3f7O0MavizvN8nMq5HVYwYotQpZps0BIFefPWJprp_pxSF_wLuzqVOY57fZ3OzBzKxupbvyTEarmpR-GcUc-qPyF9OPfUWPIqnNs050FRQpWR6UNVf/s1600/Malayotyphlops+Wynn+et+al+2016+Figs+3+4+5.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="225" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEghm_Rs4lti4jslM5Gyspd5cvQoua3f7O0MavizvN8nMq5HVYwYotQpZps0BIFefPWJprp_pxSF_wLuzqVOY57fZ3OzBzKxupbvyTEarmpR-GcUc-qPyF9OPfUWPIqnNs050FRQpWR6UNVf/s400/Malayotyphlops+Wynn+et+al+2016+Figs+3+4+5.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Malayotyphlops luzonensis </i>(L), <i>M. denrorum </i>(C), and <i>M. andyi </i>(R)<br />
From <a href="https://www.researchgate.net/profile/Rafe_Brown/publication/297599848_Two_New_Species_of_Malayotyphlops_from_the_Northern_Philippines_with_Redescriptions_of_Malayotyphlops_luzonensis_Taylor_and_Malayotyphlops_ruber_Boettger/links/575496d508ae6807fb04d068.pdf">Wynn et al. 2016</a></td></tr>
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As if that wasn't strange enough, there is a third possible candidate member of Gerrhopilidae: the species known as either <i>Typhlops manilae</i>, <i>Malayotyphlops manilae</i>, or <i>Gerrhopilus manilae</i>. The taxonomic status of this species is currently unclear. It was described by <a href="http://www.nature.com/news/taxonomy-the-spy-who-loved-frogs-1.13710?WT.mc_id=FBK_NatureNews">American herpetologist and spy Edward H. Taylor</a> in 1919, from a specimen that was <a href="http://www.biodiversitylibrary.org/item/209613#page/144/mode/2up">"discovered in the Santo Tomas Museum"</a> in Manila, although even then nobody knew when, where, or by whom it was collected. It appears to have been barely mentioned in the scientific literature until 2014, when its morphological distinctiveness from other members of the <i>Typhlops ater </i>species group/Gerrhopilidae was noted as part of <a href="https://www.researchgate.net/profile/Nicolas_Vidal/publication/259757280_A_taxonomic_framework_for_Typhlopid_snakes_from_the_Caribbean_and_other_regions_Reptilia_Squamata/links/00b4952da4869e5ecd000000.pdf">a massive review of typhlopid snakes</a> led by Pennsylvania State University blindsnake specialist and evolutionary biologist <a href="http://www.hedgeslab.org/">Blair Hedges</a>. They suggested it belonged instead to another new genus, <i>Malayotyphlops</i>, also mostly from the Philippines, because it has 28 scale rows (vs. 18 in <i>Gerrhopilus</i>) and a short tail, and because a subocular scale is not unique to <i>Gerrhopilus</i>. Later the same year, <a href="https://www.researchgate.net/profile/Van_Wallach/publication/264433188_strongSystematics_of_the_blindsnakes_Serpentes_Scolecophidia_Typhlopoidea_based_on_molecular_and_morphological_evidencestrong/links/552539420cf2561f2ac16adb/strongSystematics-of-the-blindsnakes-Serpentes-Scolecophidia-Typhlopoidea-based-on-molecular-and-morphological-evidence-strong.pdf?origin=publication_detail&ev=pub_int_prw_xdl&msrp=CH5Y5UR9SGnZbgAAEStY3_LOWOMlNouCcBYbsnB6Tj8wY2GnjSnzkCVVDe_9ekaGDls1xTZDerlZ2L10WJ3wdb2-sDw1T8BtPuKqrWA6g0k.F42ZNgBXhLDzvim9NCNw6IeRUL-qWPWl35Z7eCm9bAkGDMzSXLMpIdj5tuZ7_Ws1Yzx0T5qqdGO39_HxaHw0RA.jM7mosZnSt8jy5a58Si48d9x7GNPx5bteYbpyAJde8E8rv9L6sVys7PCu9OJzYKzlkWz0s33ah2Yz8AdUTLlQA.7lRc_DzNYwbNC4AcFsHxV08iB64UPquxK5shmRTwMrf009Wh7rUfe2Nos2EXiu8NmpHB8drOFF221ksq4lqvhg">a different study</a> disagreed and moved the species back to <i>Gerrhopilus </i>based on the statement from the original description that it has a subocular. However, <a href="https://www.researchgate.net/profile/Rafe_Brown/publication/297599848_Two_New_Species_of_Malayotyphlops_from_the_Northern_Philippines_with_Redescriptions_of_Malayotyphlops_luzonensis_Taylor_and_Malayotyphlops_ruber_Boettger/links/575496d508ae6807fb04d068.pdf">yet a third study</a> took a close look at Taylor's original description, which contains no illustration, and noted several areas of potential confusion, concluding that without examination of the original specimen, which is still in Manila, "it is not possible to determine to which genus, or even family, <i>T. manilae...</i>belongs".</div>
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhAuQvprwq7uVrLvtLRZn22X9qfa_NGgrT19ZKC82uwXptqORV0os4ANuEvaNt6Jf6FWngf5aL75pc3dwu9BpEkywjgRJTK62ouNRiMKwsz2y6XwwUYXEniJQIUue9iBsUim-aYuc05k0Ro/s1600/Xenotyphlops+grandidieri+Wegener+et+al+2014+Fig1.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="248" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhAuQvprwq7uVrLvtLRZn22X9qfa_NGgrT19ZKC82uwXptqORV0os4ANuEvaNt6Jf6FWngf5aL75pc3dwu9BpEkywjgRJTK62ouNRiMKwsz2y6XwwUYXEniJQIUue9iBsUim-aYuc05k0Ro/s320/Xenotyphlops+grandidieri+Wegener+et+al+2014+Fig1.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">The three reptile species originally described by Mocquard<br />
and re-discovered at Baie de Sakalava in northern Madagascar<br />
after more than 100 years without records.<br />
The blindsnake <i>Xenotyphlops grandidieri </i>(pink), and two<br />
legless skink species: <i>Paracontias minimus</i> (brown with<br />
longitudinal lines of dark spots) and <i>P. rothschildi</i><br />
(beige with black flanks). From <a href="https://www.researchgate.net/profile/Zoltan_Nagy4/publication/259339506_Morphological_variation_and_taxonomic_reassessment_of_the_endemic_Malagasy_blind_snake_family_Xenotyphlopidae_Serpentes_Scolecophidia/links/0c96052b198f65aa56000000/Morphological-variation-and-taxonomic-reassessment-of-the-endemic-Malagasy-blind-snake-family-Xenotyphlopidae-Serpentes-Scolecophidia.pdf">Wegener et al. 2013</a></td></tr>
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Before you get too discouraged, remember that <a href="http://snakesarelong.blogspot.com/2014/05/snakes-long-lost.html">snake biology is replete with tales of rediscovery</a>. Case in point: the other "new" family, Xenotyphlopidae. This bizarre snake has completely lost any traces of visible eyes. It was known solely from the type specimens, described by French zoologist François Mocquard in 1905 and 1906, for more than 100 years. Their precise locality was unknown. However, <a href="http://brownanole.weebly.com/people.html">Hanna Wegener</a> and a term of German, Belgian, and American herpetologists <a href="https://www.researchgate.net/profile/Zoltan_Nagy4/publication/259339506_Morphological_variation_and_taxonomic_reassessment_of_the_endemic_Malagasy_blind_snake_family_Xenotyphlopidae_Serpentes_Scolecophidia/links/0c96052b198f65aa56000000/Morphological-variation-and-taxonomic-reassessment-of-the-endemic-Malagasy-blind-snake-family-Xenotyphlopidae-Serpentes-Scolecophidia.pdf">rediscovered</a> <i>Xenotyphlops</i> in 2013<i> </i>on a coastal dune under a piece of wood in the sand in a littoral forest at Baie de Sakalava in northern Madagascar, along with two endemic legless skinks in the genus <i>Paracontias</i> also described by Mocquard. Because the new specimens of <i>X. grandidieri </i>overlapped the other species in this genus (<i>X. mocquardi</i>) in most morphological characteristics, the two have now been synonymized, making the family Xenotyphlopidae monotypic (for now). <a href="http://pubmedcentralcanada.ca/pmcc/articles/PMC2936224/pdf/rsbl20100220.pdf">These blindsnakes are unique</a> in having a greatly enlarged and nearly circular rostral scale and an enlarged anal shield, and in lacking a tracheal lung.</div>
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The number of less-phylogenetically-distinct but poorly-known blindsnakes is not small. These have received renewed attention due to <a href="http://pubmedcentralcanada.ca/pmcc/articles/PMC2936224/pdf/rsbl20100220.pdf">their placement in new families</a>, but the 21st century blindsnake revolution is just getting started.</div>
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<a href="https://www.blogger.com/null" name="1"><b>1 </b></a>Péron and Lesueur also collected the first and some of the only specimens of <i><a href="https://snakesarelong.blogspot.com/2012/04/round-island-splitjaw-snakes.html">Bolyeria multicarinata</a></i> from Mauritius, which is now thought to be extinct, although they mistakenly labeled it as being from Australia.<a href="#top1"><sup>↩</sup></a><br />
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<a href="https://www.blogger.com/null" name="2"><b>2 </b></a>Today, only <i>I. braminus</i> and another introduced species, <i>I. porrectus</i>, are found on Mauritius; the latter may have also been introduced in the 1800s but was first conclusively documented only in 1993.<a href="#top2"><sup>↩</sup></a><br />
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<span class="Apple-style-span" style="font-size: x-small;"><br />
<a href="https://www.blogger.com/null" name="3"><b>3 </b></a>A few pieces of evidence against: a length of 4–5 French inches corresponds to 109–136 mm, which is just right for <i>I. braminus</i> but a tad small for the <i>Cathetorhinus </i>specimen, which measures 178 mm (6.6 French inches). <a href="https://www.researchgate.net/profile/Anthony_Cheke/publication/281839405_Is_the_enigmatic_blind-snake_Cathetorhinus_melanocephalus_Serpentes_Typhlopidae_an_extinct_endemic_species_from_Mauritius/links/55faaaf208aec948c4ac89f7.pdf?origin=publication_list">Cheke thought that</a> "Lesueur appeared to be writing from memory without the specimen actually before him, so, impressed by its small size, he may have exaggerated how tiny his snake actually was.", maybe the last time in history that somebody underestimated the size of a snake. The other point of confusion is over the exact locality: Lesueur and Péron were clearing land with an upland planter, Toussaint de Chazal, at whose estate in the area now known as Mondrain they were staying. Mondrain is on a plateau adjacent to the Tamarin Gorge, which is 9 km from Grand Bassin, where Lesueur stated that they found the snake.<a href="#top3"><sup>↩</sup></a><br />
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ACKNOWLEDGMENTS</div>
<br />
Thanks to <a href="https://www.flickr.com/photos/15023943@N05/5824871337/">Tim Colston</a>, <a href="http://www.arkive.org/jans-worm-snake/typhlops-mirus/">Ruchira Somaweera</a> and <a href="http://reptile-database.reptarium.cz/species?genus=Gerrhopilus&species=beddomii">Sumaithangi Ganesh</a> for the use of their photos.<br />
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REFERENCES</div>
<br />
<div style="text-align: justify;">
<span style="font-size: x-small;">Cheke, A. 2010. Is the enigmatic blind snake <i>Cathetorhinus melanocephalus </i>(Serpentes: Typhlopidae) an extinct endemic species from Mauritius? Hamadryad 35:101-104 <<a href="https://www.researchgate.net/profile/Anthony_Cheke/publication/281839405_Is_the_enigmatic_blind-snake_Cathetorhinus_melanocephalus_Serpentes_Typhlopidae_an_extinct_endemic_species_from_Mauritius/links/55faaaf208aec948c4ac89f7.pdf?origin=publication_list">full-text</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Duméril, C., G. Bibron, and A. Duméril. 1854. Erpetologie Générale on Histoire Naturelle Compléte des Reptiles. Librairie Encyclopédique de Roret, Paris <<a href="http://www.biodiversitylibrary.org/item/101228#page/286/mode/1up">link to <i>Cathetorhinus</i> description</a>></span></div>
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<span style="font-size: x-small;">Hedges, S., A. Marion, K. Lipp, J. Marin, and N. Vidal. 2014. A taxonomic framework for typhlopid snakes from the Caribbean and other regions (Reptilia, Squamata). Caribbean Herpetology 49:1-61 <<a href="https://www.researchgate.net/profile/Nicolas_Vidal/publication/259757280_A_taxonomic_framework_for_Typhlopid_snakes_from_the_Caribbean_and_other_regions_Reptilia_Squamata/links/00b4952da4869e5ecd000000.pdf">full-text</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Kraus, F. 2005. New species of blindsnake from Rossel Island, Papua New Guinea. Journal of Herpetology 39:591-595 <<a href="https://www.jstor.org/stable/4092848">abstract</a>></span></div>
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<span style="font-size: x-small;">Pyron, R. and V. Wallach. 2014. Systematics of the blindsnakes (Serpentes: Scolecophidia: Typhlopoidea) based on molecular and morphological evidence. Zootaxa 3829:1-81 <<a href="https://www.researchgate.net/publication/264433188_strongSystematics_of_the_blindsnakes_Serpentes_Scolecophidia_Typhlopoidea_based_on_molecular_and_morphological_evidencestrong">full-text</a>></span></div>
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<span style="font-size: x-small;">Taylor, E. H. 1919. New or rare Philippine reptiles. Philippine Journal of Science 14:105-125 <<a href="http://www.biodiversitylibrary.org/item/209613#page/144/mode/2up">full-text</a>></span></div>
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<span style="font-size: x-small;">Vidal, N., J. Marin, M. Morini, S. Donnellan, W. R. Branch, R. Thomas, M. Vences, A. Wynn, C. Cruaud, and S. B. Hedges. 2010. Blindsnake evolutionary tree reveals long history on Gondwana. Biology Letters 6:558-561 <<a href="http://pubmedcentralcanada.ca/pmcc/articles/PMC2936224/pdf/rsbl20100220.pdf">full-text</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Wallach, V. 1996. Two new Blind snakes of the <i>Typhlops ater</i> species group from Papua new Guinea (Serpentes: Typhlopidae). Russian Journal of Herpetology 3:107-118 <<a href="https://www.researchgate.net/profile/Van_Wallach/publication/277038209_Two_New_Blind_Snakes_of_the_Typhlops_ater_Species_Group_from_Papua_New_Guinea_Serpentes_Typhlopidae/links/570826dc08aed09e916d24a8.pdf">full-text</a>></span><br />
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<span style="font-size: x-small;">Wallach, V. and O. Pauwels. 2008. The systematic status of <i>Cathetorhinus melanocephalus</i> Duméril & Bibron, 1844 (Serpentes: Typhlopidae). Hamadryad 33:39-47 <<a href="https://www.researchgate.net/profile/Olivier_Pauwels/publication/236983437_The_systematic_status_of_Cathetorhinus_melanocephalus_Dumeril_Bibron_1844_Serpentes_Typhlopidae/links/02e7e51ac7d54d25cd000000/The-systematic-status-of-Cathetorhinus-melanocephalus-Dumeril-Bibron-1844-Serpentes-Typhlopidae.pdf?origin=publication_detail&ev=pub_int_prw_xdl&msrp=cGRCmIMcir9sD1AtjG4BAu6Up2PBlq_Nm_D00LAF6Q0MAsqtVkae9C1QrL7fIPuJqLn73Cl1A1zfiqbDXLN3jhZiZGPCsZbESKGrtIiZlxs.XHJwWSAnXm9BaKBbMjVJthuW7gX4XryX0G4MasxcxF0CmLt7wNXNTc7oCvQuzbr7M5ol-U26bGaNbQGHEmkLzQ.I47bax12KYTyDPgzn-jhcYc2gj4iDJ_PKY_JVXzIecWc_y80R0LC2wbK8X-EhN_2NNI2n2UfSOSusrURCBne9A.zNRpW3j16R7Vwr5lgFbDJWVqjfa6mSr5V3wW4XXTuEXBrC0PKwiO92j97URY3yFPKIlrm1ik2nxZnNdUNEGgng">full-text</a>></span></div>
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<span style="font-size: x-small;">Wegener, J. E., S. Swoboda, O. Hawlitschek, M. Franzen, V. Wallach, M. Vences, Z. T. Nagy, S. B. Hedges, J. Köhler, and F. Glaw. 2013. Morphological variation and taxonomic reassessment of the endemic Malagasy blind snake family Xenotyphlopidae. Spixiana 36:269-282 <<a href="https://www.researchgate.net/profile/Zoltan_Nagy4/publication/259339506_Morphological_variation_and_taxonomic_reassessment_of_the_endemic_Malagasy_blind_snake_family_Xenotyphlopidae_Serpentes_Scolecophidia/links/0c96052b198f65aa56000000/Morphological-variation-and-taxonomic-reassessment-of-the-endemic-Malagasy-blind-snake-family-Xenotyphlopidae-Serpentes-Scolecophidia.pdf">full-text</a>></span></div>
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<span style="font-size: x-small;">Wynn, A. H., R. P. Reynolds, D. W. Buden, M. Falanruw, and B. Lynch. 2012. The unexpected discovery of blind snakes (Serpentes: Typhlopidae) in Micronesia: two new species of <i>Ramphotyphlops </i>from the Caroline Islands. Zootaxa 3172:39–54 <<a href="https://www.fs.fed.us/psw/publications/falanruw/psw_2012_falanruw001_wynn.pdf">full-text</a>></span></div>
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<span style="font-size: x-small;">Wynn, A. H., A. C. Diesmos, and R. M. Brown. 2016. Two new species of <i>Malayotyphlops </i>from the northern Philippines, with redescriptions of <i>Malayotyphlops luzonensis</i> (Taylor) and <i>Malayotyphlops ruber</i> (Boettger). Journal of Herpetology 50:157-168 <<a href="https://www.researchgate.net/profile/Rafe_Brown/publication/297599848_Two_New_Species_of_Malayotyphlops_from_the_Northern_Philippines_with_Redescriptions_of_Malayotyphlops_luzonensis_Taylor_and_Malayotyphlops_ruber_Boettger/links/575496d508ae6807fb04d068.pdf">full-text</a>></span><br />
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="https://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
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Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com4Jena, Germany50.927054 11.58923720000007150.7669025 11.26651370000007 51.087205499999996 11.911960700000071tag:blogger.com,1999:blog-7443075087825368900.post-39333537222964448542017-03-31T15:43:00.000-06:002018-01-04T16:42:32.179-07:00Snakebite, antivenom research, and basic science<div style="text-align: justify;">
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<span style="font-size: x-small;"><a href="https://drive.google.com/file/d/1_bkugShekOqzGO9neTox0eNSBfYrTBZZ/view?usp=sharing">Haga clic aquí para leer en español</a></span><br />
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In the past few weeks, a peculiar congruence of several seemingly-unrelated events took place. (At least) two new scientific papers about snake biology were published, a new video series was announced, some scientists entered contests, and the U.S. executive branch announced <a href="https://www.wired.com/2017/03/trumps-budget-break-american-science-today-tomorrow/">a budget proposal with deep cuts to science funding</a>. However, these events aren't as unrelated at they might seem at first glance, and they have something to tell us about where snake biology, and science in general, are going in the future.<br />
<b><i><br /></i></b>
<b><i>The science: part I (puff adders)</i></b><br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhnemC1o5qVy-EoaovmfCW9E2SsT9i-1BC1AR8QjLUbsGasMIzFNk4vOvoSrk1kaJn_XfLpJm69fbpOwYF1qNMHH2i3LQ4lhkJ_9_ArcR3mht6-ekmgKRWV7PMrvTv7Y-ih9CIjJ7G6zvwi/s1600/Bitis+arietans+Markus+Oulehla.jpg" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="213" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhnemC1o5qVy-EoaovmfCW9E2SsT9i-1BC1AR8QjLUbsGasMIzFNk4vOvoSrk1kaJn_XfLpJm69fbpOwYF1qNMHH2i3LQ4lhkJ_9_ArcR3mht6-ekmgKRWV7PMrvTv7Y-ih9CIjJ7G6zvwi/s320/Bitis+arietans+Markus+Oulehla.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">A puff adder (<i>Bitis arietans</i>)</td></tr>
</tbody></table>
Puff Adders (<i>Bitis arietans</i>) are <a href="http://snakesarelong.blogspot.com/2013/01/africas-giant-gaboon-vipers.html">among Africa's most widespread vipers</a>. They are heavy-bodied snakes that are found in savannas and open woodlands. Like most vipers, they eat mostly rodents as adults, which they ambush from carefully-selected sites, <a href="https://www.researchgate.net/profile/Xavier_Glaudas/publication/311166745_Food_supplementation_affects_the_foraging_ecology_of_a_low-energy_ambush-foraging_snake/links/583ed06908ae8e63e617e3cc.pdf">which they sometimes occupy for weeks at a time</a>. Recently, <a href="https://xavierglaudas.wordpress.com/">Xavier Glaudas</a> and <a href="https://www.wits.ac.za/apes/staff/academic-staff/alexander/prof-graham-alexander/">Graham Alexander</a> published <a href="http://onlinelibrary.wiley.com/doi/10.1111/jzo.12450/full">a new study</a> showing that, even though Puff Adder strikes last less than two seconds, they can choose to either hold onto or let go of the prey depending on its size. Specifically, they hold onto small mice, <a href="http://onlinelibrary.wiley.com/store/10.1111/jzo.12450/asset/supinfo/jzo12450-sup-0004-VideoS4.mp4?v=1&s=b5c4d4c8c890b802d9e1433a87d23f4a0a74b3ba">shrews</a>, <a href="http://onlinelibrary.wiley.com/store/10.1111/jzo.12450/asset/supinfo/jzo12450-sup-0006-VideoS6.mp4?v=1&s=3d92dd175c13b47de4afb470323064fba49d742c">birds</a>, <a href="http://onlinelibrary.wiley.com/store/10.1111/jzo.12450/asset/supinfo/jzo12450-sup-0003-VideoS3.mp4?v=1&s=70b437f3c1348ce6e905af122e52234637099e17">toads</a>, and <a href="http://onlinelibrary.wiley.com/store/10.1111/jzo.12450/asset/supinfo/jzo12450-sup-0002-VideoS2.mp4?v=1&s=d4ba3c7889913e9ec7ee75ef668ace3e52b99972">lizards</a>, but strike & <a href="http://onlinelibrary.wiley.com/store/10.1111/jzo.12450/asset/supinfo/jzo12450-sup-0012-VideoS12.mp4?v=1&s=5cec6c4abbbf3f0bf33a6f0bb9e3068f8138b111">release larger rodents</a> and <a href="http://onlinelibrary.wiley.com/store/10.1111/jzo.12450/asset/supinfo/jzo12450-sup-0013-VideoS13.mp4?v=1&s=79727f3d65399bfc75394fc08944f376911f5bd2">rabbits</a>, because retaliatory <a href="http://onlinelibrary.wiley.com/store/10.1111/jzo.12450/asset/supinfo/jzo12450-sup-0020-VideoS20.mp4?v=1&s=710ddd81ee33af2272403e7132bf13e4b208ae6b">rat bites are dangerous to them</a>. After they let go of these larger prey, which usually run off a short distance before the venom kills them, they track them down again using stereotypic strike-induced chemosensory searching behavior to locate <a href="http://snakesarelong.blogspot.de/2013/03/non-toxic-venoms.html">the scent of non-toxic components of their own venom</a>. This is really similar to findings by <a href="https://www.breeputman.com/">Bree Putman</a> and <a href="http://www.bio.sdsu.edu/pub/clark/Site/Home.html">Rulon Clark</a> that <a href="https://media.wix.com/ugd/8f1659_b5ae072145384e8e8a2d5faffcf9119f.pdf">Southern Pacific Rattlesnakes (<i>Crotalus oreganus</i>) were more likely to hold onto smaller rodents than to larger ground squirrels</a>. You can watch <a href="http://onlinelibrary.wiley.com/doi/10.1111/jzo.12450/full">26 awesome videos</a> selected from an archive of thousands of hours of video taken in the wild over more than two years.<a href="#1" name="top1"><sup>1</sup></a><br />
<br />
This research matters because venomous snakes and their prey are in constant evolutionary arms races, leading to:<br />
<ol>
<li>a mosaic of new biochemical compounds <a href="http://ngm.nationalgeographic.com/2013/02/125-venom/holland-text">that are often useful in treating disease</a></li>
<li>a mosaic of new biochemical compounds <a href="http://med.fsu.edu/userFiles/file/TSL/Proteomics__Margres_et_al,_Genetics_(2015)_199_p165_176.pdf">that can make venomous snakebite really hard to treat</a></li>
</ol>
We'll come back to the second one in a minute. The obvious importance of human medicine and venomous snakebite treatment overshadow a third important reason to study snakes and what they eat. Although the beneficial role of snakes in rodent control is taken as gospel by many advocates of snake conservation, the amount of data that we actually have on what snakes eat in the wild is surprisingly small. For many species, we don't even have a general idea of what kinds of prey they like to eat. Given <a href="https://link.springer.com/article/10.1007%2Fs00114-017-1440-1">recent estimates that spiders eat about as much meat as people do worldwide</a>, and the <a href="http://comp.uark.edu/~jwillson/publications/2015%20-%20Willson%20and%20Winne%20-%202015%20-%20JZL%20-%20role%20of%20aquatic%20snakes.pdf">potential for snakes to reach very high population densities in certain habitats</a>, it's likely that the top-down effects of snakes as predators are significant ecosystem services that most humans aren't aware of and thus undervalue. Indirect effects on other aspects of the ecology of snake prey species, such as <a href="http://comp.uark.edu/~jwillson/publications/2017%20-%20Willson-2017-JAE%20-%20python%20indirect.pdf">predation release</a> and <a href="https://eco.confex.com/eco/2013/webprogram/Paper44305.html">disease transmission</a>, link snake predation even more strongly to human health. This is particularly timely in light of <a href="http://www.npr.org/sections/goatsandsoda/2017/03/06/518219485/forbidding-forecast-for-lyme-disease-in-the-northeast">recent predictions that 2017 will be a big year for white-footed mice and thus for Lyme disease</a> in the northeastern USA, <a href="http://www.nybooks.com/daily/2016/08/18/fear-of-rattlesnake-island-massachusetts/">controversy over</a> the reintroduction of Timber Rattlesnakes, one of the white-footed mouse's top predators, <a href="http://wwlp.com/tag/quabbin-rattlesnakes/">to Quabbin Island in Massachusetts</a><a href="#2" name="top2"><sup>2</sup></a>, and the <a href="http://snakesarelong.blogspot.de/2015/03/rattlesnake-roundups.html">continuation</a> of both the <a href="https://soundcloud.com/joe-phelan-194475994/from-the-ground-up-08-sweetwater-rattlesnake-roundup">infamous Sweetwater Rattlesnake Roundup</a><a href="#3" name="top3"><sup>3</sup></a> and the <a href="http://claxtonenterprise.com/festival-attendance-tips-16000/">reformed Claxton Wildlife Festival</a> and <a href="http://rattlesnakepreservationtrust.org/index.php/lone-star-rattlesnake-days">Lone Star Rattlesnake Days</a> earlier this month.<br />
<b><i><br /></i></b>
<b><i>The science: part II (how cobras got their flesh-eating venoms)</i></b><br />
<br />
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiaJXk7ThfIsXpLpV8SGL89GcWGUHH8Ls0P3a_JJhSkvPMxe8aTxZpbe3N_3Ne_Mis3K-YY0QyqWeGkTJNsB7pL7HCM8qycgSs5ziZ5oSTMH96PTezedGKSX2IuxMi6IkTtpddeWUO2ZsQq/s1600/Naja+mossambica+Colin+Donihue.jpg" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="265" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiaJXk7ThfIsXpLpV8SGL89GcWGUHH8Ls0P3a_JJhSkvPMxe8aTxZpbe3N_3Ne_Mis3K-YY0QyqWeGkTJNsB7pL7HCM8qycgSs5ziZ5oSTMH96PTezedGKSX2IuxMi6IkTtpddeWUO2ZsQq/s400/Naja+mossambica+Colin+Donihue.jpg" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">A Mozambique spitting cobra (<i>Naja mossambica</i>) spitting its venom</td></tr>
</tbody></table>
<a href="http://www.mdpi.com/2072-6651/9/3/103"></a><a href="http://snakesarelong.blogspot.com/2015/04/spitting-cobras.html">Spitting cobras</a> are even more well-known than puff adders because of their defensive venom spitting abilities, showcased on the <a href="https://www.youtube.com/watch?v=JiCqXfHIpVY">BBC's </a><a href="https://www.youtube.com/watch?v=JiCqXfHIpVY" style="font-style: italic;">Life in Cold Blood</a>. They are<i> </i>found in Africa and Asia and are thought to have evolved two or three times from non-spitting cobras. <a href="http://www.mdpi.com/2072-6651/9/3/103">A new paper</a> from the lab of <a href="http://www.venomdoc.com/">Bryan Fry at the University of Queensland</a> sheds some light on when and why venom spitting evolved. Elapid snakes, including cobras, have venoms rich in neurotoxins, which are highly potent toxins that are very effective at paralyzing their prey. Cobras also have less potent cytotoxins that kill cells directly, which is a bit weird. What is the function of these toxins?<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhO4ioYjaoWE4u1aC3Rm-XqRIG7D0VP8r5yJe3JGBbQ3WG0gQR4yp8VBleZ78kBFAV-9SDnmqe54xrR50ywKLjAjxHU_InElunyZQnnQGbaWGtwLZFU8OaVM-bbzwDc-j3c8_TnOTaK-h4y/s1600/Panagides+et+al+2017+Fig1.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="400" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhO4ioYjaoWE4u1aC3Rm-XqRIG7D0VP8r5yJe3JGBbQ3WG0gQR4yp8VBleZ78kBFAV-9SDnmqe54xrR50ywKLjAjxHU_InElunyZQnnQGbaWGtwLZFU8OaVM-bbzwDc-j3c8_TnOTaK-h4y/s400/Panagides+et+al+2017+Fig1.png" width="258" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Toxicity of snake venom to human cells grown in culture.<br />
Warm colors indicate higher toxicity.<br />
From <a href="http://www.mdpi.com/2072-6651/9/3/103">Panagides et al. 2017</a></td></tr>
</tbody></table>
The hypothesis put forth here is that the first step towards venom spitting was the evolution of hooding behavior and morphology, which happened twice in elapids: once in "regular" cobras and once in King Cobras, which are more closely related to mambas. Only once a conspicuous visual display was present was there selective pressure for cytotoxic venom components delivered to the eyes of potential predators via spitting. Although the venom of both groups is cytotoxic, <i>Hemachatus</i> (rinkhals) and <i>Naja </i>cobras use three-finger toxins, whereas King Cobras use L-amino acid oxidase enzymes, consistent with the undirected, opportunistic nature of <a href="http://phys.org/news/2013-12-rapid-evolution-cobra-venom.html">our current model of venom evolution by gene duplication and mutation</a>. The authors suggest that further elevations in cytotoxicity are linked to bright bands and other aposematic colors or hood markings, although their paper did not attempt to quantify these attributes of cobra displays, which can be quite diverse even within species. Further evidence in support of the hypothesis is that <i>Naja naja</i> and <i>Naja oxiana</i> seem, based on their nested position, to have lost spitting but to have retained cytotoxicity, and their close relatives <i>Naja atra</i> and <i>Naja kaouthia</i> might represent steps down this evolutionary path, being capable of spitting only in some populations and with less accuracy than the African and southeast Asian clades of true spitting cobras.<br />
<br />
This is an extremely cool and popular topic. It was covered by <a href="http://www.iflscience.com/plants-and-animals/how-cobras-learned-to-spit-venom/">IFLS</a>, <a href="https://thewire.in/117248/cobra-venom-cytotoxin-cancer/">The Wire</a>, <a href="http://gizmodo.com/the-deadliest-cobras-also-look-the-most-terrifying-1793269750">Gizmodo</a>, and <a href="https://www.washingtonpost.com/news/morning-mix/wp/2017/03/15/evolutionary-study-unravels-mystery-of-how-cobras-evolved-their-flesh-eating-venom/">the Washington Post</a>. It goes to show that people worldwide are fascinated by venomous snakes, and the Fry lab has done a great job capitalizing on that interest (among other accolades, Fry's graduate student <a href="https://www.facebook.com/JordiDebono/">Jordan Debono</a> recently won the <a href="https://www.thinkable.org/submission_entries/D96ywN8Y">Queensland Women in Science Peoples' Choice Award</a> [a contest that was decided by an online popular vote; more on this later] for her research on <a href="http://snakesarelong.blogspot.com/2013/11/the-truth-about-snakebite.html">global snakebite</a> treatments). One reason for this fascination has to do with the question of who, exactly, these cobras are defending themselves from? The most reasonable hypothesis, given the timing and geography of the diversification of spitting cobras and the precision with which they can target forward-facing eyes and hominoid faces, is primates. Us, and our ancestors, <a href="https://pdfs.semanticscholar.org/cc06/7d26152799da8c55570e2fb560b9ca794cb9.pdf">who have eaten and been eaten by snakes for millions of years</a>. Studying spitting cobras is a window into our own evolutionary past, a way for us to learn about ourselves. But, let us not be misled into thinking that interactions between humans and cobras are a thing of the past.</div>
<div style="text-align: justify;">
<b><i><br /></i></b>
<b><i>The upshot: the truth about snakebite</i></b><br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiDP9rrNtmSiq7Pg650zPFlpTNIgePAphyy4RZ3vKiBViIzdcHbhS7yJBeJ9z0mRKlCRlvTOde1GwwmS-X56rvddo29DfiiF41AqdvT2RL2G1NpxoHfGzCqil9V7YrEI9F2xoeIwmkzmnlx/s1600/venimologie.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiDP9rrNtmSiq7Pg650zPFlpTNIgePAphyy4RZ3vKiBViIzdcHbhS7yJBeJ9z0mRKlCRlvTOde1GwwmS-X56rvddo29DfiiF41AqdvT2RL2G1NpxoHfGzCqil9V7YrEI9F2xoeIwmkzmnlx/s320/venimologie.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="font-size: 12.8px; text-align: center;">You can follow the ASV <a href="https://twitter.com/venimologie">@Venimologie</a></td></tr>
</tbody></table>
If you haven't read <a href="http://scorpiondoc.silvrback.com/">the blog by medical toxinologist Leslie Boyer</a>, you really should. Earlier this month she wrote about the <a href="http://scorpiondoc.silvrback.com/the-vicious-circle-of-antivenom-shortage">vicious circle of antivenom shortage in sub-Saharan Africa</a>, where millions of people are bitten by venomous snakes every year, <a href="http://www.nature.com/nbt/journal/v25/n2/full/nbt0207-173.html">many of which die or suffer awful injuries</a> because they lack access to good antivenom. This crisis has prompted the creation of the <a href="http://www.sav-asv.org/en/home-2/">African Society of Venimology</a> and a <a href="https://scorpiondoc.silvrback.com/introducing-a-new-series-of-snakebite-training-videos">new series of snakebite training videos</a> in English, French, and Spanish. The politics and economics of antivenom are complicated and reflect larger issues in medicine, education, quality control, supply and demand, and how global economics and corporations have failed to respond to the needs of local communities and consumers. In a nutshell, the issue is that antivenom manufacturers don't make enough good antivenom, because not enough people buy it. People don't buy it because it's expensive, and it's expensive because not that much is made. This is despite a huge need for it—but not everybody with a snakebite goes to a hospital and gets antivenom in Africa, partially because it's not certain there will be any and partially because a lot of patients and doctors don't know about antivenom, because it's not in widespread use (which is mostly because of the reasons above). Other exacerbating problems include that it's often not certified, fake products can price the real antivenom out of the market, and the infrastructure for distributing antivenom and information in Africa is sub-optimal (but improving). Fixing any one or even most of these problems won't fix the whole system—if any one of them break down, supply and demand will be out of balance and people won't get the care they need.<br />
<br />
A lot of the same issues used to be present in Mexico, but product improvements, government outreach, and massive education efforts in the 1980s and 1990s dramatically reduced mortality from venomous snakebite and led Mexico to become <a href="http://www.npr.org/2011/08/06/139023806/mexico-to-the-rescue-in-americas-venom-belt">a major producer and consumer of high-quality, affordable antivenom</a>, so much so that the USA now imports some of these drugs from Mexico. The Mexican government enabled the Mexican antivenom industry to be competitive and reach its market, which is much larger than the domestic market for American antivenom manufacturers—medically-serious venomous snakebites (and scorpion stings) in the USA are mostly confined to the southwest, and <a href="https://www.blogger.com/snakesarelong.blogspot.com/2013/11/the-truth-about-snakebite.html">the per-capita risk of snakebite is the lowest in the world</a>. This creates its own unique problems. You may have heard about the controversy surrounding <a href="http://www.oriannesociety.org/blog/coral-snake-antivenom">the discontinued coralsnake antivenom</a> made by Wyeth, and there are compelling arguments that the Mexican polyvalent antivenoms Anavip (made by Bioclon for humans) and ViperSTAT (made by Veteria Labs for cats and dogs) are <a href="https://www.researchgate.net/profile/Leslie_Boyer/publication/255957134_Subacute_coagulopathy_in_a_randomized_comparative_trial_of_Fab_and_Fab%272_antivenoms/links/00b7d52332e01c1d08000000/Subacute-coagulopathy-in-a-randomized-comparative-trial-of-Fab-and-Fab2-antivenoms.pdf">more effective</a> and <a href="https://www.researchgate.net/profile/Leslie_Boyer/publication/281226188_On_1000-fold_pharmaceutical_price_markups_and_why_drugs_cost_more_in_the_US_than_in_Mexico/links/55de5ebd08ae45e825d39d4b.pdf">much less expensive (although this is due almost exclusively to the idiosyncrasies of the US healthcare finance system)</a> than the only FDA-approved viper antivenom, CroFab (although BTG, the maker of CroFab, <a href="http://opa.ahsc.arizona.edu/sites/opa.ahsc.arizona.edu/files/uploads/science-2014-wade-16-72.pdf">filed a complaint asserting that these Mexican products infringe on its patent</a>).<br />
<br />
Finally, the global importance of the availability of high-quality, affordable antivenom for Latin American, African, and other exotic snakes is only going to increase as venomous snakes become more popular as pets and in zoos. This is particularly true in parts of the world <a href="https://snakesarelong.blogspot.com/2015/10/are-there-any-countries-without-snakes.html">completely lacking venomous snakes</a> or with only very benign, non-life-threatening species, such as <a href="http://vipersgarden.at/PDF/snakebites2011a.PDF">northern Europe</a>, <a href="https://lup.lub.lu.se/search/ws/files/2851010/623719.pdf">Scandinavia</a> and northern North America, where doctors may be totally unprepared for a snakebite emergency and may not have appropriate antivenom on hand. This is exactly the kind of situation where government funding, in the form of <a href="https://rarediseases.info.nih.gov/guides/pages/124/finding-funding-opportunities">orphan disease R&D grants</a>, could play a role in making it affordable for researchers and doctors to save lives.<br />
<br />
For a great introduction to and more in-depth coverage of these issues, you should watch <a href="http://thevenominterviews.com/#section-contents">The Venom Interviews</a> or read <a href="http://thevenominterviews.com/2017/03/11/solving-snakebite-in-sub-saharan-africa/">their coverage of the recent video series</a>.<br />
<br />
<b><i>The future: <a href="http://www.pacb.com/smrt-science/smrt-grant/pag2017/">sequence the Temple Pitviper genome</a></i></b><br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg9TmSYbe_c6AMVSpIpW0f69hVgjBkQu6SLXDPA2Q8OtHEq8ha7lzUDj4TVK8fZZRm7X_IereKNx2t23ojayfVunABXQEGgZF_zPdbcUjJm7Rio7DM4JNWR3jrOkoMq_gCyB-ALK0Jo0Kes/s1600/Tropidolaemus+wagleri+Ian+Glover.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="191" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg9TmSYbe_c6AMVSpIpW0f69hVgjBkQu6SLXDPA2Q8OtHEq8ha7lzUDj4TVK8fZZRm7X_IereKNx2t23ojayfVunABXQEGgZF_zPdbcUjJm7Rio7DM4JNWR3jrOkoMq_gCyB-ALK0Jo0Kes/s320/Tropidolaemus+wagleri+Ian+Glover.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Temple or Wagler's Pitvipers (<i>Tropidolaemus wagleri</i>)<br />
at <a href="http://www.malaysiasite.nl/snaketempleng.htm">the famous Temple of the Azure Cloud in Penang, Malaysia</a><br />
<b>You can vote to sequence their genome<a href="http://www.pacb.com/smrt-science/smrt-grant/pag2017/"> here</a>!</b></td></tr>
</tbody></table>
<a href="http://snakesarelong.blogspot.com/2014/12/big-deal-snake-genomes.html">Genomics</a> of snakes is taking off in a big way, and we stand to learn <i>a lot more</i> about the evolution and function of snake venoms and the treatment of their effects. But, funding for basic science isn't a priority for many people, and more and more scientists are turning to crowd-funding their research or <a href="https://blogs.scientificamerican.com/the-curious-wavefunction/are-we-entering-a-golden-era-of-private-science-funding/">relying on limited funding from private foundations</a>, which often decide which projects to fund through a crowd-sourced voting process. This isn't necessarily a bad thing; in fact, I think it's a great thing in many cases. But, it's important to realize that government funding for science is different from private funding in two crucial ways: 1) there is a lot more of it (at least for now), and 2) it's not driven by specific, private interests. A great example is <a href="http://www.oriannesociety.org/">the Orianne Society</a>, a non-profit reptile conservation organization whose founding purpose was <a href="http://snakesarelong.blogspot.de/2012/09/recent-conservation-successes-with.html">preventing the extinction of Eastern Indigo Snakes (<i>Drymarchon couperi</i>)</a>. Thanks to generous donations from private funding sources, the Society succeeded in purchasing large areas of critical habitat for this endangered snake and protecting them in perpetuity, probably the most effective and laudable conservation goal in existence. Another good example is <a href="https://www.durrell.org/blog/the-translocation-of-the-round-island-keelscaled-boa2/">the work of the Durrell Wildlife Conservation Trust</a>, who have essentially saved a globally-rare snake, <i><a href="https://snakesarelong.blogspot.com/2012/04/round-island-splitjaw-snakes.html">Casarea dussumieri</a></i>, from extinction in the wild. I wish the quality conservation work that these organizations have become well-known for were more common, but to date their donors are some of the only large private backers of reptile research and conservation in the world.<br />
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Snakes are part of human economics, albeit to a lesser extent than many insects, fishes, birds, and mammals—they are hunted for food (although there are <a href="http://snakesarelong.blogspot.com/2016/05/rattlesnake-roundups-revisited.html">many issues</a> surrounding <a href="https://www.iucn.org/sites/dev/files/content/documents/2016/natusch_et_al_2016_sustainable_management_of_the_trade_in_reticulated_python_skins.pdf">better management</a> of <a href="https://s3.amazonaws.com/academia.edu.documents/42663317/Sea_Snake_Harvest_in_the_Gulf_of_Thailan20160213-463-nv40rx.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1490963882&Signature=vMRPIVB88D2I3nkWoMkqFoYJaYQ%3D&response-content-disposition=inline%3B%20filename%3DSea_snake_harvest_in_the_gulf_of_Thailan.pdf">unsustainable harvests</a>), kept as pets, their skins made into leather, and their venom harvested to make antivenom and other drugs. But, in their current form, these industries place very little emphasis on finding out more about snake biology in the wild; it just isn't necessary for them to make a profit, even though the information is important for what they do. Antivenom manufacturers are accountable to their shareholders, but trying to block FDA approval of Mexican antivenom is certainly not going to result in better treatment for snakebite victims in the USA, and American companies aren't investing in any research to create new, better products themselves, since drug development is expensive and risky, and <a href="https://www.wemjournal.org/article/S1080-6032(15)00410-X/fulltext">they already have a monopoly on antivenom in the USA</a>.<br />
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It's no secret that snakes and snake research have <a href="http://www.cebc.cnrs.fr/publipdf/2002/BTEE17.pdf">a PR problem: even scientific journals are less likely to publish research articles about snakes than about mammals and birds</a> (although the bias is likely subliminal). Many people prefer cute fuzzy animals that are similar to humans, but research into the biology of un-fuzzy animals is equally important. There's a parallel to the divide between funding for basic and applied science. <a href="https://www.forbes.com/sites/chadorzel/2017/03/23/the-most-important-science-to-fund-is-the-hardest-to-explain/">Basic science isn't usually as sexy as the exciting, fun applications that come later</a>, like saving lives, curing diseases, or discovering new complex biological phenomena. However, important applied science like antivenom creation cannot happen without basic science, in particular basic science on snakes. Private companies <a href="https://www.forbes.com/sites/chadorzel/2017/03/23/the-most-important-science-to-fund-is-the-hardest-to-explain/">can't afford to invest in basic science</a> the way they once did. Which leaves government funding and that from a limited number of interested, private backers.<br />
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We should support public funding for science and elect politicians who will do the same; better treatment for snakebite should be the least partisan and most universally-agreed-upon goal in the world. I think the path between basic (snake ecology, venomics, and genomics) and applied (antivenom manufacturing and public health) science is shorter and clearer in this context than in many, but the same principles apply—you cannot have medicine, conservation, and the other good parts of civilization without science.<br />
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You can <a href="http://www.pacb.com/smrt-science/smrt-grant/pag2017/">vote now through April 5th 2017</a> for a project sequencing the entire genome of the Temple Pitviper (<i>Tropidolaemus wagleri</i>) co-led by <a href="https://herpsocsg.wordpress.com/2017/03/22/genomics-and-venomics-of-the-sexually-dimorphic-temple-pitviper/">Ryan McCleary</a>.</div>
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<i>Stay tuned for more about the role of snake venom proteins in treating human diseases, and the role of snakes as predators in ecosystems.</i><br />
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<a href="https://www.blogger.com/null" name="1"><b>1 </b></a>Naturally, I wanted to link to the full-text of the paper so that anyone interested in learning more could read it, but the publisher (Wiley) has a 12-month embargo on posting the PDF anywhere online. They actually expect you to pay between $6 and $38 to read the article. Now, I think it's great research, and it probably cost Glaudas, Alexander, and their university thousands of dollars and thousands of hours to do it. But, if you pay Wiley to read their paper, none of that money will go to them, nor to the scientists who peer-reviewed their work for free. It will go to Wiley, who Xav paid (maybe) to publish. <a href="http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1469-7998/homepage/FundedAccess.html">They could have paid $3,000 to make it open access</a>, but you can understand why they didn't. No wonder most <a href="https://theconversation.com/academics-can-change-the-world-if-they-stop-talking-only-to-their-peers-55713">most science is read by fewer than 10 people</a>. It's an outdated model that can't go away fast enough. In contrast, <a href="http://www.mdpi.com/2072-6651/9/3/103">the spitting cobra paper is open access</a>, which <a href="http://www.mdpi.com/about/apc">cost its authors over $1,500</a>. This is typical; academic authors <a href="https://www.quora.com/How-much-money-do-academic-authors-typically-make-from-academic-publishing/answer/Zen-Faulkes">almost always lose money on a publication</a>.<a href="#top1"><sup>↩</sup></a><br />
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<a href="https://www.blogger.com/null" name="2"><b>2 </b></a>Recent update <a href="http://www.gazettenet.com/100-people-show-more-than-20-comment-on-state-rattlesnake-plan-in-Ware-8854207">here</a>; you can write the governor of Massachusetts <a href="http://www.mass.gov/governor/constituent-services/contact-governor-office/">here</a>.<a href="#top2"><sup>↩</sup></a><br />
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<a href="https://www.blogger.com/null" name="3"><b>3 </b></a><a href="http://www.ktxs.com/news/59th-annual-rattlesnake-roundup-in-sweetwater-going-on-this-weekend/390392665">Reports</a> suggest that <a href="http://www.dallasnews.com/life/texana/2017/03/16/texas-rattlesnake-festivals-create-buzz-controversy-hunting-techniques">this year</a>, <a href="http://snakesarelong.blogspot.de/2016/05/rattlesnake-roundups-revisited.html">like last year</a>, a much larger number of live rattlesnakes were collected than markets could support, and <a href="http://www.upi.com/Archives/1987/03/25/A-man-bitten-by-a-large-rattlesnake-he-was/7568305364460/">at least one person died from a snakebite</a> sustained while trying to capture a rattlesnake for a roundup.<a href="#top3"><sup>↩</sup></a><br />
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ACKNOWLEDGMENTS</div>
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Thanks to Bryan Fry for alerting me in advance of his publication, and to Colin Donahue, <a href="https://www.flickr.com/photos/markus_oulehla/15180251328/">Markus Oulehla</a>, and <a href="https://www.flickr.com/photos/irglover/18477096343/">Ian Glover</a> for the use of their photos.</div>
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<span style="font-size: x-small;">Putman, B. J., M. A. Barbour, and R. W. Clark. 2016. The foraging behavior of free-ranging Rattlesnakes (<i>Crotalus oreganus</i>) in California Ground Squirrel (<i>Otospermophilus beecheyi</i>) colonies. Herpetologica 72:55-63 <<a href="https://media.wix.com/ugd/8f1659_b5ae072145384e8e8a2d5faffcf9119f.pdf">full-text</a>></span><br />
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<span style="font-size: x-small;">Stock, R. P., A. Massougbodji, A. Alagon, and J.-P. Chippaux. 2007. Bringing antivenoms to Sub-Saharan Africa. Nature Biotechnology 25:173-177 <<a href="http://www.nature.com/nbt/journal/v25/n2/full/nbt0207-173.html">full-text</a>></span><br />
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<span style="font-size: x-small;">Wade, L. 2014. For Mexican antivenom maker, US market is a snake pit. Science 343:16-17 <<a href="http://opa.ahsc.arizona.edu/sites/opa.ahsc.arizona.edu/files/uploads/science-2014-wade-16-72.pdf">full-text</a>></span><br />
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<span style="font-size: x-small;">Willson, J. D. 2016. Indirect effects of invasive Burmese pythons on ecosystems in southern Florida. Journal of Applied Ecology 10.1111/1365-2664.12844 <<a href="http://comp.uark.edu/~jwillson/publications/2017%20-%20Willson-2017-JAE%20-%20python%20indirect.pdf">full-text</a>></span><br />
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="https://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
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Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com3Jena, Germany50.927054 11.58923720000007150.7669025 11.26651370000007 51.087205499999996 11.911960700000071tag:blogger.com,1999:blog-7443075087825368900.post-34051870716334083342017-02-28T07:34:00.002-07:002017-03-17T10:19:55.448-06:00Shield-tailed snakes (Uropeltidae)<div style="text-align: center;">
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<span style="font-size: x-small;"><a href="https://drive.google.com/file/d/0B2jxDXUJh_X7aDRaNjNWbGtISTQ/view?usp=sharing">Esta publicación es disponible en español</a></span></div>
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi71wjwwvGySINUwyUY0eGs1uu4IrTBKzJOLvj0vUCciTRS9YLBXVBo_UNTEBiefw0kP40I-4FURDhbWam599oQDMcn-qaR29GaNMvun9mZl1gHx_EcKImhyphenhyphenczxo8G-z_3wGCOq01MWaNRh/s1600/Uropeltis+macrolepis+Satyen+Mehta+tongue+flick.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="262" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi71wjwwvGySINUwyUY0eGs1uu4IrTBKzJOLvj0vUCciTRS9YLBXVBo_UNTEBiefw0kP40I-4FURDhbWam599oQDMcn-qaR29GaNMvun9mZl1gHx_EcKImhyphenhyphenczxo8G-z_3wGCOq01MWaNRh/s400/Uropeltis+macrolepis+Satyen+Mehta+tongue+flick.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Large-scaled Earth Snake <span style="font-size: 12.8px;">(</span><i style="font-size: 12.8px;">Uropeltis macrolepis</i><span style="font-size: 12.8px;">)</span></td></tr>
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Shield-tailed snakes (family Uropeltidae) are poorly studied, fossorial snakes endemic to montane regions of peninsular India & Sri Lanka. T<span style="text-align: start;">ogether with the families Cylindrophiidae (14 species) and <a href="http://snakesarelong.blogspot.com/2012/04/dwarf-pipesnakes.html">Anomochilidae (3 species)</a> they make up the superfamily Uropeltoidea, which is named for them because they are the most diverse subgroup, with </span><span style="text-align: start;">55 species in 8 genera (<a href="http://sciencepress.mnhn.fr/sites/default/files/articles/pdf/z2016n4a2.pdf">9 until recently</a>, more below). Most phylogenetic studies suggest that </span><span style="text-align: start;"><a href="http://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">Uropeltoidea</a> is the sister group to </span><span style="text-align: left;"><a href="http://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">Pythonoidea</a>, although these two lineages share only a few obvious features and likely diverged at least 60 and possibly up to 85 million years ago. If it's true, this relationship is pretty interesting because it means that the familiar giant pythons are more closely related to the ~18" long burrowing uropeltoids than they are to their most obvious ecological analogues, the giant Neotropical boas. However, this kind of relationship is not unprecedented: the emerging picture of <a href="http://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">henophidian</a> taxonomy is that constriction and large gape size have evolved at least three or four times within snakes, a good example being the sister relationship now widely accepted between <a href="http://snakesarelong.blogspot.com/2015/02/anilius-pipesnake-that-wasnt.html">the "pipesnake" family Aniliidae</a> and <a href="http://snakesarelong.blogspot.com/2015/01/dwarf-boas.html">the "dwarf boa" family Tropidophiidae</a> (the "Amerophidia"), both of which were formerly considered members of the Uropeltoidea. Even <a href="http://science.sciencemag.org/content/200/4337/74">some species of <i>Cylindrophis</i></a> (and, possibly, <i><a href="http://snakesarelong.blogspot.de/2015/01/dwarf-boas.html#3">Anilius</a></i>) immobilize their prey with their coils</span><span style="text-align: left;">, although they have small gapes.</span><br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi1OKA1u7RWesQcsQJ4Pr_h-e7aVruWGkkS9edLyb3CwJXeIfvigsPd9C8nf-_1J6YN5WOr6ksfN7p0Jwf0voi0syb5XcaFhX-IdXscAU6plJHOzsjmzWW8jzLUrJMlSBgFjN0RR385XVV3/s1600/Western+Ghats+map.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="400" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi1OKA1u7RWesQcsQJ4Pr_h-e7aVruWGkkS9edLyb3CwJXeIfvigsPd9C8nf-_1J6YN5WOr6ksfN7p0Jwf0voi0syb5XcaFhX-IdXscAU6plJHOzsjmzWW8jzLUrJMlSBgFjN0RR385XVV3/s400/Western+Ghats+map.jpg" width="308" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">The southern Western Ghats</td></tr>
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<span style="text-align: left;"><a href="https://www.researchgate.net/profile/Sr_Ganesh/publication/308748190_Shieldtail_snakes_Reptilia_Uropeltidae_-_the_Darwin's_finches_of_south_Indian_snake_fauna/links/57ee2c0008ae8da3ce482d69.pdf">Uropeltids are the Darwin's Finches of snakes</a>. They have radiated spectacularly across an archipelago of "sky islands", reaching their highest diversity and endemism in the mountain ranges </span>of India's southern tip. These volcanic mountains run parallel to the coast, creating a <a href="http://ngm.nationalgeographic.com/2009/11/india-rain/india-map">rain shadow</a> of dry plains to the east and generating torrential rainfall within their hills as trade winds blow monsoonally wet air northeast from the Arabian Sea. Known as the Great Escarpment of India, these mountains are an ancient coastline, formed during the break-up of the supercontinent Gondwana some 150 million years ago. Uropeltids are especially diverse in the high, wet 'shola' forests of the Anai Malai Hills (also known as the Elephant Hills), but are also highly diverse in the the Pothigai (Agasthyar Malai), Nilgiri, and Cardamom Hills, as well as the northern and <a href="http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.708.7062&rep=rep1&type=pdf">central Western Ghats</a>, the Eastern Ghats, and on <a href="http://reptile-database.reptarium.cz/advanced_search?taxon=uropeltidae&location=Sri+Lanka&submit=Search">Sri Lanka</a>. Many of these mountain ranges are part of the UNESCO World Network of Heritage Sites and are among the "hottest hot-spots" of biological diversity in the world. They're home to the <a href="http://wiienvis.nic.in/Database/eri_8226.aspx">world's largest wild population of Indian Elephants</a>, the <a href="http://s3.amazonaws.com/academia.edu.documents/5996288/statusof_tigers2008.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1487329072&Signature=xlVmG7BDa4oPeHf4suHzNzBY8eE%3D&response-content-disposition=inline%3B%20filename%3DStatus_of_Tigers_Co-predators_and_Prey_i.pdf">second largest wild tiger population</a>, and even more critically-endangered, endemic mammals such as <a href="https://en.wikipedia.org/wiki/Nilgiri_tahr">Nilgiri tahr</a> and <a href="https://en.wikipedia.org/wiki/Malabar_large-spotted_civet">Malabar large-spotted civets</a>. <a href="https://books.google.de/books?hl=en&lr=&id=BUPcAwAAQBAJ&oi=fnd&pg=PA35&dq=%22Teretrurus+sanguineus%22&ots=AHKzeruoz4&sig=96n02EPaqV3Sz-xsAam1KPh98k8#v=onepage&q&f=false">Sixty-five percent of reptile species in this area are found nowhere else</a>. In addition to uropeltids, the Ghats are <a href="http://s3.amazonaws.com/academia.edu.documents/38744432/FB_04_Science.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1487155987&Signature=BaPppnVW0BbqSbnfZ%2BxD%2B%2BM2tp0%3D&response-content-disposition=inline%3B%20filename%3DLocal_Endemism_Within_the_Western_Ghats-.pdf">home to diverse radiations of</a> endemic freshwater crabs and shrimps, minnows and carps, tree frogs, and caecilians. Other endemic reptiles include the cane turtle (<i>Vijayachelys silvatica</i>) and travancore tortoise (<i>Indotestudo travancorica</i>), two genera of skinks (<i>Ristella</i> and <i>Kaestlea</i>), spiny tree lizards (genus <i>Salea</i>), and <a href="http://snakesarelong.blogspot.com/2016/02/dragonsnakes-and-filesnakes-revisited.html">wood snakes</a> (genus <i>Xylophis</i>, <a href="http://www.biodiversitylibrary.org/item/90451#page/154/mode/1up">also named by Beddome</a>, which were thought to be <a href="http://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">xenodermids</a> but from which we have just recently obtained the first DNA sequences, placing <a href="https://sararuane.files.wordpress.com/2013/12/ruane_et_al-2017-molecular_ecology_resources.pdf">at least one species with pareids</a> and <a href="https://academic.oup.com/mbe/article-abstract/33/10/2483/2925599/Visual-Pigments-Ocular-Filters-and-the-Evolution">another with natricines</a>). Besides <i>Xylophis </i>and uropeltids, there are at least 29 other species of endemic snakes, ranging from <a href="http://reptile-database.reptarium.cz/species?genus=Gerrhopilus&species=beddomii&search_param=%28%28species%3D%27beddomii%27%29%29">blindsnakes</a> to <a href="http://www.biodiversitylibrary.org/item/190685#page/106/mode/1up">sand boas</a>, <a href="http://reptile-database.reptarium.cz/species?genus=Ahaetulla&species=dispar&search_param=%28%28genus%3D%27ahaetulla%27%29%29">vine snakes</a>, <a href="http://www.mapress.com/zootaxa/2012/f/zt03437p068.pdf">coral snakes</a>, and <a href="http://reptile-database.reptarium.cz/species?genus=Tropidolaemus&species=huttoni&search_param=%28%28species%3D%27huttoni%27%29%29">vipers</a>.<br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEglx3zHo1V2qed3JYVc7cuB0Qg6VQTU480BMLRgAc8OPzkWU2q68AnRQuPdsB1tDxu386acFuDUPU7Di0zbrq2t8S2YzEC3zHN5SX5bhb__ya75fgjsARcEZ-fXz7w6kFY8mwhT1wnhaH0u/s1600/Beddome.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEglx3zHo1V2qed3JYVc7cuB0Qg6VQTU480BMLRgAc8OPzkWU2q68AnRQuPdsB1tDxu386acFuDUPU7Di0zbrq2t8S2YzEC3zHN5SX5bhb__ya75fgjsARcEZ-fXz7w6kFY8mwhT1wnhaH0u/s320/Beddome.jpg" width="285" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Col. Richard H. Beddome</td></tr>
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Perhaps it isn't surprising that most species of uropeltids <a href="http://sciencepress.mnhn.fr/sites/default/files/articles/pdf/z2016n4a2.pdf">"were described in a burst of activity in the 19th century"</a>, because British colonials cleared large swaths of mountain forest for timber and tea, coffee, and teak plantations between 1860 and 1950. Forty percent (22) of the known species were discovered and described by <a href="https://www.academia.edu/2430910/A_tribute_to_Col._Richard_Henry_Beddome">Colonel Richard Henry Beddome</a>, a naturalist who <i>"…exploited the South Indian Hills, including the Palni Hills, to such purpose in the seventies and eighties of the last century, that he has hardly left a snake for any later enthusiast to discover"</i> wrote distinguished herpetologist <a href="http://www.biodiversitylibrary.org/item/95242#page/707/mode/1up">Frank Wall in 1922</a>.<a href="#1" name="top1"><sup>1</sup></a> Perhaps somewhat ironically, <a href="http://gwydir.demon.co.uk/jo/genealogy/beddome/richardhenry.htm">Beddome's 1911 obituary</a> states that his career as a naturalist and forester coincided with <i>"the first systematic steps to save the forests of Southern India from the denudation at the hands of the rural population to which they had long been exposed"</i>. However, Beddome used his tours in the montane forests to carefully document, describe, and make exhaustive and useful collections of plants, land snails, amphibians, and reptiles. In his lifetime Beddome <a href="http://gwydir.demon.co.uk/jo/genealogy/beddome/richardhenry.htm">described over a thousand species of animals and plants</a>, and many others have been named for him.<a href="#2" name="top2"><sup>2</sup></a><br />
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<div style="text-align: left;">
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhDX1KEupH1UTvkoypjbr2U4GDFJv79XIxtIOJZ2-Knk9GdZxonFMawsIERUlxKfcYhHFZT6nyjahkXfiyQqPGw_RLYTltzStHD1Ii8h7oPnj7eZZEiOQiK2bxQqbLPr9hNzq7murRilbnf/s1600/Uropeltid+heads+%2526+tails+Pyron+et+al+2016.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="271" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhDX1KEupH1UTvkoypjbr2U4GDFJv79XIxtIOJZ2-Knk9GdZxonFMawsIERUlxKfcYhHFZT6nyjahkXfiyQqPGw_RLYTltzStHD1Ii8h7oPnj7eZZEiOQiK2bxQqbLPr9hNzq7murRilbnf/s400/Uropeltid+heads+%2526+tails+Pyron+et+al+2016.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><div style="font-size: 12.8px;">
Heads and tails of uropeltids. </div>
<div style="font-size: 12.8px;">
The blade-like, "boomerang rostral" and polygonal eye shield</div>
<div style="font-size: 12.8px;">
of <i>Rhinophis punctatus </i>(top left [top] and top right [side]).</div>
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<span style="font-size: 12.8px;">Shield-like tail of </span><i style="font-size: 12.8px;">Uropeltis </i><i style="font-size: 12.8px;">rubromaculata</i><span style="font-size: 12.8px;"> (bottom left)<br />and</span><i style="font-size: 12.8px;"> </i><i style="font-size: 12.8px;">Rhinophis philippinus </i><span style="font-size: 12.8px;">(bottom right). </span><span style="font-size: 12.8px;">From</span><span style="font-size: 12.8px;"> </span><a href="http://sciencepress.mnhn.fr/sites/default/files/articles/pdf/z2016n4a2.pdf" style="font-size: 12.8px;">Pyron et al. 2016</a></div>
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</tbody></table>
Uropeltids are supremely adapted for burrowing, perhaps more so than any other snake. They construct a network of burrows during the rainy season, when the soil is soft, and wander through them after they harden. They can dig as deep as two meters and for extended periods of time. They have stout, relatively lizard-like skulls with few teeth, and conical, slender heads that are much narrower than their thick bodies. The eyes of some species <a href="http://snakesarelong.blogspot.com/2013/11/how-snakes-see-through-closed-eyes.html">are protected by polygonal scales</a>. <i>Rhinophis</i> and some <i>Uropeltis </i>have keeled, blade-like rostral scales that give the head a distinct pointed appearance.<span style="text-align: start;"> </span>Uropeltids have <a href="http://snakesarelong.blogspot.com/2012/11/identifying-snake-sheds-part-iii.html">narrow ventral scales similar to their dorsal scales</a>, and short, blunt, often shield-like tails, from which they get their common name. Uropeltid tail morphology ranges from relatively normal (in <i>Brachyophidium</i>, <i>Platyplectrurus</i>, and <i>Teretrurus</i>) or somewhat compressed with a multi-pointed scute on the end (in <i>Melanophidium</i>, <i>Plectrurus</i>, and <i>Pseudoplectrurus</i>), through decidedly unusual, including tails terminating in a projecting, rugose, keratinous disc (in <i>Rhinophis </i>and <i>Pseudotyphlops</i>), to the classic, highly modified “shield” tail of some <i>Uropeltis</i>, in which the body appears to have been sliced off at a ~45° angle, leaving a flattened disc covered with rugose scales. However, the real specializations for burrowing are hidden within.<br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhW_dF5iByjgDgj-5ploawRKTtpB69BRy_QJHUGRHoS0Nj9HuxTeSE-ilRi27_83XRx9B2qKl6lfvu5J69bLS9asHu2Fc4sOq18mQIW50S3aes0bpaQdT1Qu7DZQbCLZFZLwZVk4TuJCRzD/s1600/Rhinophis+drummondhayi+Gans+et+al+1977+Fig1.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="284" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhW_dF5iByjgDgj-5ploawRKTtpB69BRy_QJHUGRHoS0Nj9HuxTeSE-ilRi27_83XRx9B2qKl6lfvu5J69bLS9asHu2Fc4sOq18mQIW50S3aes0bpaQdT1Qu7DZQbCLZFZLwZVk4TuJCRzD/s320/Rhinophis+drummondhayi+Gans+et+al+1977+Fig1.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Diagram of a dissected <i>Rhinophis drummondhayi</i><br />
showing the extent <span style="font-size: 12.8px;">of red and white muscle along the body<br />and in </span><span style="font-size: 12.8px;">two cross-sections. From </span><a href="http://science.sciencemag.org/content/199/4325/189.long" style="font-size: 12.8px;">Gans et al. 1978</a></td></tr>
</tbody></table>
<div style="text-align: left;">
<div style="text-align: justify;">
Firstly, the heads of uropeltids are battering rams that are used against the soil. They are the only amniotes <a href="http://www.biodiversitylibrary.org/page/3191051#page/422/mode/1up">whose skulls are supported at the base by two vertebrae</a>: that is, both the first and second vertebra (the human atlas and axis) articulate directly with the occipital condyle at the base of the skull. Furthermore, <a href="http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0032450">their braincases are reinforced</a> and <a href="http://www.academia.edu/download/33060550/2002_-_Rieppel_and_Zaher_2002_The_skull_of_the_Uropeltinae_with_special_reference_to_the_otico-occipital_region.pdf">many other skull bones</a> are strong and stout, especially for a snake. The anatomy and physiology of the anterior third of a uropeltid's body is adapted for driving this strong head forward into the soil. <a href="http://science.sciencemag.org/content/199/4325/189.long">The muscles along the anterior portion of the trunk</a> are large, thick, deep red, and rich in myoglobin, <span style="text-align: left;">catalytic enzymes, and mitochondria, all <a href="https://link.springer.com/book/10.1007/978-1-4613-4609-8">biochemical or cellular adaptations that permit sustained activity</a>. </span><span style="text-align: left;">These muscles are loosely attached to the rest of the body, </span><span style="text-align: left;">so they can simultaneously push the sides of the body against the tunnel walls and move the head forward, without</span> pushing the rest of their bodies backwards. To accomplish this, muscles in the posterior body squeeze the anterior vertebral column into a sequence of hairpin turns, <a href="http://snakesarelong.blogspot.com/2012/07/snake-eating-snakes.html">not unlike those formed in the vertebrae of large, elongate prey when they are eaten by snakes</a>.<a href="#3" name="top3"><sup>3</sup></a> Because the tip of the nose creates a narrow burrow that is later widened by the flexing of the body, uropeltids can burrow effectively among rocks and roots.<a href="#4" name="top4"><sup>4</sup></a> Like a freight train, the anterior fifth of the body is like a locomotive in that it contains almost all of the propulsive machinery, and pulls along behind it the mostly-inert posterior trunk like the other train cars, containing viscera, embryos, food, etc., all protected on the end by a caboose-like caudal shield. You can get some idea of how it works <a href="https://youtu.be/wJ3HCnTZqL8">in this video</a> (compare the forceful muscle contractions <a href="https://youtu.be/wJ3HCnTZqL8?t=68">here</a> with how the rest of the body is simply dragged underground <a href="https://youtu.be/wJ3HCnTZqL8?t=152">here</a>).</div>
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhVldi9KhKlxP2hGS36RTZD8NJMkIHqm4-J55C2RF7f2g1CptuIjQtdPxuf3sxJKYpKIAiC5PvumzGbC39s4wV42nUYM5B68eC8TYntYJmAig7EJbFoNgYDIaPUQ4v7fbH0rvGsz_rXVgP1/s1600/uropeltid+burrowing+diagram+Gans+et+al+1978.jpg" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="171" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhVldi9KhKlxP2hGS36RTZD8NJMkIHqm4-J55C2RF7f2g1CptuIjQtdPxuf3sxJKYpKIAiC5PvumzGbC39s4wV42nUYM5B68eC8TYntYJmAig7EJbFoNgYDIaPUQ4v7fbH0rvGsz_rXVgP1/s400/uropeltid+burrowing+diagram+Gans+et+al+1978.jpg" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Schematic diagram of a uropeltid burrowing, from <a href="http://science.sciencemag.org/content/199/4325/189.long">Gans et al. 1978</a>.<br />
The dark black areas between the snake and the tunnel wall indicate<br />
firm contact. In A) the snake's vertebral column is curved and pushing<br />
against the sides of the tunnel. In B) the firm contact between the curved spine<br />
and the tunnel walls acts as a base against which the head can push,<br />
extending the tunnel forward. The widened body narrows as the spine uncurves.<br />
In C) the snake pulls its vertebral column forward and reintroduces<br />
the curves, which widen the body and the tunnel. The rest of the body<br />
is pulled along without doing any work or needing to resist any force.</td></tr>
</tbody></table>
<div style="text-align: justify;">
This division of labor is <a href="http://www.nature.com/nature/journal/v234/n5325/abs/234150a0.html">similar to that seen in some caecilians</a>, which burrow in a similar way and thereby create tunnels that are wider than their bodies. <span style="text-align: left;"> And, </span><span style="text-align: left;">unlike scolecophidians and amphisbaenians<a href="#5" name="top5"><sup>5</sup></a>, they can burrow without pushing against their tails, which leads to the question of what exactly their weird, shield-like tails are for, if not being pushed against? </span>It is thought that the function of these eponymous tails is to collect dirt as the snakes burrow, forming a "plug" that protects the snake from behind. The <a href="https://pdfs.semanticscholar.org/ac05/3d6b8f1bbee7e3f918e2cd3181b48860b559.pdf">scale texture</a> of the tail shield scales is deeply ridged, in sharp contrast to the texture of the body scales, which instead bear regular microstructure that inhibits wetting, sheds dirt, reduces friction, and produces iridescent colors. There is also evidence that the tail disc develops over the lifetime of some species, because juveniles do not have modified tails (although they do have large, deep red axial muscles like those of adults).</div>
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg611NWonSqclIuP7L2zQ_UcpVOWUPfteWQPN2WsjFUl1DUpNsAy9R5OwvvDumNgsON75yZCeTdej9yQqEDfaqB5vKg3FCuPoeqON6Y9yN3duiPphrkAGXT0DL4pFRJxDiFhmCeHVKeSfA3/s1600/uropeltids+DB%2526D+1854.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="480" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg611NWonSqclIuP7L2zQ_UcpVOWUPfteWQPN2WsjFUl1DUpNsAy9R5OwvvDumNgsON75yZCeTdej9yQqEDfaqB5vKg3FCuPoeqON6Y9yN3duiPphrkAGXT0DL4pFRJxDiFhmCeHVKeSfA3/s400/uropeltids+DB%2526D+1854.png" width="294" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Uropeltids from <a href="http://www.biodiversitylibrary.org/item/99515#page/153/mode/1up">Duméril, Bibron, & Duméril's<br />1854 Erpetologie Générale</a><br />
—<br />
Top left and top right: <i>Rhinophis philippinus</i><br />
Center left and center right: <i>Rhinophis saffragamus</i><br />
(<a href="http://sciencepress.mnhn.fr/sites/default/files/articles/pdf/z2016n4a2.pdf">formerly</a> <i>Pseudotyphlops philippinus</i>)<br />
Bottom left and bottom right: <i>Uropeltis ceylanica</i><br />
Center top and center bottom: <i>Plectrurus perroteti</i></td></tr>
</tbody></table>
<a href="http://sciencepress.mnhn.fr/sites/default/files/articles/pdf/z2016n4a2.pdf">A new phylogeny</a>, the most comprehensive yet, nevertheless includes DNA from just five of the eight genera of uropeltids. The most diverse and well-known genera are <i>Uropeltis</i> and <i>Rhinophis</i>, containing 24 and 19 species respectively. These are also the most highly specialized for burrowing. <i>Rhinophis </i>is so bizarre that it was originally described as a subgenus of the legless lizard genus <i>Anguis</i>. In contrast, the smaller and more poorly-known genera <i>Brachyophidium </i>(1 species), <i>Melanophidium </i>(4 species), <i>Platyplectrurus </i>(2 species), <i>Plectrurus </i>(4 species), and <i>Teretrurus </i>(1 species) have less highly modified heads, tails, and body musculature. Apparently these species are unable to tunnel in dry grassland soils, instead remaining belowground until rain softens the soil. Although the 'shola' forests have been greatly reduced, in recent years many of the remnants have been protected. In contrast, the high-altitude grasslands favored by certain species have, like grasslands all over the world, been largely ignored from a conservation standpoint. A single species in an eighth genus, <i>Pseudoplectrurus</i>, is known only from the original specimens collected by Beddome in 1870, from atop the 6000' Mount Kudremukh. It seems that uropeltids first evolved in India at least 37 million years ago, and crossed only once onto Sri Lanka, <a href="https://scholar.google.com/scholar_url?url=http://xa.yimg.com/kq/groups/22368033/1011644859/name/Phylogeny_SL_snakes_Pyron_etal_2013.pdf&hl=en&sa=T&oi=gsb-gga&ct=res&cd=0&ei=h4e1WL-RJsW8jAHdpZagBw&scisig=AAGBfm0qjFpWV0c2Pq_Cs764qiVIV6F4Rw">an island with one of the most phylogenetically diverse snake faunas in the world</a>, but which has maintained its distinctiveness from the Indian mainland despite several extended periods of land connection during the past 500,000 years.<br />
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<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgBwsZSb4tWSVVeMF60eskW49MdmjsvJHou-M5x7He6W0C1j3KCy5zTR3HWGwQ1CUz0l0dmVPXp78t6i6Bn6KnzEuwND93x-tfpe0WmJapuuCGKn-zE8xIg1lbwrIkg20sbWcN9YYL4HceV/s1600/Uropeltis+macrolepis+eating+earthworm+M.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="155" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgBwsZSb4tWSVVeMF60eskW49MdmjsvJHou-M5x7He6W0C1j3KCy5zTR3HWGwQ1CUz0l0dmVPXp78t6i6Bn6KnzEuwND93x-tfpe0WmJapuuCGKn-zE8xIg1lbwrIkg20sbWcN9YYL4HceV/s320/Uropeltis+macrolepis+eating+earthworm+M.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Uropeltis macrolepis</i> eating an earthworm</td></tr>
</tbody></table>
Unfortunately, we still know precious little about the ecology of uropeltids. Most species eat 80-90% earthworms, but they may snack upon the occasional earwigs, termites, or caterpillars. They are eaten by kraits (genus <i>Bungarus</i>) and <a href="https://www.youtube.com/watch?v=rU9N0aiKqWc">vinesnakes</a> (genus <i>Ahaetulla</i>), as well as wild boars, mongoose, owls, and galliform birds. They mate during the rainy season and females <a href="http://www.ias.ac.in/article/fulltext/reso/001/08/0064-0070">give birth to 3-9 live young at a time</a>. Like many fossorial snakes, some species are brightly colored on the underside, especially on the tail and neck. These colors may send warning signals to predators, including possibly mimicking the coloration of some venomous kraits or centipedes. It's likely that a high amount of diversity remains to be described. If you want to read about the current state of our knowledge of uropeltid diversity and taxonomy, including outlines of the genus-level groups that are supported by molecular and morphological phylogenies, not to mention numerous color photographs, you can do so <a href="http://sciencepress.mnhn.fr/sites/default/files/articles/pdf/z2016n4a2.pdf">here</a>.</div>
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<div style="text-align: justify;"><hr width="80%"><p><span class="Apple-style-span" style="font-size: x-small;"><br />
<a name="1"><b>1 </b></a>In the same issue, sandwiched between <i>"Alpine Orthoptera from central Asia"</i> and <i>"Hand-list of the Birds of India, Part IV"</i>, appears an article with the nonchalant title <a href="http://www.biodiversitylibrary.org/item/95242#page/154/mode/1up" style="font-style: italic;">"A few hints on crocodile shooting (with two Plates)"</a>, as well as a short note by a Miss Kennion called <a href="http://www.biodiversitylibrary.org/item/95242#page/397/mode/1up" style="font-style: italic;">"Crocodile shooting in Nepal"</a>. Sport hunting of predators was common during the British colonial period, and <a href="http://threatenedtaxa.org/index.php/JoTT/article/view/1790/3121">evidently human babies were sometimes used as bait</a>. It's a good thing Beddome and Wall were paying attention to uropeltids back then, because nobody else was.<a href="#top1"><sup>↩</sup></a><br />
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<hr width="80%"><p><span class="Apple-style-span" style="font-size: x-small;"><br />
<a name="2"><b>2 </b></a>Interestingly, <a href="http://gwydir.demon.co.uk/jo/genealogy/beddome/richardhenry.htm#description">a children's book written in 1947</a> by Vera Barclay contains a possible description of Col. Beddome. The book is called <i>"They Met a Wizard"</i> and the titular wizard is a zoologist living in colonial India with a special interest in snakes. Ms. Barclay was the great niece of Col. R.H. Beddome and it's likely that she knew him growing up and based her description of the zoologist in the story at least in part on her memories of him.<a href="#top2"><sup>↩</sup></a><br />
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<hr width="80%"><p><span class="Apple-style-span" style="font-size: x-small;"><br />
<a name="3"><b>3 </b></a>As a result, some early descriptions of uropeltids, such as <a href="http://www.biodiversitylibrary.org/item/24572#page/218/mode/1up">Günther's <i>The Reptiles of British India</i></a> or <a href="http://www.biodiversitylibrary.org/item/114007#page/66/mode/1up">Wall's <i>Ophidia Taprobanica</i></a>, contained erroneous claims that the neck was "swollen and knuckled" or that the head was very frequently bent to one side, as a result of the snake being preserved with the axial muscles contracted and unconstrained by tunnel walls.<a href="#top3"><sup>↩</sup></a><br />
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<a name="4"><b>4 </b></a>I cannot improve upon <a href="http://science.sciencemag.org/content/199/4325/189.long">the ingenious phrasing used by Carl Gans</a> to describe the burrowing of uropeltids: <i>"The burrowing method provides an ideal tunneling device for an unpredictably inhomogeneous substratum. The initial divot driven by the head is quite narrow and will be deflected by roots or rocks. When it passes close to such effectively nondeformable and nondisplacable objects, the opposite wall of the tunnel will be compressed unevenly so that the final tunnel achieves its full if meandering diameter by extra asymmetric compression of the softer zones."</i><a href="#top4"><sup>↩</sup></a><br />
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<hr width="80%"><p><span class="Apple-style-span" style="font-size: x-small;"><br />
<a name="5"><b>5 </b></a>Most amphisbaenians bite pieces out of their prey rather than swallowing it whole, so they are less likely to be impeded by a food bolus while burrowing.<a href="#top5"><sup>↩</sup></a><br /></div>
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ACKNOWLEDGMENTS</div>
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Thanks to <a href="https://sararuane.wordpress.com/">Sara Ruane</a>, <a href="https://www.flickr.com/photos/satyenmehta/12789159475/">Satyen Mehta</a>, and <a href="https://www.flickr.com/photos/milindsshutterbug/14630816360/">M</a> for the use of their photos, and to the <a href="http://www.retplants.org/index.php">Rare, Endangered and Threatened Plants of Southern Western Ghats database</a> for sharing their beautiful map.<br />
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REFERENCES</div>
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEimY9jDGBaDg2Rdx4hmPUmolqljhB8MFvllwJf4PvYa5CJFrMApnAh7Q7oZu5G6qftGLcqB3tC2_x1lLJdfaYpp0ZT-e_T6K9NguhC4tEC-usDX0JC2adEozmUfJyTp2a_OtwvY4A3Wxg_w/s1600/Uropeltis+macrorhynchus+Sara+Ruane.jpg" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEimY9jDGBaDg2Rdx4hmPUmolqljhB8MFvllwJf4PvYa5CJFrMApnAh7Q7oZu5G6qftGLcqB3tC2_x1lLJdfaYpp0ZT-e_T6K9NguhC4tEC-usDX0JC2adEozmUfJyTp2a_OtwvY4A3Wxg_w/s320/Uropeltis+macrorhynchus+Sara+Ruane.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Extremely similar head (top) and tail (bottom) of<br />
<i>Uropeltis macrorhynchus</i></td></tr>
</tbody></table>
<span style="font-size: x-small;">Beddome, R. H. 1886. An account of the earth-snakes of the peninsula of India and Ceylon. Annals and Magazine of Natural History 17:3-33 <<a href="http://www.biodiversitylibrary.org/page/15688563#page/17/mode/1up">Biodiversity Heritage Library</a>></span><br />
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<span style="font-size: x-small;">Bossuyt, F., M. Meegaskumbura, N. Beenaerts, D. J. Gower, R. Pethiyagoda, K. Roelants, A. Mannaert, M. Wilkinson, M. M. Bahir, K. Manamendra-Arachchi, K. L. N. Peter, C. J. Schneider, V. O. Oommen, and M. C. Milinkovitch. 2004. Local endemism within the Western Ghats-Sri Lanka biodiversity hotspot. Science 306:479-481 <<a href="http://s3.amazonaws.com/academia.edu.documents/38744432/FB_04_Science.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1487155987&Signature=BaPppnVW0BbqSbnfZ%2BxD%2B%2BM2tp0%3D&response-content-disposition=inline%3B%20filename%3DLocal_Endemism_Within_the_Western_Ghats-.pdf">download</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Comeaux, R. S., J. C. Olori, and C. J. Bell. 2010. Cranial osteology and preliminary phylogenetic assessment of <i>Plectrurus aureus</i> Beddome, 1880 (Squamata: Serpentes: Uropeltidae). Zoological Journal of the Linnaean Society of London 160:118-138 <<a href="https://www.researchgate.net/profile/Jennifer_Olori/publication/230019209_Cranial_osteology_and_preliminary_phylogenetic_assessment_of_Plectrurus_aureus_Beddome_1880_Squamata_Serpentes_Uropeltidae/links/57dff34b08aee5d2c6dd98b0.pdf">ResearchGate</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Gans, C. and D. Baic. 1977. Regional specialization of reptilian scale surfaces: relation of texture and biologic role. Science 195:1348-1350 <<a href="http://science.sciencemag.org/content/195/4284/1348">abstract</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Gans, C., H. C. Dessauer, and D. Baic. 1978. Axial differences in the musculature of uropeltid snakes: the freight-train approach to burrowing. Science 199:189-192 <<a href="http://science.sciencemag.org/content/199/4325/189.long">abstract</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Ganesh, S. 2010. Richard Henry Beddome and south India’s herpetofauna—a tribute on his centennial death anniversary. Cobra 4:1-11 <<a href="http://gwydir.demon.co.uk/jo/genealogy/beddome/rhbeddometribute.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Ganesh, S. 2015. Shieldtail snakes (Reptilia: Uropeltidae)–the Darwin’s finches of south Indian snake fauna? Pages 13-24 in P. Kannan, editor. Manual on identification and preparation of keys of snakes with special reference to their venomous nature in India. Proceedings by Govt. Arts College, Udhagamandalam, Tamilnadu, India <<a href="https://www.researchgate.net/profile/Sr_Ganesh/publication/308748190_Shieldtail_snakes_Reptilia_Uropeltidae_-_the_Darwin's_finches_of_south_Indian_snake_fauna/links/57ee2c0008ae8da3ce482d69.pdf">ResearchGate</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Ganesh, S. R. and S. R. Chandramouli. 2013. Endangered and Enigmatic Reptiles of Western Ghats – An Overview. Pages 35-61 in N. Singaravelan, editor. Rare Animals of India. Bommanampalayam Bharathiyar University (Post), Tamil Nadu, India <<a href="https://books.google.de/books?hl=en&lr=&id=BUPcAwAAQBAJ&oi=fnd&pg=PA35&dq=%22Teretrurus+sanguineus%22&ots=AHKzeruoz4&sig=96n02EPaqV3Sz-xsAam1KPh98k8#v=onepage&q&f=false">Google book</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Gaymer, R. 1971. New method of locomotion in limbless terrestrial vertebrates. Nature 234:150-151 <<a href="http://www.nature.com/nature/journal/v234/n5325/abs/234150a0.html">abstract</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Gower, D. J. 2003. Scale microornamentation of uropeltid snakes. Journal of Morphology 258:249-268 <<a href="https://pdfs.semanticscholar.org/ac05/3d6b8f1bbee7e3f918e2cd3181b48860b559.pdf">full-text</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Günther, A. 1864. The Reptiles of British India. Robert Hardwick, London <<a href="http://www.biodiversitylibrary.org/item/24572#page/9/mode/1up">Biodiversity Heritage Library</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Olori, J. C. and C. J. Bell. 2012. Comparative skull morphology of uropeltid snakes (Alethinophidia: Uropeltidae) with special reference to disarticulated elements and variation. PLoS ONE 7:e32450 <<a href="http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0032450">full-text</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Smith, M. A. 1943. The Fauna of British India. Volume III. Serpentes. Taylor & Francis, London <<a href="http://faunaofindia.nic.in/PDFVolumes/fbi/016/index.pdf">full-text</a>></span></div>
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<span style="font-size: x-small;"><br /></span></div>
<div style="text-align: justify;">
<span style="font-size: x-small;">Pyron, R. A., S. R. Ganesh, A. Sayyed, V. Sharma, V. Wallach, and R. Somaweera. 2016. A catalogue and systematic overview of the shield-tailed snakes (Serpentes: Uropeltidae). Zoosystema 38:453-506 <<a href="http://sciencepress.mnhn.fr/sites/default/files/articles/pdf/z2016n4a2.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Rajendran, M. 1985. Studies in uropeltid snakes. Madurai Kamaraj University, Madurai.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Rieppel, O. and H. Zaher. 2002. The skull of the Uropeltinae (Reptilia, Serpentes), with special reference to the otico-occipital region. Bulletin of the Natural History Museum: Zoology 68:123 <<a href="http://www.academia.edu/download/33060550/2002_-_Rieppel_and_Zaher_2002_The_skull_of_the_Uropeltinae_with_special_reference_to_the_otico-occipital_region.pdf">download</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Shanker, K. 1996. Nature watch: secrets of the shieldtails. Resonance 1:64-70 <<a href="http://www.ias.ac.in/article/fulltext/reso/001/08/0064-0070">full-text</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Wall, F. 1921. A new snake of the family Uropeltidae. Journal of the Bombay Natural History Society 28:41-42 <<a href="http://www.biodiversitylibrary.org/page/30112553#page/111/mode/1up">Biodiversity Heritage Library</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Wall, F. 1921. Ophidia Taprobanica, or the Snakes of Ceylon. H. R. Cottle, Govt. Printer, Colombo <<a href="http://www.biodiversitylibrary.org/item/114007#page/13/mode/1up">Biodiversity Heritage Library</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Wall, F. 1922. Acquisition of four more specimens of the snake <i>Brachyophidium rhodogaster </i>Wall. Journal of the Bombay Natural History Society 28:556-557 <<a href="http://www.biodiversitylibrary.org/item/95242#page/705/mode/1up">Biodiversity Heritage Library</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Williams, E. E. 1959. The occipito-vertebral joint in the burrowing snakes of the family Uropeltidae. Breviora 106:1-10 <<a href="http://www.biodiversitylibrary.org/page/3191051#page/419/mode/1up">Biodiversity Heritage Library</a>></span><br />
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="https://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
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Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com3Jena, Germany50.927054 11.58923720000007150.7669025 11.26651370000007 51.087205499999996 11.911960700000071tag:blogger.com,1999:blog-7443075087825368900.post-51674370337212977172017-01-31T08:24:00.002-07:002017-02-02T02:27:29.841-07:00Do snakes have a third eye?<div style="text-align: center;">
<span style="font-size: x-small;">This post will soon be available in Spanish</span></div>
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<tr><td class="tr-caption" style="text-align: center;">Limerick written by Annie Simminger about her nephew,<br />
<a href="https://en.wikipedia.org/wiki/Richard_M._Eakin">Richard Marshall Eakin</a>, and published in his 1983 book <span text-align: start;"><a href="https://books.google.de/books?id=pkDgJBd4O6oC&q=limerick#v=snippet&q=limerick&f=false" style="font-style: italic;">The Third Eye</a><i>.</i><br />Eakin and Robert C. Stebbins performed and published many<br />experiments on the structure and function of the third eyes of lizards</span></td></tr>
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Many lizards have a parietal eye, also known as a third eye or pineal eye. This "eye" is a photosensory organ located on the top of the skull, in the center. It has <a href="https://www.researchgate.net/profile/Gianluca_Tosini/publication/228648887_The_pineal_complex_of_reptiles_Physiological_and_behavioral_roles/links/56e020ef08aee77a15fe8a78.pdf">a well-defined lens, cornea, and retina</a>, and is lined on the inside with photosensitive cells that resemble the cones of the lateral eyes and contain <a href="https://www.scientificamerican.com/article/fact-or-fiction-carrots-improve-your-vision/">the light-sensitive pigment vitamin A</a><a href="#1" name="top1"><sup>1</sup></a>. These cells are connected by the parietal nerve to the pineal organ in the brain, which produces melatonin<a href="#2" name="top2"><sup>2</sup></a>, the hormone that controls sleep patterns, <a href="http://darkwing.uoregon.edu/~mosquito/articles/bradshaw_annrevecolsys2008.pdf">circadian rhythms</a>, and seasonal cycles such as mating, migration, and hibernation. The parietal eye can see light and is primarily used to sense changes in day length. Many lizards have a parietal eye, although it is most well-developed in tuataras, even serving as <a href="http://www.tuatarabrewing.co.nz/the-third-eye/about">the inspiration for a New Zealand brewery</a>.</div>
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Well, they're not nostrils. I don't think this has been scientifically evaluated before but maybe sensory related. Let's ask <a href="https://twitter.com/SssnakeySci">@SssnakeySci</a>. <a href="https://t.co/uwfvdLuYwI">https://t.co/uwfvdLuYwI</a></div>
— David Steen, Ph.D. (@AlongsideWild) <a href="https://twitter.com/AlongsideWild/status/821170254063792129">January 17, 2017</a></blockquote>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj_1bCzllZTGLtVtMUpQ7xRivpQ2yowYNbx3_yH4FVPErHwDElZYA9kkkRcjXNZIu4nY-6QwC6dPW-edB93WgW7PEL5qPQAleHUb0LQuGenjHxqy9cyH5oBv8IUZXKqniCplthwBMZTtuJy/s1600/Labra+et+al+2010+Fig2+Liolaemus+parietal+eye.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="243" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj_1bCzllZTGLtVtMUpQ7xRivpQ2yowYNbx3_yH4FVPErHwDElZYA9kkkRcjXNZIu4nY-6QwC6dPW-edB93WgW7PEL5qPQAleHUb0LQuGenjHxqy9cyH5oBv8IUZXKqniCplthwBMZTtuJy/s320/Labra+et+al+2010+Fig2+Liolaemus+parietal+eye.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Parietal eye (black outline) and parietal scale (white outline)<br />
of <i>Liolaemus bisignatus </i>(Philippi's Tree Iguana).<br />
<i></i>From <a href="http://repositorio.uchile.cl/bitstream/handle/2250/128896/Labra_Antonieta.pdf?sequence=1">Labra et al. 2010</a></td></tr>
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Recently, a Twitter conversation led me to evaluate the evidence for a parietal eye in snakes. As with many things, you would assume that if lizards have parietal eyes, then snakes have them too, since snakes are just one group of legless lizards. And, as with many things, you'd be wrong (probably; read on). It turns out that studies on the parietal eyes of snakes are almost non-existent. Maybe this isn't surprising, considering how little we know about other basics of snake physiology, like <a href="http://snakesarelong.blogspot.com/2015/09/can-snakes-hear.html">how well they can hear</a> or <a href="http://snakesarelong.blogspot.com/2015/08/do-snakes-sleep.html">whether or not they sleep</a>. The evidence for the existence of a parietal eye in snakes is scant at best, and despite evidence for its absence, <a href="http://carlgans.org/bor-view/?borv=9&borpage=367&borp=379">"amazingly few species have been studied"</a>, just seven as of 1979, and barely any others since then. <a href="http://link.springer.com/article/10.1007/BF00331245">Detailed studies</a> have been made on the pineal organ of just one species, <i>Natrix natrix</i>, the European Grass Snake, in contrast to <a href="http://carlgans.org/bor-view/?borv=9&borpage=-1">a large body of work on the pineal complex of lizards and tuataras</a>. The tale of the <a href="https://books.google.de/books?id=pkDgJBd4O6oC&lpg=PR9&dq=sphenodon%20pineal%20eye&lr&pg=PA13#v=onepage&q&f=false">evolution of the parietal eye</a> is helpful in understand the assumptions made about snake parietal eyes, in the absence of much direct research on them.</div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjC19ZDr5Rc4k258wMlSwfbeITdBaaPwZM3X_0ynj3aVWz0KNlMoTWskMhr55flfvOMNTRzYMS3tCcSbAP0nCHAVDvs7ye3Kusl5hj0xtvGSwg0oDbdb01J5GKfOjvzMA-yrSsvJ4SLW_if/s1600/Haughton+1929+Hipposaurus+composite.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="400" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjC19ZDr5Rc4k258wMlSwfbeITdBaaPwZM3X_0ynj3aVWz0KNlMoTWskMhr55flfvOMNTRzYMS3tCcSbAP0nCHAVDvs7ye3Kusl5hj0xtvGSwg0oDbdb01J5GKfOjvzMA-yrSsvJ4SLW_if/s400/Haughton+1929+Hipposaurus+composite.png" width="276" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">The "chimney-like" pineal foramen of<br />
the extinct 3-foot-long 250 million year old<br />
South African fossil therapsid <i>Hipposaurus.</i><br />
From <a href="http://www.biodiversitylibrary.org/item/126862#page/85/mode/1up">Haughton 1929</a></td></tr>
</tbody></table>
Tuataras and many lizards have relatively well-developed parietal eyes. These organs face upwards between the parietal bones of the skull, and were it not for their covering of skin they would effectively connect the outside world with the brain (more or less as our "normal" [lateral] eyes do). There are no eyelids, or rather, <a href="http://snakesarelong.blogspot.com/2013/11/how-snakes-see-through-closed-eyes.html">like the lateral eyes of snakes</a>, there is a fused, clear eyelid. In young lizards, the opening in the skull is large and T-shaped, like <a href="https://en.wikipedia.org/wiki/Fontanelle">the familiar soft fontanelle of an infant human's skull</a>, whereas in adult lizards the opening becomes ossified and may close completely late in life. Some extinct reptiles had bony protuberances around the margins of their parietal eyes: one, <i>Hipposaurus</i>, had a <a href="http://www.biodiversitylibrary.org/item/126862#page/85/mode/1up">"chimney-like structure"</a>. For a long time paleontologists debated whether holes in the parietal bones of fossil skulls were necessarily evidence of ancient parietal eyes. The German-American paleoneurologist and "fossil brain" expert <a href="http://academics.wellesley.edu/Biology/Faculty/Emily/BRB48(4).pdf">Tilly Edinger</a><a href="#3" name="top3"><sup>3</sup></a> snarkily wrote that <a href="http://www.biodiversitylibrary.org/item/26724#page/13/mode/1up">"if one doubts that this association existed also in extinct vertebrates, one may as well doubt that the orbits of fossil skulls contained eyes"</a>, because other than lizards and tuatara, no other living animals possess such holes. Many extinct tuatara relatives had even larger and more well-developed parietal eyes than do living tuataras—in some extinct pliosaurs, the opening was as large as 50 mm (2") by 20 mm (almost 1"), whereas in living tuataras it rarely exceeds 3 mm and modern lizards 1 mm. A more useful comparison may be the size of the parietal eye relative to that of the braincase: in living tuataras, this is about 1:7, whereas in living lizards it varies from 1:21 to 1:36 or less. The absence of any traces of musculature in fossil skulls or signs of more a complex past during embryonic development suggests that parietal eyes have never been any more elaborate in structure than they are in modern lizards and tuatara, although the larger parietal eyes of extinct reptiles were probably better at seeing than the tiny ones of living species.<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgLzIk7Q9gc-HABLv14e5Zrsf9VMl8HvwEXQTXQ_DypPqxttcLeeBfHa-GdhwGGEfmaC91ED4hGEBoiXHjOUfHj1j_9GyAoA6_8GM0y29NpLtz7mgaYRgOVJnjJ0t4pC6-TDbHdA5nuOJuO/s1600/Edinger+1955+cast+Lystrosaurus+brain.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="253" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgLzIk7Q9gc-HABLv14e5Zrsf9VMl8HvwEXQTXQ_DypPqxttcLeeBfHa-GdhwGGEfmaC91ED4hGEBoiXHjOUfHj1j_9GyAoA6_8GM0y29NpLtz7mgaYRgOVJnjJ0t4pC6-TDbHdA5nuOJuO/s400/Edinger+1955+cast+Lystrosaurus+brain.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Endocast of the brain of the dog-sized 250-million-year-old<br />
dicynodont <i><a href="http://phenomena.nationalgeographic.com/2013/05/28/lystrosaurus-the-most-humble-badass-of-the-triassic/">Lystrosaurus</a></i>, the "humble badass of the Triassic",<br />
showing the large parietal eye (dark structure at the top).<br />
From <a href="http://www.biodiversitylibrary.org/item/26724#page/13/mode/1up">Edinger 1955</a></td></tr>
</tbody></table>
Iguanids, agamids, varanids, cordylids, lacertids, and shinisaurids are diurnal, surface-active lizards that <a href="https://www.jstor.org/stable/1562791">have well-developed parietal eyes</a>. Several families of lizards that are mostly nocturnal and/or spend a great deal of time underneath cover or beneath the ground also have parietal eyes, including scincids, anguids, anniellids, xantusiids, amphisbaenids, and xenosaurids. <a href="http://onlinelibrary.wiley.com/doi/10.1002/ar.1091850404/abstract">Chameleons</a> have a degenerated parietal eye that lies above the foramen; presumably it is redundant with the lateral eyes of chameleons, which can move independently and cover 180° horizontally and 90° vertically. Some surface-active (teiids). burrowing (dibamids), and intermediate nocturnal (geckos) and diurnal (helodermatids and lanthanotids) lineages lack parietal eyes. Many of the lizard genera lacking a parietal eye have more equatorial geographic distributions. It has been suggested that a long evolutionary history in the tropics could lead to the loss of the parietal eye, because changes in day length are so minor close to the Equator. Even though there are seasons in the tropics (normally wet and dry), they are not associated with day length or light level cues that animals could use to know when the switch between the two is going to happen (and alter their lifestyles accordingly). There are no truly polar reptiles or amphibians, but <a href="https://books.google.de/books?id=ueb31jsWs2wC&lpg=PA1947&ots=_A1kTc2-TJ&dq=mammal%20lose%20pineal%20organ&lr&pg=PA1962#v=onepage&q&f=false">some polar mammals</a> (<i>e.g., </i>walruses, Weddell seals) have unusually large pineal organs, whereas <a href="http://link.springer.com/article/10.1007%2FBF01451040?LI=true">some tropical mammals</a> (<i>e.g., </i>sloths, pangolins) have lost their pineal organs, suggesting that the function of the pineal complex is more important where day length is more variable.<br />
<br />
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiqEEdzIDvAZzZ1eCExV69p94iyIuzPe0t1hBZWU5IrPy5hYC6d-YKMynEc1ynKOkFdMpnEfvGydjoqYg44s4Iwg6ZhZHFG34s36joNL-cD2Rc13lMDy_Kbg0occMZX0NyDWI_TvFBAUqFO/s1600/Bungarus+fasciatus+skull+Scanlon+%2526+Lee+2004.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiqEEdzIDvAZzZ1eCExV69p94iyIuzPe0t1hBZWU5IrPy5hYC6d-YKMynEc1ynKOkFdMpnEfvGydjoqYg44s4Iwg6ZhZHFG34s36joNL-cD2Rc13lMDy_Kbg0occMZX0NyDWI_TvFBAUqFO/s320/Bungarus+fasciatus+skull+Scanlon+%2526+Lee+2004.png" width="172" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Paired parietal foramina in the parietal (ptl)<br />
bone of a Banded Krait (<i>Bungarus fasciatus)</i><br />
skull. From <a href="http://www.academia.edu/download/31054601/2004Scanlon_Lee_Elapids.pdf">Scanlon & Lee 2004</a></td></tr>
</tbody></table>
It's thought that the parietal eye is retained in many burrowing lizards because these animals are occasionally exposed to light, and perhaps the parietal eye is a more suitable photoreceptor for a burrower than are lateral eyes, because it is already oriented upwards. If snakes evolved underground, <a href="https://bmcevolbiol.biomedcentral.com/articles/10.1186/s12862-015-0358-5">as the leading hypothesis suggests</a>, then it would make sense that they lost their parietal eye. Their normal eyes appear to have lost <a href="http://www.kingsnake.com/aho/pdf/menu3/rickshine/originofsnakes.pdf">some muscles</a> and <a href="http://s3.amazonaws.com/academia.edu.documents/41732364/Visual_system_evolution_and_the_nature_o20160129-8815-8hk1pl.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1485339031&Signature=3Wskl35eW%2BPkDWsDLVlNQPr9LPs%3D&response-content-disposition=inline%3B%20filename%3DVisual_system_evolution_and_the_nature_o.pdf">modern surface-dwelling snakes have lost at least two of the five visual pigment (opsin) genes</a> found in other vertebrates. Fossil and modern osteological evidence shows that a median parietal foramen like that of lizards was lost in an ancestor of all snakes (about 125 million years ago) and is not present in any living or fossil snakes. About 60 million years ago, small, laterally paired foramina evolved in early <a href="http://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">colubroids</a>, and are present in many, but not all, living elapids, viperids and other colubroid groups. As in lizards, these may be present only in juveniles, becoming obliterated externally by bone growth later in life. <a href="https://scholar.google.com/citations?hl=en&user=tkpCjdsAAAAJ">Snake osteology expert John Scanlon</a> told me that "Nobody, as far as I'm aware, has investigated whether the paired foramina [of snakes] are homologous or functionally similar to the median foramen of basal lepidosaurs [lizards]." The loss of parietal eyes is also supported by <a href="http://carlgans.org/bor-view/?borv=9&borpage=275&borp=292">developmental formation and then fusion with the pineal gland</a> in embryonic snakes, birds, and mammals<a href="#4" name="top4"><sup>4</sup></a>.<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhv5upAyBmhg6viQqFTgNd4cYiAgqZgUhHl2JV3N_F1mIYexmaCiPeTpU7fL5ckB5LTn5tLq7FwVZuO0NTAqHo4CDEs4vlQH0gPlYjnqcIYhL5diN2z_6B0oW8jhAuem51crmJUO2EjX0__/s1600/deveopment+of+parietal+eye+Eakin+1970.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="257" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhv5upAyBmhg6viQqFTgNd4cYiAgqZgUhHl2JV3N_F1mIYexmaCiPeTpU7fL5ckB5LTn5tLq7FwVZuO0NTAqHo4CDEs4vlQH0gPlYjnqcIYhL5diN2z_6B0oW8jhAuem51crmJUO2EjX0__/s400/deveopment+of+parietal+eye+Eakin+1970.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Developmental origins of the parietal ("median") eye and the lateral eye.<br />
The cilia are cellular structures that normally function for movement<br />
(<i>e.g.,</i> of debris out of the nose, of water over gills, of eggs into oviducts,<br />
of sperm cells to the egg). In the eye, <a href="https://books.google.de/books?id=pkDgJBd4O6oC&lpg=PR9&dq=sphenodon%20pineal%20eye&lr&pg=PA7#v=onepage&q=snake&f=false">they have evolved into photoreceptors</a>.</td></tr>
</tbody></table>
So, snakes join <a href="https://www.theguardian.com/science/2016/sep/07/did-milk-and-fur-evolve-before-the-earliest-mammals">most mammals</a>, birds, turtles, and most amphibians<a href="#5" name="top5"><sup>5</sup></a> in having <a href="https://books.google.com/books?id=qCDqCAAAQBAJ&pg=PA498&lpg=PA498&dq=%22Adaptive+radiation+of+the+pineal+system%22&source=bl&ots=2QTT_qUosc&sig=2xQmKsdX2rLJeSak-QOSw06oMgQ&hl=en&sa=X&ved=0ahUKEwjPjeewid3RAhXBUBQKHZqQBfkQ6AEIJDAC#v=onepage&q=%22Adaptive%20radiation%20of%20the%20pineal%20system%22&f=false">lost their parietal eyes</a> but retaining a photosensitive pineal organ in the brain that is not directly exposed to the outside of the skull. However, <a href="https://www.researchgate.net/profile/Gianluca_Tosini/publication/228648887_The_pineal_complex_of_reptiles_Physiological_and_behavioral_roles/links/56e020ef08aee77a15fe8a78.pdf">a recent review of the function of the pineal complex in reptiles</a> states that the pineal gland of adult snakes does not contain photoreceptor-like cells. Instead, the principal cells are pineal parenchymal cells, which secrete melatonin but do not sense light. Nevertheless, <a href="https://www.researchgate.net/profile/David_Crews/publication/14617750_Pinealectomy_melatonin_and_courtship_behavior_in_male_red-sided_garter_snakes_Tahmnophis_sirtalis_parietalis/links/547d18060cf285ad5b088a36/Pinealectomy-melatonin-and-courtship-behavior-in-male-red-sided-garter-snakes-Tahmnophis-sirtalis-parietalis.pdf">experiments on gartersnakes</a> have shown that removing the pineal organ of male gartersnakes in the fall, before hibernation, alters their melatonin cycle and reduces their courtship behavior when they emerge in the spring, so the pineal organ clearly functions to regulate melatonin and annual cycles in snakes.<br />
<br />
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjASAyaSZoC6SE-Wb9TwdayyCwdWcsBZiWiNCwl_iAxHjJHATdrgTB-FHEzhfk2SvLa0e73iC3f_IJCdtsXOj25aYJrNg-jd_gEJEKL3BKA9uSAe25PBrgisX13WV_ZPBbkrZ41_LKc4Mx2/s1600/lizard+parietal+eye+diagram+Solessio+%2526+Engbretson+1993.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="400" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjASAyaSZoC6SE-Wb9TwdayyCwdWcsBZiWiNCwl_iAxHjJHATdrgTB-FHEzhfk2SvLa0e73iC3f_IJCdtsXOj25aYJrNg-jd_gEJEKL3BKA9uSAe25PBrgisX13WV_ZPBbkrZ41_LKc4Mx2/s400/lizard+parietal+eye+diagram+Solessio+%2526+Engbretson+1993.png" width="221" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Diagram of the lizard parietal eye<br />
From <a href="http://www.nature.com/nature/journal/v364/n6436/abs/364442a0.html">Solessio & Engbretson 1993</a></td></tr>
</tbody></table>
At first glance, it doesn't seem to make sense to have a deep brain photoreceptor that isn't connected to the outside world, because it doesn't seem possible for it to be able to sense light or darkness from inside of your skull. But, don't forget that the two lateral eyes allow light to enter the brain; it is this light that the pineal organ is sensing. <a href="https://books.google.de/books?id=EZxEAAAAQBAJ&lpg=PA11&ots=wmIhMRY9Sv&dq=%22pineal%20parenchymal%20cells%22&pg=PA11#v=onepage&q=%22pineal%20parenchymal%20cells%22&f=false">Humans have pineal organs too</a>, and clearly we have no third eye (except in Greek mythology
and Grimm's fairytales). Think of how sleepy you feel when you have to get up before the sun, or how awake you feel when a bright light is turned on at night. This is because your pineal organ senses the ambient light or darkness and adjusts your melatonin levels, telling you (if it's bright) to wake up or (if it's dark) to stay asleep. Melatonin is also synthesized directly by the parietal eye of lizards. Although the pineal organ can only sense light and dark, there is <a href="http://www.tandfonline.com/doi/abs/10.1080/08927014.1992.9525353">evidence that the the parietal eye can also detect different colors of light</a>, including <a href="http://europepmc.org/abstract/med/7388628">ultraviolet</a> but not infrared light, and that <a href="http://www.nature.com/nature/journal/v364/n6436/abs/364442a0.html">it may be especially sensitive to the order of appearance of light of different wavelengths</a>, enabling lizards to detect dawn and dusk with great precision. <a href="https://www.jstor.org/stable/1562791">Detailed anatomical studies</a> have shown that the pineal organs of <a href="http://onlinelibrary.wiley.com/doi/10.1002/ar.1091850404/abstract">certain lizards</a> possess either a finger-like projection that extends toward the parietal eye, or convolutions of the pineal wall, both of which result in exposing and orienting more photoreceptor cells towards the skull roof, where they can detect light. Although these are sometimes occluded by cartilage or blood sinuses, their existence suggests that the pineal organ of lizards is a more important photoreceptor than previously realized.<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjkdy0am2whzPqSr9YKCiqFgUTslhBXxzJq7f2EGf2mh_IBsHXgnxdoPv2iR7OPLhnL81Q2Q8goTmOGwkE1eYrz-NXdYvw5xvh2St_FbXROU15vZkXqPQpqZ9aP7NtIhClITt1xuJZ4LgvU/s1600/Edinger+1955+Comparison+parietal+complex.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="292" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjkdy0am2whzPqSr9YKCiqFgUTslhBXxzJq7f2EGf2mh_IBsHXgnxdoPv2iR7OPLhnL81Q2Q8goTmOGwkE1eYrz-NXdYvw5xvh2St_FbXROU15vZkXqPQpqZ9aP7NtIhClITt1xuJZ4LgvU/s400/Edinger+1955+Comparison+parietal+complex.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Comparative morphology of the pineal complex in A) lamprey,<br />
B) frog, C) lizard, and D) human. From <a href="http://www.biodiversitylibrary.org/item/26724#page/13/mode/1up">Edinger 1955</a></td></tr>
</tbody></table>
</div>
<div style="text-align: justify;">
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhOYyXjCTOyaxAavGn_JHfSJ4odD1iCFmQsfJAKGNglUAFlfYtLXTen8HiDZkMfyzhBikt_YA50xdQS8e4yQl-KhLYwRfYSO9ohwq6I3rsOORUvSN-gmUdutTChZSbXk0p1aKYKVUjtiieL/s1600/Sceloporus+parietal+eye+Eakin+1970.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="222" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhOYyXjCTOyaxAavGn_JHfSJ4odD1iCFmQsfJAKGNglUAFlfYtLXTen8HiDZkMfyzhBikt_YA50xdQS8e4yQl-KhLYwRfYSO9ohwq6I3rsOORUvSN-gmUdutTChZSbXk0p1aKYKVUjtiieL/s400/Sceloporus+parietal+eye+Eakin+1970.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">The parietal eye of a Western Fence Lizard (<i>Sceloporus occidentalis</i>)<br />
C = cornea; CC = connective tissue; L = lumen;<br />
LS = lens; PN = parietal nerve; R = retina<br />
Light micrograph from <a href="http://www.jstor.org/stable/27828931">Eakin 1970</a></td></tr>
</tbody></table>
Many hypotheses have been put forth to explain the exact function of the parietal eye, which in some ways is still unclear. Rejected hypotheses include that the parietal eye is used for detection or deterrence of aerial predators. Even in tuataras, the parietal eye is barely noticeable (<a href="http://link.springer.com/article/10.1007/BF02955847">it wasn't described until the 1870s</a>), so predator deterrence is unlikely. It may play a minor role in predator detection, because the photoreceptive cells can respond to changes in light intensity as quickly as those of the lateral eyes, but sending sleepiness signals by initiating a melatonin cascade would be counterproductive to predator avoidance, to say the least. The most straightforward hypothesis is that it measures light intensity, functioning in regulating seasonal <a href="http://digitallibrary.amnh.org/bitstream/handle/2246/4659//v2/dspace/ingest/pdfSource/nov/N1870.pdf?sequence=1&isAllowed=y">seasonal behavior</a>, <a href="https://www.researchgate.net/profile/Gianluca_Tosini/publication/228648887_The_pineal_complex_of_reptiles_Physiological_and_behavioral_roles/links/56e020ef08aee77a15fe8a78.pdf">physiology</a>, and <a href="http://www.esf.edu/EFB/turner/publication%20pdfs/Pineal%20body%20%26%20thermoregulation.pdf">thermoregulation</a>. Although reptiles do have thermally sensitive neurons in their brains, we now know that the pineal complex does not directly sense heat. Instead, reptiles have <a href="http://journals.plos.org/plosone/article/file?id=10.1371/journal.pone.0000281&type=printable">specially-adapted transient receptor potential ion channels</a> (TRPs), which are proteins found throughout the body that act as internal thermometers and external temperatures sensors. Blocking the genes that make these proteins causes crocodiles to abandon their typical regime of behavioral thermoregulation and leads to significantly altered body temperature patterns. <a href="https://www.researchgate.net/profile/Victor_Hutchison/publication/248534232_Influence_of_thyroid_hormones_on_the_thermal_selection_of_African_house_snakes_Lamprophis_fuliginosus/links/0a85e53c42da0b53dd000000/Influence-of-thyroid-hormones-on-the-thermal-selection-of-African-house-snakes-Lamprophis-fuliginosus.pdf">Changes in melatonin levels also affect the body temperatures selected by some reptiles</a>, but in <a href="https://www.researchgate.net/profile/Gianluca_Tosini/publication/228648887_The_pineal_complex_of_reptiles_Physiological_and_behavioral_roles/links/56e020ef08aee77a15fe8a78.pdf">opposite ways in lacertids and iguanids</a>. There is also a great deal of evidence that the parietal eye is sensitive to polarized light: blocking the parietal eye <a href="https://www.researchgate.net/profile/John_Phillips2/publication/226107453_Orientation_in_a_desert_lizard_Uma_notata_time-compensated_compass_movement_and_polarotaxis/links/5581763a08aed40dd8cec5ae/Orientation-in-a-desert-lizard-Uma-notata-time-compensated-compass-movement-and-polarotaxis.pdf">disrupts sun-compass orientation</a> and <a href="https://www.researchgate.net/profile/Michael_Freake/publication/226662396_Homing_behaviour_in_the_sleepy_lizard_Tiliqua_rugosa_The_role_of_visual_cues_and_the_parietal_eye/links/554908550cf205bce7ac003f/Homing-behaviour-in-the-sleepy-lizard-Tiliqua-rugosa-The-role-of-visual-cues-and-the-parietal-eye.pdf">homing ability of displaced individuals</a> in several lizard species. This makes sense because there is no evidence that lizards can see polarized light with their lateral eyes. </div>
<div style="text-align: justify;">
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhYkae6aHTLxdgMyv22dpKV58f0sU16ksmg-TSGjJJKXrcuBPOalK0-CZxYrxllIUKKiO13s8jsi-0u7j_wuZlX2s5Ewdv_kcag1tVf2W7cay5hfz13AyJ-xICsc6K1hhETlN2vRJaYSeOU/s1600/Agkistrodon+contortrix+parietal+spots+JDW.JPG" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhYkae6aHTLxdgMyv22dpKV58f0sU16ksmg-TSGjJJKXrcuBPOalK0-CZxYrxllIUKKiO13s8jsi-0u7j_wuZlX2s5Ewdv_kcag1tVf2W7cay5hfz13AyJ-xICsc6K1hhETlN2vRJaYSeOU/s320/Agkistrodon+contortrix+parietal+spots+JDW.JPG" width="240" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Parietal spots of a Copperhead<br />
(<i>Agkistrodon contortrix</i>)</td></tr>
</tbody></table>
<a href="http://repositorio.uchile.cl/bitstream/handle/2250/128896/Labra_Antonieta.pdf?sequence=1">One study of thirty species of South American <i>Liolaemus </i>lizards</a> found that parietal eye size did not vary meaningfully with latitude, altitude, environmental temperature, thermal
tolerance, or body size, and that there was no evidence of phylogenetic inertia and high intraspecific
variation in parietal eye size, suggesting that parietal-eye size may not be under strong selection for accuracy. <a href="http://onlinelibrary.wiley.com/doi/10.1111/j.1751-1097.1976.tb07247.x/full">Another detailed study</a> found that removal of the parietal eye and pineal organ did not prevent 8 species of lizards from four families from carrying out their normal circadian rhythms. They concluded that other photoreceptors within the
brain were compensating, although the aforementioned extensions of the pineal organ may also be a factor in the occasional “failure” of parietalectomy experiments. It's actually <a href="http://darkwing.uoregon.edu/~mosquito/articles/bradshaw_annrevecolsys2008.pdf">not clear that we even have enough baseline data</a> on seasonal changes in snake circadian rhythms to correctly interpret the results of experiments that attempted to manipulate the pineal organs of snakes.<br /></div>
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<div style="text-align: justify;">
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="margin-left: auto; margin-right: auto; text-align: center;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhlMByABCGbt2bhjJlF8h49vfrDKJxLjh0alhpsW38LXhdrBQSOhGeJOnrzqLSI1XuMeiFDtoVM5EdFyVFNFSkQFZj309qJv3LbLqPZS34OlknuPU_AlcPFhchty75P4yyY8mxJaU7wzLoU/s1600/Agkistrodon+contortrix+skull+DigiMorph.png" imageanchor="1" style="margin-left: auto; margin-right: auto;"><img border="0" height="195" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhlMByABCGbt2bhjJlF8h49vfrDKJxLjh0alhpsW38LXhdrBQSOhGeJOnrzqLSI1XuMeiFDtoVM5EdFyVFNFSkQFZj309qJv3LbLqPZS34OlknuPU_AlcPFhchty75P4yyY8mxJaU7wzLoU/s320/Agkistrodon+contortrix+skull+DigiMorph.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Dorsal view of a Copperhead skull, from <a href="http://digimorph.org/specimens/Agkistrodon_contortrix/">DigiMorph</a></td></tr>
</tbody></table>
<br />
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhrlnqPRzYlUq7Jchd_linCUn1LfEZ6hn6WNMqnyIciksxI7XGsI_K7ephXUMZzpSuA65lj_ZjyIoG4OR0l8MyICDN4mU2qB6y4vntRrwQFbkqCexA8-cyM3O4BJ593tT7OkVXtZSRQ5bRB/s1600/IMG_1099.JPG" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhrlnqPRzYlUq7Jchd_linCUn1LfEZ6hn6WNMqnyIciksxI7XGsI_K7ephXUMZzpSuA65lj_ZjyIoG4OR0l8MyICDN4mU2qB6y4vntRrwQFbkqCexA8-cyM3O4BJ593tT7OkVXtZSRQ5bRB/s320/IMG_1099.JPG" width="294" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Pigmented apical pits of a ratsnake</td></tr>
</tbody></table></div>
<div style="text-align: left;">
But could the paired parietal formaina of some snakes function as parietal eyes? The question that started me looking into this was about Copperheads (<i>Agkistrodon contortrix</i>), which usually have a pair of small dark spots on their parietal scales. Evidently a <a href="https://www.youtube.com/watch?v=FJ_RA_T5bgg&app=desktop">National Geographic documentary</a> called them nostrils, which is totally absurd. But, the spots do seem to be in the approximate location of the parietal foramina in other snakes. <a href="http://digimorph.org/specimens/Agkistrodon_contortrix/">The DigiMorph scan of a copperhead skull</a> does not show any parietal foramina, although if it is of an adult specimen (not stated) then they may have closed up on top. A few other snake species also have such spots, and many snakes have pigmented sensory or apical scale pits elsewhere on their bodies. The parietal eyes of some lizards are also differentially pigmented. Do we need to open our (lateral) eyes to some new possibilities? I think it's clear that snake photoreception, although well-known in <a href="http://snakesarelong.blogspot.com/2012/09/snakes-that-can-see-without-eyes.html">species with pit organs</a>, is still relatively poorly understood for snakes as a whole.</div>
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<br /></div>
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<hr width="80%"><p><span class="Apple-style-span" style="font-size: x-small;"><br />
<a name="1"><b>1 </b></a>The function of vitamin A in eyesight was the basis for <a href="http://www.smithsonianmag.com/arts-culture/a-wwii-propaganda-campaign-popularized-the-myth-that-carrots-help-you-see-in-the-dark-28812484/">a WWII propaganda campaign that eating more carrots could improve human night vision</a>. Although it's true that carrots and vitamin A are essential for good eyesight, the extent to which eating more carrots can improve a person's eyesight was apparently greatly exaggerated in 1940 to create a cover story for the novel abilities of Allied pilots to pinpoint Axis fighter jets at night, which in reality was due to on-board Airborne Interception Radar (although there is <a href="https://www.scientificamerican.com/article/fact-or-fiction-carrots-improve-your-vision/">in turn some disagreement among historians as to how purposeful the deception was and how much both sides knew about the other side's radar capabilities</a>).<a href="#top1"><sup>↩</sup></a><br />
</span>
<hr width="80%"><p><span class="Apple-style-span" style="font-size: x-small;"><br />
<a name="2"><b>2 </b></a>Melatonin is synthesized from the amino acid tryptophan, which is the origin of another common myth: <a href="https://www.scientificamerican.com/article/fact-or-fiction-does-turkey-make-you-sleepy/">that eating a ton of turkey causes you feel sleepy</a>.<a href="#top2"><sup>↩</sup></a><br />
</span>
<hr width="80%"><p><span class="Apple-style-span" style="font-size: x-small;"><br />
<a name="3"><b>3 </b></a>Tilly Edinger was among the very last scientists of Jewish ancestry to leave pre-WWII Germany. A 1938 letter to the U. S. State Department in support of her immigration application from George Gaylord Simpson read "She is a research scientist of the first rank and is favorably known as such all over the world. She is everywhere recognized as the leading specialist on the study of the brain and nervous system of extinct animals and on the evolution of the gross structure of the brain. She is so preeminent in this field that she may really be said to have created a new branch of science, that of paleo-neurology, a study of outstanding value and importance”. She was the first female president of the Society of Vertebrate Paleontology, and authored over 1200 scientific papers and books, many sprinkled with sharp-witted, humorous phrases and observations. Her pioneering work in paleoneurology is well-chronicled <a href="http://academics.wellesley.edu/Biology/Faculty/Emily/BRB48(4).pdf">here</a>.<a href="#top3"><sup>↩</sup></a><br />
</span>
<hr width="80%"><p><span class="Apple-style-span" style="font-size: x-small;"><br />
<a name="4"><b>4 </b></a>During embryonic development, the parietal eye and the pineal organ form together from a pocket formed in the brain ectoderm. The ancestral state is presumed to have been a possibly paired photosensory organ, as seen in extant lampreys. The parietal eye and the pineal gland of tetrapods are probably the descendants of the left and right parts of this organ, respectively. Some Devonian fishes have two parietal foramina in their skulls, suggesting an ancestral bilaterality of parietal eyes.<a href="#top4"><sup>↩</sup></a><br />
</span>
<hr width="80%"><p><span class="Apple-style-span" style="font-size: x-small;"><br />
<a name="5"><b>5 </b></a>Crocodilians and some tropical lineages of mammals (<a href="http://link.springer.com/article/10.1007%2FBF01451040?LI=true">some xenarthrans</a> [sloths], <a href="https://books.google.de/books?id=ueb31jsWs2wC&lpg=PA1947&ots=_A1kTc2-TJ&dq=mammal%20lose%20pineal%20organ&lr&pg=PA1962#v=onepage&q&f=false">pangolins</a>, <a href="http://onlinelibrary.wiley.com/doi/10.1111/j.1463-6395.1985.tb00647.x/full">sirenians</a> [manatees & dugongs], some marsupials [<a href="https://books.google.de/books?id=eYOgce4cVH4C&lpg=PA25&ots=Gn41nuW8Wa&lr&pg=PA29#v=onepage&q&f=false">sugar gliders</a>]) have lost both their parietal eye and their pineal organ. <a href="http://onlinelibrary.wiley.com/doi/10.1111/j.1751-1097.1976.tb07250.x/abstract">All amphibians have a pineal organ</a>, but some frogs and toads also have what is called a "frontal organ", which is essentially a parietal eye. The word "pineal" comes from the shape of the human pineal organ, which resembles a pine cone.<a href="#top5"><sup>↩</sup></a><br />
</span></div>
<div style="text-align: justify;">
<br /></div>
<div style="text-align: center;">
ACKNOWLEDGMENTS</div>
<br />
<div style="text-align: justify;">
Thanks to <a href="https://twitter.com/PHSJonas">Daniel</a>, <a href="https://twitter.com/SssnakeySci">Helen Plylar</a>, and <a href="https://twitter.com/AlongsideWild">David Steen</a> for initiating a discussion of this topic on Twitter, to <a href="https://scholar.google.com/citations?hl=en&user=tkpCjdsAAAAJ">John Scanlon</a> for providing additional details about the evolution of parietal bone anatomy in squamates, and to Sandy Durso and <a href="https://www.flickr.com/people/57809070@N03/?rb=1">J. D. Willson</a> for the use of their photos.</div>
<br />
<div style="text-align: center;">
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<span style="font-size: x-small;">Linder, H. 1913. Beiträge zur Kenntnis der Plesiosaurier-Gattungen <i>Peloneustes </i>und <i>Pliosaurus</i>. Neues Jahrbuch für Geologie und Paläontologie–Abhandlungen 15:337-409.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Lutterschmidt, D. I., W. I. Lutterschmidt, and V. H. Hutchison. 1997. Melatonin and chlorpromazine: thermal selection and metabolic rate in the bullsnake, <i>Pituophis melanoleucus</i>. Comparative Biochemistry and Physiology Part C: Pharmacology, Toxicology and Endocrinology 118:271-277 <<a href="http://www.sciencedirect.com/science/article/pii/S0742841397001060">abstract</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Lutterschmidt, D. I., W. I. Lutterschmidt, and V. H. Hutchison. 2003. Melatonin and thermoregulation in ectothermic vertebrates: a review. Canadian Journal of Zoology 81:1-13 <<a href="https://www.researchgate.net/publication/237972712_Melatonin_and_thermoregulation_in_ectothermic_vertebrates_A_review">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Lutterschmidt, D. I., W. I. Lutterschmidt, N. B. Ford, and V. H. Hutchison. 2002. Behavioral thermoregulation and the role of melatonin in a nocturnal snake. Hormones and Behavior 41:41-50 </span><span style="font-size: x-small;"><</span><a href="https://www.researchgate.net/profile/Victor_Hutchison/publication/248534232_Influence_of_thyroid_hormones_on_the_thermal_selection_of_African_house_snakes_Lamprophis_fuliginosus/links/0a85e53c42da0b53dd000000/Influence-of-thyroid-hormones-on-the-thermal-selection-of-African-house-snakes-Lamprophis-fuliginosus.pdf" style="font-size: x-small;">link</a><span style="font-size: x-small;">></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Menaker, M. and G. Tosini. 1996. The evolution of vertebrate circadian systems. Pages 39-52 in K. Honma and S. Honma, editors. Sapporo Symposium on Biological Rhythms: Circadian Organization and Oscillatory Coupling. Hokkaido University Press, Sapporo, Japan.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Menaker, M. and S. Wisner. 1983. Temperature-compensated circadian clock in the pineal of <i>Anolis</i>. Proceedings of the National Academy of Sciences 80:6119-6121 <<a href="http://www.pnas.org/content/80/19/6119.full.pdf">link</a>></span>
<span style="font-size: x-small;"><br /></span><br />
<span style="font-size: x-small;">Mendonça, M. T., A. J. Tousignant, and D. Crews. 1995. Seasonal changes and annual variability in daily plasma melatonin in the red-sided garter snake (<i>Thamnophis sirtalis parietalis</i>). General and Comparative Endocrinology 100:226-237 <<a href="http://sites.utexas.edu/crewslab/files/2016/06/SeasonalChanges.pdf">link</a>></span>
<span style="font-size: x-small;"><br /></span><br />
<span style="font-size: x-small;">Mendonca, M. T., A. J. Tousignant, and D. Crews. 1996. Pinealectomy, melatonin, and courtship behavior in male red‐sided garter snakes (<i>Thamnophis sirtalis parietalis</i>). The Journal of Experimental Zoology 274:63-74 <<a href="https://www.researchgate.net/profile/David_Crews/publication/14617750_Pinealectomy_melatonin_and_courtship_behavior_in_male_red-sided_garter_snakes_Tahmnophis_sirtalis_parietalis/links/547d18060cf285ad5b088a36/Pinealectomy-melatonin-and-courtship-behavior-in-male-red-sided-garter-snakes-Tahmnophis-sirtalis-parietalis.pdf">link</a>></span>
<span style="font-size: x-small;"><br /></span><br />
<span style="font-size: x-small;">Mendonça, M. T., A. J. Tousignant, and D. Crews. 1996. Courting and noncourting male red-sided garter snakes, <i>Thamnophis sirtalis parietalis</i>: Plasma melatonin levels and the effects of pinealectomy. Hormones and Behavior 30:176-185 <<a href="https://www.researchgate.net/profile/David_Crews/publication/14406300_Courting_and_Noncourting_Male_Red-Sided_Garter_SnakesThamnophis_sirtalis_parietalisPlasma_Melatonin_Levels_and_the_Effects_of_Pinealectomy/links/00b7d5224e9df1db44000000.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Miller, W. H. and M. L. Wolbarsht. 1962. Neural activity in the parietal eye of a lizard. Science 135:316-317 <<a href="http://science.sciencemag.org/content/135/3500/316">abstract</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Moore, A. F. and M. Menaker. 2011. The effect of light on melatonin secretion in the cultured pineal glands of <i>Anolis </i>lizards. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology 160:301-308 <<a href="http://www.sciencedirect.com/science/article/pii/S1095643311002029">abstract</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Nelson, R. J., R. T. Mason, R. W. Krohmer, and D. Crews. 1987. Pinealectomy blocks vernal courtship behavior in red-sided garter snakes. Physiology & Behavior 39:231-233 <<a href="http://masonlab.science.oregonstate.edu/files/masonlab/007pinealectomy.pdf">link</a>></span><br />
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<span style="font-size: x-small;">Petit, A. 1968. Embryogénèse de l'épiphyse et de l'organe sous-commissural de la couleuvre à collier (<i>Tropidonotus natrix </i>L.). Arch. Anat. (Strasbourg) 52:3-25.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Petit, A. 1971. L'épiphyse d'un serpent: <i>Tropidonotus natrix </i>L. Zeitschrift für Zellforschung und mikroskopische Anatomie 120:94-119 <<a href="http://link.springer.com/article/10.1007/BF00331245">abstract</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Quay, W. 1979. The parietal eye–pineal complex. Pages 245-406 in C. Gans, R. G. Northcutt, and P. Ulinski, editors. Biology of the Reptilia. Volume 9. Neurology A. Academic Press, London <<a href="http://carlgans.org/bor-view/?borv=9&borpage=245">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Quay, W., J. A. Kappers, and J. Jongkind. 1968. Innervation and fluorescence histochemistry of monoamines in the pineal organ of a snake (<i>Natrix natrix</i>). Journal of Neuro-visceral Relations 31:11-25 <<a href="http://link.springer.com/article/10.1007/BF02239174">abstract</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Ralph, C. 1975. The pineal gland and geographical distribution of animals. International Journal of Biometeorology 19:289-303.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Ralph, C. L., B. T. Firth, and J. S. Turner. 1979. The role of the pineal body in ectotherm thermoregulation. American Zoologist 19:273-293 <<a href="http://www.esf.edu/EFB/turner/publication%20pdfs/Pineal%20body%20%26%20thermoregulation.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Ralph, C. L., S. Young, R. Gettinger, and T. O'Shea. 1985. Does the manatee have a pineal body? Acta Zoologica 66:55-60 <<a href="http://onlinelibrary.wiley.com/doi/10.1111/j.1463-6395.1985.tb00647.x/full">abstract</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Scanlon, J. D. and M. S. Lee. 2004. Phylogeny of Australasian venomous snakes (Colubroidea, Elapidae, Hydrophiinae) based on phenotypic and molecular evidence. Zoologica Scripta 33:335-366 <<a href="http://www.academia.edu/download/31054601/2004Scanlon_Lee_Elapids.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Seebacher, F. and S. A. Murray. 2007. Transient receptor potential ion channels control thermoregulatory behaviour in reptiles. PLoS ONE 2:e281 <<a href="http://journals.plos.org/plosone/article/file?id=10.1371/journal.pone.0000281&type=printable">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Simões, B. F., F. L. Sampaio, C. Jared, M. M. Antoniazzi, E. R. Loew, J. K. Bowmaker, A. Rodriguez, N. S. Hart, D. M. Hunt, J. C. Partridge, and D. J. Gower. 2015. Visual system evolution and the nature of the ancestral snake. Journal of Evolutionary Biology 28:1309-1320 <<a href="http://s3.amazonaws.com/academia.edu.documents/41732364/Visual_system_evolution_and_the_nature_o20160129-8815-8hk1pl.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1485339031&Signature=3Wskl35eW%2BPkDWsDLVlNQPr9LPs%3D&response-content-disposition=inline%3B%20filename%3DVisual_system_evolution_and_the_nature_o.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Solessio, E. and G. A. Engbretson. 1993. Antagonistic chromatic mechanisms in photoreceptors of the parietal eye of lizards. Nature 364:442-445 <<a href="http://www.nature.com/nature/journal/v364/n6436/abs/364442a0.html">abstract</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Stebbins, R. C. 1958. An experimental study of the" third eye" of the tuatara. Copeia 1958:183-190 <<a href="https://www.jstor.org/stable/1440585">abstract</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Stebbins, R. C. and R. M. Eakin. 1958. The role of the" third eye" in reptilian behavior. American Museum Novitates 1870:1-40 <<a href="http://digitallibrary.amnh.org/bitstream/handle/2246/4659//v2/dspace/ingest/pdfSource/nov/N1870.pdf?sequence=1&isAllowed=y">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Tilden, A. R. and V. H. Hutchison. 1993. Influence of photoperiod and temperature on serum melatonin in the diamondback water snake, <i>Nerodia rhombifera</i>. General and Comparative Endocrinology 92:347-354 <<a href="http://www.sciencedirect.com/science/article/pii/S001664808371172X">abstract</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Tosini, G. 1997. The pineal complex of reptiles: physiological and behavioral roles. Ethology Ecology & Evolution 9:313-333 <<a href="https://www.researchgate.net/profile/Gianluca_Tosini/publication/228648887_The_pineal_complex_of_reptiles_Physiological_and_behavioral_roles/links/56e020ef08aee77a15fe8a78.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Tosini, G., C. Bertolucci, and A. Foà. 2001. The circadian system of reptiles: a multioscillatory and multiphotoreceptive system. Physiology & Behavior 72:461-471 <<a href="http://www.sciencedirect.com/science/article/pii/S0031938400004236">abstract</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Trost, E. 1952. Untersuchungen über die frühe Entwicklung des Parietalauges und der Epiphyse von <i>Anguis fragilis</i>, <i>Chalcides ocellatus </i>und <i>Tropidonotus natrix</i>. Zoologischer Anzeiger 148:58-71.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Underwood, H. and M. Menaker. 1976. Extraretinal photoreception in lizards. Photochemistry and Photobiology 23:227-243 <<a href="http://onlinelibrary.wiley.com/doi/10.1111/j.1751-1097.1976.tb07247.x/full">abstract</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Underwood, H. 1989. The pineal and melatonin: regulators of circadian function in lower vertebrates. Cellular and Molecular Life Sciences 45:914-922 </span><span style="font-size: x-small;"><</span><a href="http://www.proberlab.caltech.edu/ewExternalFiles/Underwood-pinealectomy-1989-1.pdf5" style="font-size: x-small;">link</a><span style="font-size: x-small;">></span><br />
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<span style="font-size: x-small;">Ung, C. Y. J. and A. C. Molteno. 2004. An enigmatic eye: the histology of the tuatara pineal complex. Clinical & Experimental Ophthalmology 32:614-618 <<a href="http://onlinelibrary.wiley.com/doi/10.1111/j.1442-9071.2004.00912.x/full">abstract</a>></span><br />
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<span style="font-size: x-small;">Vollrath, L. 1979. Comparative morphology of the vertebrate pineal complex. Progress in Brain Research 52:25-38 <<a href="https://books.google.de/books?hl=en&lr=&id=eYOgce4cVH4C&oi=fnd&pg=PA25&ots=Gn41nuW8Wa&sig=2VO4INspcTnliCv0FJP2GKrWX7s#v=onepage&q&f=false">Google Book</a>></span><br />
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<span style="font-size: x-small;">Zimmerman, K. and H. Heatwole. 1990. Cutaneous photoreception: a new sensory mechanism for reptiles. Copeia 1990:860-862.</span><br />
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="https://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com1Jena, Germany50.927054 11.58923720000007150.7669025 11.26651370000007 51.087205499999996 11.911960700000071tag:blogger.com,1999:blog-7443075087825368900.post-61091220519853325972016-12-30T07:43:00.000-07:002017-01-06T05:00:19.288-07:00Life is Short but Snakes are Long 2016 Milestones<div style="text-align: justify;">
Dear reader,</div>
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiuzGUT26QbpnGlI0MD48KuPHzCc43NksIZMkJSiWQT4prZTSq024CbQH3C6SwCXje3jnT1UD3d7eG70Q6BVDSvUjEe-SnjdZ6edglYkOLiz4WDaLrvPkZ0fKgS3YFd-hAnUXajt25HVyU8/s1600/One+million+views+Blogger.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="179" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiuzGUT26QbpnGlI0MD48KuPHzCc43NksIZMkJSiWQT4prZTSq024CbQH3C6SwCXje3jnT1UD3d7eG70Q6BVDSvUjEe-SnjdZ6edglYkOLiz4WDaLrvPkZ0fKgS3YFd-hAnUXajt25HVyU8/s320/One+million+views+Blogger.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Screenshot from 15 November showing Blogger's estimate<br />
that <i>Life is Short but Snakes are Long</i> reached one million<br />
views, which is probably a bit too optimistic.</td></tr>
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<a href="http://snakesarelong.blogspot.com/2015/12/life-is-short-but-snakes-are-long-2015.html">As I did last year</a>, I want to thank you for your readership in 2016. <i>Life is Short but Snakes are Long</i> reached three-quarters of a million unique views on September 6th this year, by over 430,000 unique readers from nearly every country. We're currently over 860,000 views and on track to reach one million in 2017. The more liberal Blogger statistics show that we're already at one million, but I suspect that many of these are bots, and I'm sticking with the more conservative estimates provided by Google Analytics. I'm so happy to have reached so many people. Furthermore, <a href="http://reptile-database.reptarium.cz/advanced_search?taxon=Serpentes&year=2016&submit=Search">at least 34 new species of snakes were described in 2016</a>—another reason to celebrate!</div>
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In addition to defending my dissertation and moving to Germany in 2016, I also published <a href="https://www.researchgate.net/profile/Andrew_Durso">5 scientific papers</a> and co-authored a book chapter for the new 3rd edition of <i><a href="http://www.sciencedirect.com/science/book/9780721693279">Mader's Reptile Medicine and Surgery</a></i>, on the behavior of reptiles and amphibians, which will be published in 2017. I became a lot more active in the Facebook <a href="http://on.fb.me/1GLs0KK">Snake Identification</a> and <a href="https://www.facebook.com/groups/snakeED/">Wild Snakes: Education and Discussion</a> groups, which are fantastic resources for quick, reputable answers to questions about snakes. I recently accepted a position as an Associate Editor of the Snake Natural History Notes section at the journal <a href="https://ssarherps.org/publications/journals/herpetological-review/" style="font-style: italic;">Herpetological Review</a>, and I was invited to become a curator at the <a href="http://eol.org/">Encyclopedia of Life</a> project, where I've written <a href="http://eol.org/data_objects/34817626">several short summaries of snake taxa</a>.</div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgFOYzqJGKN8QV2FrPuGvxfQvQW38hQbTtq9aq2HbEpnjNMdeS4Btn95TJ8j93hu3tJ__M43Xc_L3Pb1QSKVQiisBJquF0WVG4iRSQBlS7talVv9kQppf5gmIxXQjqjLtuYVlxIlbPh15ah/s1600/Reptile_Amphibian_Badge.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="246" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgFOYzqJGKN8QV2FrPuGvxfQvQW38hQbTtq9aq2HbEpnjNMdeS4Btn95TJ8j93hu3tJ__M43Xc_L3Pb1QSKVQiisBJquF0WVG4iRSQBlS7talVv9kQppf5gmIxXQjqjLtuYVlxIlbPh15ah/s320/Reptile_Amphibian_Badge.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Life is Short but Snakes are Long was voted one of<br />
Bel-Rea Vet Tech College's Top 25 Reptile/Amphibian Blogs in 2016</td></tr>
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<i>Life is Short but Snakes are Long</i> was voted one of <a href="http://belrea.edu/blog/313-2/">Bel-Rea Vet Tech College's Top 25 Reptile/Amphibian Blogs</a>. The students and staff wrote that they particularly appreciated my efforts to reference my sources, and I was really glad to know that others appreciate my efforts to provide verifiable information (apparently there's all too little of that on the Internet these days).</div>
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I was particularly glad that the BBC's <i>Planet Earth II</i> featured <a href="https://www.youtube.com/watch?v=Rv9hn4IGofM">Galápagos Racers</a> so prominently this year, generating Internet-wide buzz about snakes and their feeding habits, a topic close to my heart. Since I <a href="http://snakesarelong.blogspot.com/2013/01/galapagos-racers.html">wrote about these interesting snakes back in 2013</a>, a lot of curious people found my blog, inspiring me to <a href="http://snakesarelong.blogspot.com/2015/12/life-is-short-but-snakes-are-long-2015.html">write an update and include much more detailed information</a>. I also revisited several other favorite topics, including <a href="http://snakesarelong.blogspot.com/2016/02/dragonsnakes-and-filesnakes-revisited.html">the relationship between dragonsnakes and filesnakes</a>, <a href="http://snakesarelong.blogspot.com/2016/05/rattlesnake-roundups-revisited.html">rattlesnake roundups</a>, <a href="http://snakesarelong.blogspot.com/2016/09/xenophidion-snake-with-mystery-penis.html">snake penises</a>, and <a href="http://snakesarelong.blogspot.com/2016/07/what-provincial-snakes-of-canada-should.html">snakes as state/provincial symbols</a>. I have some really good content planned to debut in 2017, including articles on the roles that snakes play in ecosystems, the nitty-gritty details of courtship, sex, and mating in snakes, the little-known and seldom-seen ecology of blindsnakes, profiles of some fossil snakes, and venomous bites from "non-venomous" snakes.</div>
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<i>Life is Short but Snakes are Long</i> would not be possible without support from volunteer translators <a href="http://malariaworld.org/blogs/alvaro-pemartin">Alvaro Pemartin</a> & Estefania Carrillo, from Utah State University, particularly my advisor <a href="http://frenchlab.weebly.com/">Susannah French</a> and <a href="http://ecology.usu.edu/">the Ecology Center</a>, and from my loving girlfriend and editor Kendal Morris.</div>
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Thank you, and happy 2017!</div>
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="https://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com3tag:blogger.com,1999:blog-7443075087825368900.post-43561818267227821472016-11-27T08:37:00.000-07:002018-02-12T08:49:36.196-07:00Galápagos Racers: answers to your questions about the BBC Planet Earth II iguana chase scene<div style="text-align: center;">
<span style="font-size: x-small;">This post will soon become available in Spanish</span></div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhR1yQ84JkAHxUC8MgXfldrNytR4yY8R5d-HP1mxbxAZTKWD56mqcOiZW5my86cyrlWoS3fFU9EM645FnzcxrhN1A7iks5p-KdwDUZseW8mV8nHgpVwwiYXswoIMjL3hmElCYZWIGIFr1Xw/s1600/Pseudalsophis+BBC+Planet+Earth+II+official.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="193" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhR1yQ84JkAHxUC8MgXfldrNytR4yY8R5d-HP1mxbxAZTKWD56mqcOiZW5my86cyrlWoS3fFU9EM645FnzcxrhN1A7iks5p-KdwDUZseW8mV8nHgpVwwiYXswoIMjL3hmElCYZWIGIFr1Xw/s320/Pseudalsophis+BBC+Planet+Earth+II+official.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><span style="font-size: x-small;">Galápagos Racers (<i>Pseudalsophis occidentalis</i>)<br />on Fernandina Island, from the BBC's Planet Earth II footage</span></td></tr>
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If you haven't seen <a href="https://www.youtube.com/watch?v=Rv9hn4IGofM">the incredible footage</a> of the "iguana chase scene" from <a href="http://www.bbc.co.uk/programmes/p04dgbn4/p04dgb2y">the BBC's Planet Earth II</a> <i>Islands </i>episode, I encourage you to watch it right away. In addition to being a highly dramatic cinematographic masterpiece, it raises a number of interesting questions about the biology of the snakes in the clip. For a few days after it aired, the Internet was buzzing with these questions, and I've cataloged the answers to some of the most popular ones below. If you have one that isn't listed, feel free to ask it in the comments! And, if you want to know more about the process I used to dig up some of this information, check out <a href="http://snakesarelong.blogspot.com/2014/10/how-to-teach-yourself-about-obscure.html">my tutorial for teaching oneself about obscure snakes</a>.<br />
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<b><span style="font-size: large;">What kind of snakes are they?</span></b><br /><br />Throughout the clip, Attenborough calls them "racer snakes"<a href="#1" name="top1"><sup>1</sup></a>, but herpetologists would normally call the snakes on the screen Galápagos Racers. Although these snakes are called "racers", they're not closely related to North American racers (genus <i>Coluber</i>); it's been about 45 million years since these two snakes last shared a common ancestor.<br /><br />Galápagos Racers belong to the genus <i>Pseudalsophis. </i>Depending on which sources you consult, there are between 4 and 7 species of <i>Pseudalsophis </i>in the Galápagos, as well as one in mainland South America.</div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiFaogFw5xvfcytivYUpb-e9CBsL5iTSEkOfLrRjhS3I17otKnxPOkYHTjYxE9nn1FVcL8-zN-j4m5Gv5_RVgSldnc7p2_XO1F4IDrZKwPTQu1vEZ7sKBTwDZ_iFa-lw7SdaHGAtFuxG-MH/s1600/Pseudalsophis+slevini+Andy+Kraemer.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiFaogFw5xvfcytivYUpb-e9CBsL5iTSEkOfLrRjhS3I17otKnxPOkYHTjYxE9nn1FVcL8-zN-j4m5Gv5_RVgSldnc7p2_XO1F4IDrZKwPTQu1vEZ7sKBTwDZ_iFa-lw7SdaHGAtFuxG-MH/s320/Pseudalsophis+slevini+Andy+Kraemer.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Pseudalsophis slevini</i> eating a gecko on Pinzón Island</td></tr>
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Just like Galápagos tortoises, finches, and many other organisms, there are different species of Galápagos Racers on the different Galápagos Islands (one of the concepts that sparked <a href="http://www.youtube.com/watch?v=GhHOjC4oxh8&feature=youtu.be">Darwin's theory of evolution by natural selection</a>). <span style="text-align: start;"><a href="http://www.bbc.co.uk/programmes/p04dgbn4/p04dgb2y">The film was made on Fernandina</a>, the <a href="https://www.geol.umd.edu/~jmerck/galsite/research/projects/leonard/Geospot.htm">youngest</a>, westernmost, and most volcanically-active island in the </span>Galápagos. Fernandina has two species of snakes, <i>Pseudalsophis slevini</i> and <i>Pseudalsophis occidentalis</i>. The snakes in the film must be <i>Pseudalsophis occidentalis</i>, because they are too large and not boldly banded enough to be <i>P. slevini</i>. You can read the original descriptions of both species <a href="http://www.biodiversitylibrary.org/item/98588#page/400/mode/2up">here</a>.<br />
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<a href="https://theconversation.com/in-defence-of-racer-snakes-the-demons-of-planet-earth-ii-theyre-only-after-a-meal-68514">None of the sources</a> reporting which species is shown in the film are authoritative, but without exception when the species is given it is given as <i>Pseudalsophis biserialis</i>. This is not correct under any modern taxonomy, although there is also a good explanation for why it is mistakenly being used—<i>P. occidentalis </i>was briefly a subspecies of <i>P. biserialis</i>, but has mostly been and is now treated either as a subspecies of <i>P. dorsalis</i> or as its own species. <a href="#2">See below</a> for much more (probably too much) detail.<br />
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<b><span style="font-size: large;">Why are there so many of them?</span></b></div>
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjSw69R-ia25E9Zq4ooSUGh15-_uH94sPTDOaNY_h2UOBqMG2xP5-FXnXuCTC9Idn1biSAKsAAztaLmPpiDPRnPhV6L0tU0qs56hZnWYqA041SLy6aFUD5kmkoMB-rZaoX596iDV_WECYfC/s1600/Pseudalsophis+and+marine+iguana_Jim+Moulton.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjSw69R-ia25E9Zq4ooSUGh15-_uH94sPTDOaNY_h2UOBqMG2xP5-FXnXuCTC9Idn1biSAKsAAztaLmPpiDPRnPhV6L0tU0qs56hZnWYqA041SLy6aFUD5kmkoMB-rZaoX596iDV_WECYfC/s320/Pseudalsophis+and+marine+iguana_Jim+Moulton.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Galápagos Racer (<i>P. dorsalis</i>) among adult Marine Iguanas<br />on Santa Cruz, which are much too large for it to eat</td></tr>
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<a href="http://snakesarelong.blogspot.com/2015/11/snakes-that-are-good-parents.html">Most snakes are not social</a>, and because <a href="http://snakesarelong.blogspot.com/2012/06/snakes-that-chew-their-food.html">they must swallow their food whole</a> they cannot share prey. These snakes are not found at such high densities year-round, but rather aggregate around consistent Marine Iguana nesting sites in May when the eggs are hatching.</div>
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Just as when baby sea turtles emerge from their nests, predators congregate at the temporary buffet, returning afterwards to their usual densities. Around the world, there are numerous examples of avian and snake predators exploiting emerging hatchling iguanas. Researchers working at other iguana nesting sites in <a href="https://www.researchgate.net/profile/Charles_Knapp2/publication/250067012_The_Influence_of_Landscape_Heterogeneity_and_Dispersal_on_Survival_of_Neonate_Insular_Iguanas/links/555b7e7208ae8f66f3ad7946.pdf">the Bahamas</a>, the West Indies, and <a href="https://www.researchgate.net/publication/274390801_Iguana_iguana_Green_iguana_Juvenile_predation">Venezuela</a> have hypothesized that snakes and other predators also converge on the nesting sites of these other iguanas to exploit the temporary food source. Another example of snakes congregating around abundant prey resources is that of <a href="http://www.jstor.org/stable/1565698?seq=1#page_scan_tab_contents">Puerto Rican and Cuban boas, which aggregate around the openings of massive bat caves</a>.</div>
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The rest of the year, Galápagos Racers eat lava lizards, geckos, insects, <a href="http://www.darwinfoundation.org/media/filer_public/2013/10/10/merlen__thomas_2013.pdf">marine fishes</a>, and hatchling birds, as well as introduced rats and mice.</div>
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<b><span style="font-size: large;">Are they really hunting in a pack?</span></b></div>
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Almost certainly not. Again, <a href="http://snakesarelong.blogspot.com/2015/11/snakes-that-are-good-parents.html">most snakes are not social</a>, and because <a href="http://snakesarelong.blogspot.com/2012/06/snakes-that-chew-their-food.html">they must swallow their food whole</a> they cannot share prey. Pack-hunting behavior is unknown in snakes.</div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjrC7jPSuJlhD_X5G2OdpUXfRX86gJSF-jMmqDMN5KUDuTcLrFMcEyLLxw0m2wnMZVBhSRPCq3wAXcax9Uif_dNS0XvMrsak48VkbmkSQiS3ZlH31B-Ia5TehpS4Avkp4w627wp4uuA5k4b/s1600/Pseudalsophis+occidentalis+trying+to+eat+the+same+iguana.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="171" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjrC7jPSuJlhD_X5G2OdpUXfRX86gJSF-jMmqDMN5KUDuTcLrFMcEyLLxw0m2wnMZVBhSRPCq3wAXcax9Uif_dNS0XvMrsak48VkbmkSQiS3ZlH31B-Ia5TehpS4Avkp4w627wp4uuA5k4b/s320/Pseudalsophis+occidentalis+trying+to+eat+the+same+iguana.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Two <i>P. occidentalis</i> trying to eat the same iguana<br />
Jaw-walking is a fixed action pattern in snakes and they<br />
may eat things that only vaguely resemble their food<br />
once they start jaw-walking them.<br />
From <a href="https://www.youtube.com/watch?v=2cOMjZWJyog">Planet Earth II Behind the Scenes</a></td></tr>
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<a href="http://snakesarelong.blogspot.com/2015/11/snakes-that-are-good-parents.html">Some species have surprisingly social behaviors</a>. It would be really interesting to examine social behavior in these snakes. To my knowledge no one has done so. Although they obviously cannot share a single food item, but if they are foraging in the same time and place on a limited resource, there might be an opportunity for the evolution of social cues. At least <a href="http://www.kingsnake.com/aho/pdf/November2003Update/shine2002c.pdf">one paper suggested that this might be the case with a pit viper</a>. Even though the BBC videographers saw snakes actively fighting over the same prey items and <a href="http://www.gq-magazine.co.uk/article/planet-earth-2">in some cases eating one another</a>, it's possible that more closely-related snakes are less likely to fight over food or eat one another, or that males are less likely to compete with or try to eat females. These are testable hypotheses. However, these are not well-studied snakes. I don't think they are helping each other, but there's a lot that we don't know about snakes. <a href="http://www.biodiversitylibrary.org/item/25758#page/209/mode/1up">Some snakes exhibit dominance hierarchies</a>, and <a href="http://psycnet.apa.org/journals/com/105/4/380/">one study suggested that individual recognition occurs and persists over time in gartersnakes</a>.</div>
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<a href="http://galapagosconservation.org.uk/wildlife/galapagos-racer/">Few scientists are currently studying these snakes</a>. It's a testament to the BBC that they are consistently able to film natural phenomena that are still unknown to science. Hopefully this tape will stimulate some research on this exact question, and on the ecology of Galápagos Racers. <a href="http://snakesarelong.blogspot.com/2013/01/galapagos-racers.html">When I wrote about </a><span style="text-align: justify;"><a href="http://snakesarelong.blogspot.com/2013/01/galapagos-racers.html">Galápagos Racers in 2013</a>, not much was known about their ecology, and that's still the case. I</span>t's amazing that so little research has been done on these snakes, particularly in contrast to Galápagos tortoises and marine iguanas (not to mention finches and other non-avian reptiles).</div>
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<b><span style="font-size: large;">Why don't the female Marine Iguanas just lay their eggs somewhere else, closer to the ocean maybe?</span></b></div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjx1yuyhDnPetUd44WygT72Xxu86j0XwCtw6GUp7MNCTBIkecvrRF3HCLqp9v-SxBQnRnc3qYSeUs5Pcj_LUeRpCfXP9Dh5WvQgg57qqhSVQUIv-Fx_jAYjcAZTEZKVwEuRUf3zCGvPehc3/s1600/Knapp+et+al+2010+Fig+3.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="195" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjx1yuyhDnPetUd44WygT72Xxu86j0XwCtw6GUp7MNCTBIkecvrRF3HCLqp9v-SxBQnRnc3qYSeUs5Pcj_LUeRpCfXP9Dh5WvQgg57qqhSVQUIv-Fx_jAYjcAZTEZKVwEuRUf3zCGvPehc3/s320/Knapp+et+al+2010+Fig+3.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Fates of rock iguana hatchlings, over half of which were<br />
eaten by <i>Cubophis</i> and <i>Epicrates</i> snake predators in their<br />
first month of life. From <a href="https://www.researchgate.net/profile/Charles_Knapp2/publication/250067012_The_Influence_of_Landscape_Heterogeneity_and_Dispersal_on_Survival_of_Neonate_Insular_Iguanas/links/555b7e7208ae8f66f3ad7946.pdf">Knapp et al. 2010</a></td></tr>
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Marine Iguanas have to dig nests and lay their eggs in soft sand, away from the rocky, tidal foraging grounds of the adults. They choose <a href="http://www.biodiversitylibrary.org/item/53858#page/342/mode/2up">protected lava reefs</a> for this purpose, which are in short supply on most islands. <a href="http://www.jstor.org/stable/2460852?seq=1#page_scan_tab_contents">One estimate</a> suggested that the cost of migrating to their nesting sites represented half the reproductive effort of female Galápagos land iguanas.<br />
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Many species of reptiles nest in areas where they otherwise do not spend much time, especially aquatic species (<a href="https://books.google.com/books?hl=en&lr=&id=BJc8J_l2L_UC&oi=fnd&pg=PA277&ots=3SY4OBjrCD&sig=gMtfVSJQNckmX8Xf0NVYnx5xok0#v=onepage&q&f=false">reptile eggs need to "breathe" air</a> and cannot be laid underwater). Female Marine Iguanas <a href="http://onlinelibrary.wiley.com/doi/10.1111/j.1365-2656.2007.01279.x/full">may all use the same nesting sites</a> because those are the only sites available, or they may choose to nest near one another because, just like with sea turtles, synchronous hatching of the young increases their probability of survival.<br />
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<a href="https://www.researchgate.net/profile/Charles_Knapp2/publication/250067012_The_Influence_of_Landscape_Heterogeneity_and_Dispersal_on_Survival_of_Neonate_Insular_Iguanas/links/555b7e7208ae8f66f3ad7946.pdf">In a study of Bahamian rock iguanas</a> (<i>Cyclura cychlura</i>), snake predation was the most likely cause of mortality for newborn iguanas dispersing away from their nests. They estimated that about 20-30% of hatchling iguanas survived their first month, and those that moved quickly and linearly away from their nests were the most likely to survive, perhaps because predators had learned to hang around the nesting area. <a href="http://link.springer.com/article/10.1007/BF00349191">Another study of Galápagos land iguanas</a> showed that predation attempts by Galápagos hawks were more than three times as likely to be successful when the body temperature of the iguana hatchlings was below 90°F. And, <a href="https://www.researchgate.net/publication/271696486_Population_Biology_of_Marine_Iguanas_Amblyrhynchus_cristatus_II_Changes_in_Annual_Survival_Rates_and_the_Effects_of_Size_Sex_Age_and_Fecundity_in_a_Population_Crash">baby Galápagos marine iguanas that hung around their hatching area</a> had about a 10% lower survival rate than those that moved to the coast, which the researchers attribute mostly to higher risk of predation at the nesting area.<br />
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Studies on <a href="https://www.researchgate.net/publication/271696486_Population_Biology_of_Marine_Iguanas_Amblyrhynchus_cristatus_II_Changes_in_Annual_Survival_Rates_and_the_Effects_of_Size_Sex_Age_and_Fecundity_in_a_Population_Crash">the population biology of Marine Iguanas</a> have shown that most of their mortality is caused by "predation, starvation (sometimes as a result of being trapped by a rock), crushing by a rock, being beaten against rocks by the sea, and suffocation in collapsed nest burrows. Animals may also die after being swept out to sea by offshore currents". So, actually, predation may be the best way for them to go. Besides Galápagos Racers, their other predators include Galápagos Hawks, Short-eared Owls, crabs, and Giant Hawk-fish.</div>
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<b><span style="font-size: large;">Are they venomous/dangerous to humans?</span></b></div>
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No. Like many snakes, Galápagos Racers are <a href="http://snakesarelong.blogspot.com/2013/09/basics-of-snake-fangs.html">rear-fanged</a>. This means that, although technically they are venomous, they don't pose a danger to humans. <a href="http://snakesarelong.blogspot.com/2013/09/basics-of-snake-fangs.html">Rear-fanged snakes</a> mostly have grooved teeth (rather than hollow fangs) on the back of their upper jaw (as opposed to the front); they can use these teeth to get venom into their prey once they are biting it, but they cannot strike out and deliver venom the way a viper can. A small minority of <a href="https://books.google.com/books?id=lB3FxFG5wrQC&dq=weinstein+venomous+bites&source=gbs_navlinks_s">rear-fanged snakes have delivered medically-significant bites to humans</a>, but almost all of these take place in a captive setting. You can read more about the different types of snake fangs <a href="http://snakesarelong.blogspot.com/2013/09/basics-of-snake-fangs.html">here</a>.</div>
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<b><span style="font-size: large;">I didn't know there were snakes in the Galápagos. How did they get there?</span></b></div>
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: justify;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiC0PpZspEAFknHObgCzcMQ5y4HqaVm_qNnVCtgVVRK-cRaO7fAAagfMxmvyz0p2rpAdMLXZpwRjbQQJCqRd7_8SecaDSYklLOYLTeAJM_KbhxfABEEKGdIcV9p3b1tzD7aVQ_yqk6nF9xv/s1600/Ali+%2526+Aitchison+2014+Fig1.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="243" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiC0PpZspEAFknHObgCzcMQ5y4HqaVm_qNnVCtgVVRK-cRaO7fAAagfMxmvyz0p2rpAdMLXZpwRjbQQJCqRd7_8SecaDSYklLOYLTeAJM_KbhxfABEEKGdIcV9p3b1tzD7aVQ_yqk6nF9xv/s320/Ali+%2526+Aitchison+2014+Fig1.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Map showing the estimated age of each of the<br />Galápagos Islands. From <a href="http://s3.amazonaws.com/academia.edu.documents/38570700/Galapagos.pdf?AWSAccessKeyId=AKIAJ56TQJRTWSMTNPEA&Expires=1480258009&Signature=qOdzv7XGQ8hTJYqNLs8GDZPY9J0%3D&response-content-disposition=inline%3B%20filename%3DGalapagos.pdf">Ali & Aitchison 2014</a></td></tr>
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Galápagos Racers colonized the Galápagos Islands from mainland South America, just like all of the other Galápagos fauna and flora. The modern Galápagos Islands <a href="http://pages.uoregon.edu/drt/Research/Volcanic%20Galapagos/presentation.view@_id=9889959127044&_page=1&_part=2&.html">formed from volcanoes</a> over the past 4 to 5 million years, although some of them have been building beneath the ocean surface for up to 15 million years. It is thought that there have been islands in the Galápagos for at least 8 million years, but the oldest islands have eroded and are now back beneath the ocean surface.</div>
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Because the Galápagos Islands are located only six hundred miles off the coast of Ecuador, <a href="http://www.islandbiogeography.org/">it is easier for them to be colonized by plants and animals from the mainland</a> than for a more remote island chain such as Hawaii (which is >2,500 miles away from the nearest snake-inhabited landmass).</div>
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Molecular dating of the divergence time between <span style="text-align: justify;">Galápagos Racers and their closest mainland relative, </span><a href="http://reptile-database.reptarium.cz/species?genus=Pseudalsophis&species=elegans&search_param=%28%28genus%3D%27Pseudalsophis%27%29%29" style="text-align: justify;"><i>Pseudalsophis </i><i>elegans</i></a><span style="text-align: justify;">, suggests that it has been about</span> 15 million years since they last shared a common ancestor. This suggests that the mainland ancestor of Galápagos Racers probably went extinct sometime over the last 15 million years, and that the ancestors of Galápagos Racers probably colonized the Galápagos Islands before any of the current islands existed (<a href="http://bbcd.bio.uniroma1.it/clone_bbcd/sites/default/files/file%20lezioni/Iguanas.pdf">as is also the case for the Marine Iguanas</a>). Until genetic work is done, we won't know how many times snakes colonized the Galápagos archipelago or how many distinct lineages there are. <a href="https://www.researchgate.net/project/Phylogeograpy-and-diversity-of-Galapagos-terrestrial-snakes-Pseudalsophis-spp"><i>[Edit 12/30/2016: I have recently learned that Massey University ecologist Luis Ortiz-Catedral and his colleagues are working to understand the evolution of all the species in the genus </i>Pseudalsophis<i> and definitively answer this question.]</i></a></div>
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<b><span style="font-size: large;">Could the film have been staged?</span></b></div>
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Obviously the scenes are spliced together, but in my opinion there's no chance the Galápagos National Park would allow something like this to be staged. They are among the strictest places in the world for researchers to conduct scientific work. However, more recent episodes of Planet Earth II <a href="http://www.huffingtonpost.co.uk/entry/planet-earth-2-bbc-sound-effects_uk_5836a77ee4b0b60ceea98dc3">have been criticized for incorporating fake sound effects</a>.</div>
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<a name="2"><span style="font-size: large;"><b>I want more detail on the complicated taxonomic history of these snakes (said almost no one ever).</b></span></a><br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiH819YlsZalscS5JzbG8wxJXDO6-PFpCuB_QXYXp_L14zYuz2NfL154yMKKbfSfIPsgVXiUmOMCZfLpilb9vKHprUADr_mRYMvckdLaBjg5Da2pmZczKAOvXNbkV9gAJMUnlhDxaW3lO8Q/s1600/Pseudalsophis+cladogram+from+Pyron+et+al+2013.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="340" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiH819YlsZalscS5JzbG8wxJXDO6-PFpCuB_QXYXp_L14zYuz2NfL154yMKKbfSfIPsgVXiUmOMCZfLpilb9vKHprUADr_mRYMvckdLaBjg5Da2pmZczKAOvXNbkV9gAJMUnlhDxaW3lO8Q/s400/Pseudalsophis+cladogram+from+Pyron+et+al+2013.png" width="400" /></a></td></tr>
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One of the few phylogenies to include Galápagos Racers</div>
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From <a href="http://www.biomedcentral.com/content/pdf/1471-2148-13-93.pdf">Pyron et al. 2013</a></div>
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<b></b>Broadly, <i>Pseudalsophis</i> is nested within a large clade of Caribbean, Central, and South American <a href="https://www.google.com/url?sa=t&rct=j&q=&esrc=s&source=web&cd=1&cad=rja&uact=8&ved=0ahUKEwjEqNuAo8nQAhVEIJoKHYQ3AJ4QFggbMAA&url=http%3A%2F%2Fsnakesarelong.blogspot.com%2F2013%2F05%2Fbasics-of-snake-taxonomy.html&usg=AFQjCNF8FypHFQcfYIF8mF20wuIzYciTQw&sig2=cHMYfMDojzmF7aKK1avFuQ&bvm=bv.139782543,d.bGs">xenodontine snakes</a> including, among numerous others, the genus <i>Alsophis</i>, which once contained Galápagos Racers and after which their current genus is named. They have been in a variety of genera since their description, especially <i>Dromicus</i>, which is no longer in use, from 1876 to 1997.<br />
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In 1973, herpetologist Charles Myers wrote: <a href="http://digitallibrary.amnh.org/bitstream/handle/2246/2728//v2/dspace/ingest/pdfSource/nov/N2522.pdf?sequence=1&isAllowed=y">"The classification of colubrid snakes in general, and of South American colubrids in particular, is in a notoriously unsatisfactory state."</a> Unfortunately, we are not that much better off today when it comes to Galápagos Racers. It seems pretty clear that the nearest relative of <i>P. biserialis</i>,<i> P. dorsalis</i>, and <i>P. occidentalis </i>is <a href="http://reptile-database.reptarium.cz/species?genus=Pseudalsophis&species=elegans&search_param=%28%28genus%3D%27Pseudalsophis%27%29%29"><i>Pseudalsophis </i><i>elegans</i></a>, the only species in the genus found on the mainland (in Ecuador, Peru, and extreme northern Chile). Beyond that, there isn't a lot of clarity about their next-closest relatives. They are possibly most closely related to obscure South American "groundsnakes" in the genus <i>Psomophis</i>, or to the even more obscure genus <i>Saphenophis</i>, which was described by Myers as "quite lacking in peculiar or unique features" and so named "in allusion to one incontrovertible fact about these snakes...from the Greek saphenes (evident truth, clear) + ophis (a serpent), meaning 'clearly a snake'". We don't really have a great hypothesis about how the different lineages of Galápagos Racers are related to one another, or even if they are all descended from a single common ancestor, because <a href="https://www.ncbi.nlm.nih.gov/nuccore/?term=pseudalsophis">we only have DNA from one of them</a> so far.<br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhCWrCjGUSVB7jxOGXGPNwISYAZWUJZ5LtO4Q5K5nqT17pR_pyn_z0DHA0IWFGWJ62F0WXh_ftu3BziiQGhgR2kjrF3UEH_2wZTyjzwU1bb0mha-r_XtacwJrqlYJDA8HfmWd0NE16mTUV4/s1600/Ali+%2526+Aitchison+2014+Fig8.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="400" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhCWrCjGUSVB7jxOGXGPNwISYAZWUJZ5LtO4Q5K5nqT17pR_pyn_z0DHA0IWFGWJ62F0WXh_ftu3BziiQGhgR2kjrF3UEH_2wZTyjzwU1bb0mha-r_XtacwJrqlYJDA8HfmWd0NE16mTUV4/s400/Ali+%2526+Aitchison+2014+Fig8.png" width="343" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Hypothesized scenario for the evolution of <i>Pseudalsophis</i> snakes<br />So far, we have no DNA evidence that would support or refute this model<br />From <a href="http://s3.amazonaws.com/academia.edu.documents/38570700/Galapagos.pdf?AWSAccessKeyId=AKIAJ56TQJRTWSMTNPEA&Expires=1480258009&Signature=qOdzv7XGQ8hTJYqNLs8GDZPY9J0%3D&response-content-disposition=inline%3B%20filename%3DGalapagos.pdf">Ali & Aitchison 2014</a></td></tr>
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Two reviews based on morphology addressed this question in the late 1990s. <a href="https://www.researchgate.net/publication/280610746_Galapagos_Terrestrial_Snakes_Biogeography_and_Systematics_Herp_Nat_Hist_51_19-40_1997">The first (Thomas 1997) focused exclusively on Galápagos Racers</a> and suggested that <i>P. biserialis</i>,<i> P. dorsalis</i>, and <i>P. occidentalis</i> are descended from a shared common ancestor with <i>P. elegans</i>, but that <i>P. hoodensis</i> is more closely related to the mainland species <i>Philodryas chammissonis</i>, and that <i>P. slevini</i> and <i>P. steindachneri</i> are most closely related to Caribbean species. <a href="http://digitallibrary.amnh.org/bitstream/handle/2246/1646//v2/dspace/ingest/pdfSource/bul/B240a01.pdf?sequence=1&isAllowed=y">The other study (Zaher 1999)</a>, which looked at <a href="http://snakesarelong.blogspot.com/2014/03/why-do-snakes-have-two-penises.html">hemipene morphology</a> over a much larger group of snakes, disagreed, finding a shared derived character—an inflated papillate ridge, placed far medially, on the medial surface of the lobes—linking the Galápagos Racers together with the mainland species <i>P. elegans</i>. Statements that Galápagos Racers have “very similar hemipenes” notwithstanding, Zaher <a href="http://www.mapress.com/zootaxa/2009/f/zt02067p028.pdf">was criticized</a> for not describing the specific characters uniting the Galápagos species to the exclusion of others.<br />
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<a href="http://www.biodiversitylibrary.org/page/4753171#page/13/mode/1up">Maglio (1970)</a> noted that the tooth counts and arrangement and the and shape of the premaxilla bone was most similar among the three Galápagos species that he examined (<i>P. biserialis</i>, <i>P. dorsalis</i>, and <i>P. slevini</i>), and different from the West Indian species that Taylor later suggested are <i>P. slevini</i>'s closest relatives. More recently, <a href="https://www.researchgate.net/profile/Hussam_Zaher/publication/236963821_Molecular_phylogeny_of_the_New_World_Dipsadidae_(Serpentes_Colubroidea)_a_reappraisal/links/00b7d51a78e574f124000000.pdf">a study led by Grazziotin</a> claimed that they "unequivocally support...Zaher' s (1999) hypothesis based on morphology that continental <i>Pseudalsophis elegans</i> is closely related to the Galápagos Island species of Xenodontinae (herein represented by <i>Pseudalsophis dorsalis</i>), rather than to West Indian <i>Alsophis </i>and <i>Antillophis</i>, and mainland <i>Philodryas </i>(Thomas, 1997)." However, they obviously didn't read Thomas's paper very carefully, because he also hypothesizes that <i>P. dorsalis</i> is closely related to <i>P. elegans</i>, and the Grazziotin paper didn't sequence any DNA from <i>P. slevini</i>,<i> P. </i><i>steindachneri</i>, or <i>P. hoodensis</i>, and therefore didn't test any hypotheses about them.<br />
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As for whether or not the snakes in Planet Earth II should be called <i>P. occidentalis</i> or <i>P. dorsalis occidentalis</i>, that's really a lumper/splitter question. But, both the <a href="http://www.iucnredlist.org/details/full/190540/0">IUCN</a> and the <a href="https://books.google.com/books?id=zZ3NBQAAQBAJ">2014 edition of <i>Snakes of the World</i></a><i> </i>recognize <i>P. occidentalis</i> as a full species; it was originally described as such <a href="http://www.biodiversitylibrary.org/item/98588#page/392/mode/2up">by Van Denburgh in 1912</a>, sunk to a subspecies of <i>P. dorsalis</i> by Mertens in 1960, and re-elevated to a full species <a href="http://digitallibrary.amnh.org/bitstream/handle/2246/1646//v2/dspace/ingest/pdfSource/bul/B240a01.pdf?sequence=1&isAllowed=y">in a 1999 paper by Zaher</a> that was not primarily concerned with taxonomy and appears to have subsequently been neglected. <a href="http://reptile-database.reptarium.cz/advanced_search?genus=Pseudalsophis&submit=Search">The Reptile Database</a> is currently a holdout for the subspecies designation, which has not been disputed but which is also not explicitly supported by unambiguous data. Perhaps wisely, <a href="http://galapagosconservation.org.uk/wildlife/galapagos-racer/">the official webpage of Galápagos National Park</a> chooses not to use scientific names and refers to the Fernandina racers as the "western subspecies". The truth is that, until more research is done, we won't be able to settle on an accurate taxonomy for these snakes.</div>
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<a name="1"><b>1 </b></a>This sounds a bit redundant to a snake biologist, but it isn't incorrect. The one thing that I wish BBC programs would do is identify the species in them more precisely. I'm advocating for a "biologist mode" that can be activated which would show the location and identity of species in all clips, similar to <a href="https://en.wikipedia.org/wiki/Pop-Up_Video">the old MTV show <i>Pop-up Video</i></a>.<a href="#top1"><sup>↩</sup></a><br />
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ACKNOWLEDGMENTS</div>
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Thanks to <a href="https://www.flickr.com/photos/38984611@N03/16081314279">Andy Kraemer</a> and <a href="https://www.flickr.com/photos/79076791@N00/7103341337/in/album-72157629510285568/">Jim Moulton</a> for the use of their photographs.</div>
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REFERENCES</div>
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<span style="font-size: x-small;">Ali, J. R. and J. C. Aitchison. 2014. Exploring the combined role of eustasy and oceanic island thermal subsidence in shaping biodiversity on the Galápagos. Journal of Biogeography 41:1227-1241 <<a href="http://s3.amazonaws.com/academia.edu.documents/38570700/Galapagos.pdf?AWSAccessKeyId=AKIAJ56TQJRTWSMTNPEA&Expires=1480258009&Signature=qOdzv7XGQ8hTJYqNLs8GDZPY9J0%3D&response-content-disposition=inline%3B%20filename%3DGalapagos.pdf">full-text</a>></span><br />
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<span style="font-size: x-small;">Bisconti, M., W. Landini, G. Bianucci, G. Cantalamessa, G. Carnevale, L. Ragaini, and G. Valleri. 2001. Biogeographic relationships of the Galapagos terrestrial biota: parsimony analyses of endemicity based on reptiles, land birds and Scalesia land plants. Journal of Biogeography 28:495-510 <<a href="https://www.researchgate.net/profile/Giorgio_Carnevale/publication/230229760_Biogeographic_relationships_of_the_Galapagos_terrestrial_biota_parsimony_analyses_of_endemicity_based_on_reptiles_land_birds_and_Scalesia_land_plants/links/02e7e528373908ddc2000000.pdf">full-text</a>></span><br />
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<span style="font-size: x-small;">Carpenter, C. C. 1966. The marine iguana of the Galapagos Islands, its behavior and ecology. Proceedings of the California Academy of Sciences (Series 4) 34:329-376 <<a href="http://www.biodiversitylibrary.org/item/53858#page/342/mode/2up">full-text</a>></span><br />
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<span style="font-size: x-small;">Carpenter, C. C. 1984. Dominance in snakes. Special Publication, University of Kansas Museum of Natural History 10:195-202 <<a href="http://www.biodiversitylibrary.org/item/25758#page/209/mode/1up">full-text</a>></span><br />
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<span style="font-size: x-small;">Christian, K. A. and C. R. Tracy. 1981. The effect of the thermal environment on the ability of hatchling Galapagos land iguanas to avoid predation during dispersal. Oecologia 49:218-223 </span><span style="font-size: x-small;"><</span><a href="http://link.springer.com/article/10.1007/BF00349191" style="font-size: small;">abstract</a><span style="font-size: x-small;">></span><br />
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<span style="font-size: x-small;">Geist, D., H. Snell, H. Snell, C. Goddard, and M. Kurz. 2014. A paleogeographic model of the Galápagos Islands and biogeographical and evolutionary implications. The Galápagos: a natural laboratory for the Earth Sciences. American Geophysical Union, Washington DC, USA:145-166 <<a href="http://www.umsl.edu/~parkerp/geistdraft.pdf">full-text</a>></span><br />
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<span style="font-size: x-small;">Grazziotin, F. G., H. Zaher, R. W. Murphy, G. Scrocchi, M. A. Benavides, Y.-P. Zhang, and S. L. Bonattoh. 2012. Molecular phylogeny of the New World Dipsadidae (Serpentes: Colubroidea): a reappraisal. Cladistics 28:437-459 <<a href="https://www.researchgate.net/profile/Hussam_Zaher/publication/236963821_Molecular_phylogeny_of_the_New_World_Dipsadidae_(Serpentes_Colubroidea)_a_reappraisal/links/00b7d51a78e574f124000000.pdf">full-text</a>></span></div>
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<span style="font-size: x-small;">Grehan, J. 2001. Biogeography and evolution of the Galápagos: integration of the biological and geological evidence. Biological Journal of the Linnean Society 74:267-287 <<a href="http://www.ib.usp.br/~silvionihei/pdf/Grehan%202001_Galapagos.pdf">full-text</a>></span></div>
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<span style="font-size: x-small;">Günther, A. 1860. On a new snake from the Galápagos islands. The Annals and Magazine of Natural History 3:78-79 <<a href="http://www.biodiversitylibrary.org/page/2267027#page/98/mode/1up">full-text</a>></span></div>
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<span style="font-size: x-small;">Hedges, S. B., A. Couloux, and N. Vidal. 2009. Molecular phylogeny, classification, and biogeography of West Indian racer snakes of the Tribe Alsophiini (Squamata, Dipsadidae, Xenodontinae). Zootaxa 2067:1-28 <<a href="http://www.mapress.com/zootaxa/2009/f/zt02067p028.pdf">full-text</a>></span></div>
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<span style="font-size: x-small;">Knapp, C. R., S. Alvarez-Clare, and C. Perez-Heydrich. 2010. The influence of landscape heterogeneity and dispersal on survival of neonate insular iguanas. Copeia 2010:62-70 <<a href="https://www.researchgate.net/profile/Charles_Knapp2/publication/250067012_The_Influence_of_Landscape_Heterogeneity_and_Dispersal_on_Survival_of_Neonate_Insular_Iguanas/links/555b7e7208ae8f66f3ad7946.pdf">full-text</a>></span><br />
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<span style="font-size: x-small;">Laurie, W. and D. Brown. 1990. Population biology of marine iguanas (<i>Amblyrhynchus cristatus</i>). II. Changes in annual survival rates and the effects of size, sex, age and fecundity in a population crash. Journal of Animal Ecology 59:529-544 <<a href="https://www.researchgate.net/publication/271696486_Population_Biology_of_Marine_Iguanas_Amblyrhynchus_cristatus_II_Changes_in_Annual_Survival_Rates_and_the_Effects_of_Size_Sex_Age_and_Fecundity_in_a_Population_Crash">full-text</a>></span></div>
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<span style="font-size: x-small;">Maglio, V. J. 1970. West Indian xenodontine colubrid snakes: their probable origin, phylogeny, and zoogeography. Bulletin of the Museum of Comparative Zoology 141:1-54 <<a href="http://www.biodiversitylibrary.org/page/4753171#page/13/mode/1up">full-text</a>></span></div>
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<span style="font-size: x-small;">Merlen, G. and R. A. Thomas. 2013. A Galapagos ectothermic terrestrial snake gambles a potential chilly bath for a protein-rich dish of fish. Herpetological Review 44:415-417 <<a href="http://www.darwinfoundation.org/media/filer_public/2013/10/10/merlen__thomas_2013.pdf">full-text</a>></span></div>
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<span style="font-size: x-small;">Mertens, R. 1960. Über die schlangen der Galápagos. Senckenbergiana Biologica 41:133-141 <not available online></span></div>
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<span style="font-size: x-small;">Myers, C. W. 1973. A new genus for Andean snakes related to <i>Lygophis boursieri </i>and a new species (Colubridae). American Museum Novitates 2522 <<a href="http://digitallibrary.amnh.org/bitstream/handle/2246/2728//v2/dspace/ingest/pdfSource/nov/N2522.pdf?sequence=1&isAllowed=y">full-text</a>></span><br />
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<span style="font-size: x-small;">Parent, C. E., A. Caccone, and K. Petren. 2008. Colonization and diversification of Galápagos terrestrial fauna: a phylogenetic and biogeographical synthesis. Philosophical Transactions of the Royal Society B: Biological Sciences 363:3347-3361 <<a href="http://rstb.royalsocietypublishing.org/content/363/1508/3347.short">full-text</a>></span></div>
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<span style="font-size: x-small;">Pyron, R. A., F. Burbrink, and J. J. Wiens. 2013. A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology 13:93 <<a href="http://bmcevolbiol.biomedcentral.com/articles/10.1186/1471-2148-13-93">full-text</a>></span><br />
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<span style="font-size: x-small;">Pyron, R. A., J. Guayasamin, N. Peñafiel, L. Bustamante, and A. Arteaga. 2015. Systematics of Nothopsini (Serpentes, Dipsadidae), with a new species of <i>Synophis </i>from the Pacific Andean slopes of southwestern Ecuador. ZooKeys 541:109-147 <<a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4714381/">full-text</a>></span></div>
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<span style="font-size: x-small;">Radder, R. S. and R. Shine. 2007. Why do female lizards lay their eggs in communal nests? Journal of Animal Ecology 76:881-887 <<a href="http://onlinelibrary.wiley.com/doi/10.1111/j.1365-2656.2007.01279.x/full">full-text</a>></span><br />
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<span style="font-size: x-small;">Rassmann, K. 1997. Evolutionary age of the Galápagos iguanas predates the age of the present Galápagos Islands. Molecular Phylogenetics and Evolution 7:158-172 </span><span style="font-size: x-small;"><</span><a href="http://www.jstor.org/stable/1565698" style="font-size: x-small;">full-text</a><span style="font-size: x-small;">></span><br />
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<span style="font-size: x-small;">Rodríguez-Durán, A. 1996. Foraging ecology of the Puerto Rican boa (<i>Epicrates inornatus</i>): bat predation, carrion feeding, and piracy. Journal of Herpetology 30:533-536<span style="font-size: x-small;"><</span><a href="http://bbcd.bio.uniroma1.it/clone_bbcd/sites/default/files/file%20lezioni/Iguanas.pdf" style="font-size: x-small;">full-text</a><span style="font-size: x-small;">></span></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Shine, R., L. X. Sun, M. Fitzgerald, and M. Kearney. 2002. Accidental altruism in insular pit-vipers (<i>Gloydius shedaoensis</i>, Viperidae). Evolutionary Ecology 16:541-548 <<a href="http://www.kingsnake.com/aho/pdf/November2003Update/shine2002c.pdf">full-text</a>></span><br />
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<div style="text-align: justify;">
<span style="font-size: x-small;">Steindachner, F. 1876. Die schlangen und eidechsen der Galapagos-inseln. Zoologisch-botanischen Gesellschaft, Wien, Germany <<a href="https://books.google.com/books?hl=en&lr=&id=pgQjL4iSaEIC&oi=fnd&pg=PT3&dq=Die+schlangen+und+eidechsen+der+Galapagos-inseln.+Zoologisch-botanischen+Gesellschaft,+Wien,+Germany.&ots=1M0B2y_1cS&sig=jPHIyIx6o1AitAn9qH0wy0LCj_M#v=onepage&q&f=false">Google book</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Swash, A. and R. Still. 2000. Birds, Mammals and Reptiles of the Galapagos Islands. Pica Press <<a href="https://www.amazon.com/Birds-Mammals-Reptiles-Gal%C3%A1pagos-Islands/dp/0300115326">Amazon</a>></span></div>
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<span style="font-size: x-small;">Thomas, R. 1997. Galapagos terrestrial snakes: biogeography and systematics. Herpetological Natural History 5:19-40 <<a href="https://www.researchgate.net/publication/280610746_Galapagos_Terrestrial_Snakes_Biogeography_and_Systematics_Herp_Nat_Hist_51_19-40_1997">full-text</a>></span></div>
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<div style="text-align: justify;">
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Van Denburgh, J. 1912. Expedition of the California Academy of Sciences to the Galápagos Islands, 1905-1906. IV. The snakes of the Galapagos Islands. Proceedings of the California Academy of Sciences (Series 4) 1:323-374 <<a href="http://www.biodiversitylibrary.org/item/98588#page/392/mode/2up">full-text</a>></span></div>
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<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Wallach, V. W., Kenneth J. and J. Boundy. 2014. Snakes of the World: A Catalogue of Living and Extinct Species. CRC Press, Boca Raton, Florida, USA <<a href="https://books.google.com/books?id=zZ3NBQAAQBAJ">Google book</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Weinstein, S. A., D. A. Warrell, J. White, and D. E. Keyler. 2011. "Venomous" Bites from Non-Venomous Snakes: A Critical Analysis of Risk and Management of "Colubrid" Snake Bites. Elsevier, Amsterdam <<a href="https://books.google.com/books?id=lB3FxFG5wrQC&printsec=frontcover&source=gbs_ge_summary_r&cad=0#v=onepage&q&f=false">Google book</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Werner, D. I. 1983. Reproduction in the iguana <i>Conolophus subcristatus </i>on Fernandina Island, Galapagos: clutch size and migration costs. American Naturalist 121:757-775 <<a href="https://www.jstor.org/stable/2460852">abstract</a>></span><br />
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<span style="font-size: x-small;">Yeager, C. P. and G. M. Burghardt. 1991. Effect of food competition on aggregation: evidence for social recognition in the plains garter snake (<i>Thamnophis radix</i>). Journal of Comparative Psychology 105:380-386 <<a href="http://psycnet.apa.org/journals/com/105/4/380/">abstract</a>></span><br /><br />
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<span style="font-size: x-small;">Zaher, H. 1999. Hemipenial morphology of the South American xenodontine snakes, with a proposal for a monophyletic Xenodontinae and a reappraisal of colubroid hemipenes. Bulletin of the American Museum of Natural History 240:1-168 <<a href="http://digitallibrary.amnh.org/bitstream/handle/2246/1646//v2/dspace/ingest/pdfSource/bul/B240a01.pdf?sequence=1&isAllowed=y">full-text</a>></span></div>
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<span style="font-size: x-small;">Zaher, H., F. G. Grazziotin, J. E. Cadle, R. W. Murphy, J. C. Moura-Leite, and S. L. Bonatto. 2009. Molecular phylogeny of advanced snakes (Serpentes, Caenophidia) with an emphasis on South American Xenodontines: A revised classification and descriptions of new taxa. Papeis Avulsos de Zoologia (Sao Paulo) 49:115-153 <<a href="http://www.scielo.br/scielo.php?pid=S0031-10492009001100001&script=sci_arttext">full-text</a>></span></div>
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="http://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
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Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com14Jena, Germany50.927054 11.58923720000007150.7669025 11.26651370000007 51.087205499999996 11.911960700000071tag:blogger.com,1999:blog-7443075087825368900.post-29789782085127437922016-10-28T11:03:00.001-06:002017-03-25T03:10:01.732-06:00Snakes with feet, anti-goo saliva, and more recent updates<div style="text-align: center;">
<span style="font-family: inherit; font-size: x-small;">This post will soon be available in Spanish</span></div>
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<span style="font-family: inherit;">More of the latest snake news and research (for other recent updates, see posts from <a href="http://snakesarelong.blogspot.com/2016/03/state-snakes-linnaean-names-and-other.html">March</a>, <a href="http://snakesarelong.blogspot.com/2016/06/virgin-birth-color-of-fossil-snakes-and.html">June</a>, and <a href="http://snakesarelong.blogspot.com/2016/08/why-snakes-are-long-and-other-recent.html">August</a>)—and, perhaps the most exciting news of all is that I have defended my dissertation and will be returning to writing more in-depth content in the next few months!</span><br />
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<b><span style="font-family: inherit;">Rattlesnake Roundups (<a href="http://snakesarelong.blogspot.com/2015/03/rattlesnake-roundups.html">I</a> and <a href="http://snakesarelong.blogspot.com/2016/05/rattlesnake-roundups-revisited.html">II</a>)</span></b><br />
<b><span style="font-family: inherit;"><br /></span></b>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiyRnFQlX54irwuCVS9OCYlXT6nyciGHxXzJaq0RUQu8teI5YTKse_qYmK5O3kHlUsd45G2XBivVgdAmiBTxRsYPWacbO9slJ-W4Ua_iHJbB2Z52Oy3oBwcVksj8D8TDzCJ5ua926TuQ0Se/s1600/rattlesnake_license_plate.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="153" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiyRnFQlX54irwuCVS9OCYlXT6nyciGHxXzJaq0RUQu8teI5YTKse_qYmK5O3kHlUsd45G2XBivVgdAmiBTxRsYPWacbO9slJ-W4Ua_iHJbB2Z52Oy3oBwcVksj8D8TDzCJ5ua926TuQ0Se/s320/rattlesnake_license_plate.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><a href="http://www.conservationplate.org/">A Texas conservation licence plate</a> ironically depicting<br />
a Western Diamond-backed Rattlesnake (<i>Crotalus atrox</i>).<br />
Funds from these plates support <a href="http://www.conservationplate.org/projects.phtml#">a variety of valuable<br />conservation projects in Texas</a> under the <a href="http://tpwd.texas.gov/publications/pwdpubs/pwd_pl_w7000_1187a/">Texas<br />Wildlife Action Plan</a>, although none are specific to snakes.</td></tr>
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<span style="font-family: inherit;">Advocates for increasing state oversight of rattlesnake roundups in Texas <a href="http://www.statesman.com/news/potential-snake-gassing-ban-killed-texas-parks-and-wildlife/IyEb1P2XcG2vNmI8r7CxkJ/">received disappointing news this week</a> when the <a href="http://tpwd.texas.gov/about/tpw-commissioners">Texas Parks and Wildlife Commission</a> decided that they would not support <a href="http://tpwd.texas.gov/huntwild/wild/wildlife_diversity/nongame/media/TPWD-SHWG-Report.pdf">a proposed ban</a> on using gasoline fumes to collect rattlesnakes<span style="text-align: start;">. Rather than reviewing and voting on the issue at their bi-annual meeting next month, the TPW Commission decided to remove it from their agenda entirely, citing "</span><span style="text-align: start;">insufficient support from legislative oversight or the potentially regulated community". </span></span><span style="font-family: inherit;">This decision marks the second time reviewing the ban has been put off, and unfortunately it is likely to be the last until the effort to reform roundups is re-initiated. The announcement included the statement that </span><span style="font-family: inherit;"><i>"TPWD [Texas Parts and Wildlife Department] staff still believe that there are better options for collecting snakes that do not adversely impact non-target species, and we will continue to work with the snake collecting community to develop and implement best practices that reduce potential impacts to these species"</i>, although in the absence of specific details it is hard to believe that this issue will remain at the fore of wildlife management in Texas without continued pressure from advocates of scientific rattlesnake management. </span><span style="font-family: inherit;">However, Representative Susan King of Sweetwater's 2015 <a href="http://www.legis.state.tx.us/tlodocs/84R/billtext/html/HB00763I.htm">house bill 763</a> requires that </span>petitions to state agencies (including TPWD) that are signed by <51% Texas residents are not valid, which means that the ability of non-Texans to influence policy on this issue is now greatly diminished.<br />
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<span style="font-family: inherit;"><br /></span><span style="font-family: inherit;">If you're not familiar with the issues surrounding the gassing ban, I encourage you to read the <a href="http://tpwd.texas.gov/huntwild/wild/wildlife_diversity/nongame/media/TPWD-SHWG-Report.pdf">2016 Snake Harvest Working Group report</a>, the same document that was available to the TPW Commission prior to their decision this week. Among other topics, it contains data on </span><a href="http://tpwd.texas.gov/huntwild/wild/wildlife_diversity/nongame/media/TPWD-SHWG-References.pdf">the adverse impacts of gassing on non-target endangered species</a>, which is the primary impetus for the ban. It hints at human health impacts of consuming meat from gassed rattlesnakes. The SHWG report also summarizes previously unavailable data on roundup economics, <a href="http://snakesarelong.blogspot.com/2016/05/rattlesnake-roundups-revisited.html">showing that profits are not related to the number of rattlesnakes at an event</a> and did not decline after gassing was banned in Alabama and Oklahoma. Stakeholder survey responses and <a href="http://tpwd.texas.gov/huntwild/wild/wildlife_diversity/nongame/media/TPWD-SHWG-References.pdf">the vast majority (>90%) of public comments</a> from Texans were in favor of the gassing ban, as are many TWPD employees.<br />
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The <a href="http://tpwd.texas.gov/about/tpw-commissioners">TPW Commission</a> is solely responsible for this decision. <span style="font-family: inherit;">You can let </span><a href="http://tpwd.texas.gov/about/tpw-commissioners" style="font-family: inherit;">the TPW Commission</a><span style="font-family: inherit;"> and</span><span style="font-family: inherit;"> </span><a href="http://www.house.state.tx.us/members/member-page/email/?district=71&session=84" style="font-family: inherit;">Texas State Representative Susan King of Sweetwater</a> (or your own state representative, if you live in Texas) <span style="font-family: inherit;">know whether you think they are acting in the best interest of the majority of the public and <a href="https://strangebehaviors.wordpress.com/2016/10/28/texas-shows-its-too-scared-to-stop-folks-from-gassing-wildlife/">in accordance with game management principles</a></span><span style="font-family: inherit;"> at the links provided (if you no longer have a fax machine, you can send a fax over the Internet </span><a href="http://faxzero.com/" style="font-family: inherit;">here</a><span style="font-family: inherit;">).</span></div>
<span style="font-family: inherit;"><br /></span>
<b><span style="font-family: inherit;"><a href="http://snakesarelong.blogspot.com/2012/08/goo-eating-snakes-and-eggs-that-evade.html">Goo-eating Snakes and the Eggs that Evade Them</a> and <a href="http://snakesarelong.blogspot.com/2013/09/basics-of-snake-fangs.html">Basics of Snake Fangs</a></span></b><br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjhO0Pwrodtyvel3VdI4hE9YXZOoKfxY-boMfGmBOf4KGnA_NRGqIdEPjK1w2RVBtpNno-AqwYS8E1eDap_-JHrREThcQwlyyfb2TNUgSgol9Qv-5nNsiS7toSnbEH3FcmZHb3qef-9zYt8/s1600/Dipsas+alternans+mandibular+glands+Zaher+et+al+2014+Fig5.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><span style="font-family: inherit;"><img border="0" height="273" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjhO0Pwrodtyvel3VdI4hE9YXZOoKfxY-boMfGmBOf4KGnA_NRGqIdEPjK1w2RVBtpNno-AqwYS8E1eDap_-JHrREThcQwlyyfb2TNUgSgol9Qv-5nNsiS7toSnbEH3FcmZHb3qef-9zYt8/s320/Dipsas+alternans+mandibular+glands+Zaher+et+al+2014+Fig5.png" width="320" /></span></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><span style="font-family: inherit;">Mandibular glands of <i>Dipsas alternans</i></span><br />
<span style="font-family: inherit;">From <a href="http://bmcevolbiol.biomedcentral.com/articles/10.1186/1471-2148-14-58">Zaher et al. 2014</a></span></td></tr>
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<span style="font-family: inherit;">This discovery is from 2014, but it's newer than either of the past posts to which it's germane and I just found out about it. Perhaps you've seen <a href="http://video.nationalgeographic.com/video/news/160615-treefrog-tadpoles-escape-hatching-vin?utm_source=Facebook&utm_medium=Social&utm_content=link_fb20160620video-tadpolesvod&utm_campaign=Content&sf29141940=1">the incredible rapid hatching behavior that treefrog eggs have evolved to escape</a> from snake predators, including cat-eyed snakes (genus <i>Leptodeira</i>), blunt-headed tree snakes (genus <i>Imantodes</i>), and snail-sucking snakes (genera <i>Sibon</i> and <i>Dipsas</i>). These snakes also eat a variety of other gooey prey, such as earthworms, leeches, <a href="https://static-content.springer.com/esm/art%3A10.1186%2F1471-2148-14-58/MediaObjects/12862_2013_2568_MOESM5_ESM.wmv">snails</a>, slugs, adult frogs, caecilians, and, more rarely, non-gooey prey like lizards and reptile eggs. They have a number of adaptations that help them consume their sticky, viscous prey, including long, slender teeth, skull bones and muscles modified for extreme lower jaw extrusion, and a short-snouted, large-eyed look that resembles a snake embryo. <a href="http://bmcevolbiol.biomedcentral.com/articles/10.1186/1471-2148-14-58">Recently, a team of scientists from Brazil discovered a new one</a>: a protein-secretion delivery system in the lower jaw.</span><br />
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<span style="font-family: inherit;">Are the secretions venom? No. <i>Dipsas</i> and its relatives always extract snails <a href="https://static-content.springer.com/esm/art%3A10.1186%2F1471-2148-14-58/MediaObjects/12862_2013_2568_MOESM5_ESM.wmv">using a sudden strike</a>, followed by fast, alternating probing motions of the mandible inside the shell; this behavior could hardly depend on a chemical reaction of any kind. Instead, the gland secretions probably play a role in mucus control and prey transport rather than immobilization or killing of the prey. Although the glands in some species are associated with muscles, they are <a href="http://snakesarelong.blogspot.com/2013/09/basics-of-snake-fangs.html">not connected to any teeth</a>, but rather open onto the floor of the mouth, which in some species is covered with extensively loose, folded skin. <a href="https://www.researchgate.net/publication/257814973_Unusual_Labial_Glands_in_Snakes_of_the_Genus_Geophis_Wagler_1830_Serpentes_Dipsadinae?el=1_x_8&enrichId=rgreq-b8861264b928d76a311e7ff933cdfaf2-XXX&enrichSource=Y292ZXJQYWdlOzI2MTA2Njg3ODtBUzo5OTkzMTMxNDQ1ODYzN0AxNDAwODM2ODkwODA2">Hypertrophied infralabial glands</a> have been known from some dipsadine species since the 1960s, but the new paper describes the muscles and other soft tissues surrounding them and documents their variation among several dozen species of this very speciose group of snakes. On the other side of the world, <a href="http://snakesarelong.blogspot.com/2012/04/asymmetrical-snakes.html">pareatid snail-eating snakes</a> have independently evolved a similar lifestyle, complete with <a href="https://www.researchgate.net/publication/250068393_On_the_rictal_structures_of_some_snakes?el=1_x_8&enrichId=rgreq-b8861264b928d76a311e7ff933cdfaf2-XXX&enrichSource=Y292ZXJQYWdlOzI2MTA2Njg3ODtBUzo5OTkzMTMxNDQ1ODYzN0AxNDAwODM2ODkwODA2">upper jaw glands of perhaps similar function</a>.</span><br />
<br />
<a href="http://snakesarelong.blogspot.com/2016/08/why-snakes-are-long-and-other-recent.html"><b>Why snakes are long</b></a> and <b><a href="http://snakesarelong.blogspot.com/2014/03/why-do-snakes-have-two-penises.html">Why do snakes have two penises?</a></b><br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhJ1FPj0EEo9k78equkKHojEO3mWIq-9psAnwaF2SanA60StLww9iQ9pKWx5bpp_sGxtZM3SpzmqCwpKgu8cTs6j2o0pVUA2gGfCcvRugzaTNJRDsk4moyTv5s-IHTI3XWNGqSImuuMu8I5/s1600/Leal+%2526+Cohn+2016+Fig4+crop.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="351" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhJ1FPj0EEo9k78equkKHojEO3mWIq-9psAnwaF2SanA60StLww9iQ9pKWx5bpp_sGxtZM3SpzmqCwpKgu8cTs6j2o0pVUA2gGfCcvRugzaTNJRDsk4moyTv5s-IHTI3XWNGqSImuuMu8I5/s400/Leal+%2526+Cohn+2016+Fig4+crop.jpg" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Pelvic girdles (dark blue) and hind limbs (red) of lizards,<br />
living snakes, and extinct snakes with fully-developed limbs.<br />
ZRS is the name of the SHH enhancer gene<br />
that has been partially deleted in snakes.<br />
From <a href="http://www.sciencedirect.com/science/article/pii/S0960982216310697">Leal & Cohn 2016</a></td></tr>
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Many people are familiar with the tiny vestigial legs or "spurs" of boas, pythons, and other <a href="http://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">henophidian</a> snakes. These structures are sexually dimorphic and are used by male boas and pythons <a href="http://www.jstor.org/stable/3891678">in male-male combat</a> and also to <a href="http://www.jstor.org/stable/1444292">titillate females before and during mating</a>. <a href="http://www.sciencedirect.com/science/article/pii/S0960982216310697">New data from the University of Florida</a> describes how the spurs are formed: a weak flicker of activity by a gene called <i>Sonic hedgehog</i> (SHH) during the first few hours of embryonic development, in contrast to strong, sustained activity of this gene in lizard embryos throughout their development, forming legs. In snakes, unique genetic deletions from an enhancer of SHH explain its weak activity; transgenic mouse embryos with the same deletions showed similarly weak SHH activity, whereas mouse embryos grown with a lizard enhancer developed normally. <a href="http://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">Caenophidian snakes</a>, such as vipers, gartersnakes, and cobras, had more extreme deletions and mutations, with the cobra barely retaining any of the SHH enhancer gene.<br />
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Amazingly, the researchers also found that HOXD13, the part of the limb-building gene that's responsible for building hands and feet, was unaltered in python embryos, and that python embryos develop not just a pelvic girdle and femur, which form the spur in adulthood, but cartilaginous templates of a tibia, fibula, and foot, which are reabsorbed prior to hatching. Although living snakes appear to follow a gradual pattern of limb shrinkage and loss, some extinct snakes that are thought to have been more similar to boas and pythons than they were to blindsnakes also had fully-developed, albeit small, limbs, complete with feet, as adults. This new discovery helps explain the apparent evolutionary "re-appearance" of these structures; they were never completely lost in the first place. As for the reason why not, <a href="http://www.evodevo.net/uploads/1/8/1/3/18132731/leal_python.pdf">snake HOXD genes and their regulators appear to be equally important to the development of their paired hemipenes</a>, structures of obvious importance.<br />
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REFERENCES</div>
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<span style="font-size: x-small;">Oliveira, L., A. L. Costa Prudente, and H. Zaher. 2014. Unusual labial glands in snakes of the genus <i>Geophis </i>Wagler, 1830 (Serpentes: Dipsadinae). Journal of Morphology 275:87-99 <<a href="https://www.researchgate.net/publication/257814973_Unusual_Labial_Glands_in_Snakes_of_the_Genus_Geophis_Wagler_1830_Serpentes_Dipsadinae?el=1_x_8&enrichId=rgreq-b8861264b928d76a311e7ff933cdfaf2-XXX&enrichSource=Y292ZXJQYWdlOzI2MTA2Njg3ODtBUzo5OTkzMTMxNDQ1ODYzN0AxNDAwODM2ODkwODA2">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Leal, F. & Cohn, M.J. 2016. Loss and re-emergence of legs in snakes by modular evolution of Sonic hedgehog and HOXD enhancers. Current Biology DOI:10.1016/j.cub.2016.09.020 <<a href="http://www.sciencedirect.com/science/article/pii/S0960982216310697">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Leal, F. & Cohn, M.J. 2014. Development of hemipenes in the ball python snake <i>Python regius</i>. Sexual Development, 9, 6-20 <<a href="http://www.evodevo.net/uploads/1/8/1/3/18132731/leal_python.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;"> Savitzky, A.H. 1983. Coadapted character complexes among snakes: fossoriality, piscivory, and durophagy. American Zoologist, 23, 397-409 <<a href="http://az.oxfordjournals.org/content/23/2/397.abstract">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Texas Parks and Wildlife Department. 2016. Snake Harvest Working Group Final Report <<a href="http://tpwd.texas.gov/huntwild/wild/wildlife_diversity/nongame/media/TPWD-SHWG-Report.pdf">link</a>> <<a href="http://tpwd.texas.gov/huntwild/wild/wildlife_diversity/nongame/media/TPWD-SHWG-References.pdf">references</a>> <<a href="http://tpwd.texas.gov/huntwild/wild/wildlife_diversity/nongame/media/TPWD-SHWG-Executive-Summary.pdf">summary</a>><br /><br />Zaher, H., de Oliveira, L., Grazziotin, F.G., Campagner, M., Jared, C., Antoniazzi, M.M. & Prudente, A.L. 2014. Consuming viscous prey: a novel protein-secreting delivery system in neotropical snail-eating snakes. BMC Evolutionary Biology, 14, 1-28 <<a href="http://bmcevolbiol.biomedcentral.com/articles/10.1186/1471-2148-14-58">link</a>></span></div>
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<span style="font-family: inherit; font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="https://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com0Jena, Germany50.927054 11.58923720000007150.7669025 11.26651370000007 51.087205499999996 11.911960700000071tag:blogger.com,1999:blog-7443075087825368900.post-9834878255040233082016-09-28T10:43:00.000-06:002020-05-04T07:41:45.887-06:00Xenophidion: The Snake with the Mystery Penis<div style="text-align: center;">
<span style="font-size: x-small;">This post will soon be available in Spanish</span></div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhZC7WCck3Tu8_9UVcabdlNGrCe42gXGKBPWh7LTaG5213oOq5cXE7-PzaDcxo2NObkKqkVR-_fz8WwEOkkyQYaSv9effwH1B8ilEVcB-X-DFu4Qqtm1rH5WfPwgvYOUzPKhvsOAaGK_ojU/s1600/Das+2010+Xenophidion+schaeferi+art+by+Szabolcs+Kokay.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="168" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhZC7WCck3Tu8_9UVcabdlNGrCe42gXGKBPWh7LTaG5213oOq5cXE7-PzaDcxo2NObkKqkVR-_fz8WwEOkkyQYaSv9effwH1B8ilEVcB-X-DFu4Qqtm1rH5WfPwgvYOUzPKhvsOAaGK_ojU/s400/Das+2010+Xenophidion+schaeferi+art+by+Szabolcs+Kokay.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Xenophidion schaeferi</i><br />
From <a href="https://books.google.com/books?id=LRRpBwAAQBAJ&dq=A+field+guide+to+the+reptiles+of+South-East+Asia&lr=&source=gbs_navlinks_s">Das 2010</a>, painted by <a href="http://kokay.hu/eng_index.htm" style="text-align: justify;">Szabolcs Kókay</a></td></tr>
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For a combination of phylogenetically distinct, taxonomically confusing, and poorly known, you simply cannot beat the spinejaw snakes, genus <i>Xenophidion</i>. <a href="https://www.researchgate.net/profile/Ulrich_Manthey/publication/233570693_Xenophidion_a_new_genus_with_two_new_species_of_snakes_from_Malaysia_Serpentes_Colubridae/links/5602a86308ae3b544e3511e2.pdf">Described in 1995</a>, there are two species, each known from a single specimen<a href="#1" name="top1"><sup>1</sup></a>. That makes even dwarf pipesnakes (family Anomochilidae), of which <a href="http://snakesarelong.blogspot.com/2012/04/dwarf-pipesnakes.html">we've obtained several new color photos recently,</a> seem relatively well-represented. Putting together this article <a href="http://snakesarelong.blogspot.com/2014/10/how-to-teach-yourself-about-obscure.html">strained my research powers</a>—xenophidiids don't even have an English language Wikipedia page (yet). Google the name of the family and it asks whether you meant Xenophilius, the first name of a minor character from <i>Harry Potter</i>, who has more than twice as many results. <i>Xenophidion</i> means "small strange snakes" in Greek, and indeed we have barely scratched the surface of how strange these snakes probably are. And, to top it all off, no one has ever seen its penis. Read on to find out why.</div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgKQr5n7P1ZcYIbNFNAWiVGtZmpuJpYRarlPRtkNshPMVXts9jU9hUXoEDxXsmMXeQHd191L8HhkFSzpWCli-2z6kq3I7yq6sgUZBmenkjKgF7DDhtbpPSmpGAIKcbSC0OgZQN4ekhd8kjm/s1600/Southeast-Asia-map-outlines.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="288" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgKQr5n7P1ZcYIbNFNAWiVGtZmpuJpYRarlPRtkNshPMVXts9jU9hUXoEDxXsmMXeQHd191L8HhkFSzpWCli-2z6kq3I7yq6sgUZBmenkjKgF7DDhtbpPSmpGAIKcbSC0OgZQN4ekhd8kjm/s320/Southeast-Asia-map-outlines.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Collection locations of the only specimens of<br />
<i>Xenophidion acanthognathus</i> (red)<br />
and <i>Xenophidion schaeferi </i>(green)</td></tr>
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The story of <i>Xenophidion</i> begins on the morning of November 20th, 1987. It was 8:15 AM when Chicago Field Museum Herpetologist <a href="https://www.fieldmuseum.org/about/staff/profile/636">Robert F. Inger</a> found a snake beneath some moss on rock during field work in a selectively-logged forest near Mendolong, in Sabah's Sipitang District on the island of Borneo. Inger, an expert in the herpetology of southeast Asia who <a href="https://www.fieldmuseum.org/science/blog/lifetime-curiosity">by that time in his life had "made thorough searches of thousands of square meters of forest floor litter with the help of very sharp-sighted local men"</a>, had never seen a snake like this before, and he brought it back to Chicago and placed it in the Field Museum collection.<br />
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Almost a year later, at 10:00 PM on November 5th, 1988, German amateur herpetologist Christian Schäfer collected and photographed a snake at the edge of a trail near Templer Park, about 12 miles north of Kuala Lumpur in peninsular Malaysia. Schäfer donated his specimen to the Zoological Museum in Berlin in the spring of 1993. Curators <a href="https://www.naturkundemuseum.berlin/en/einblicke/mitarbeiter/rainer.guenther">Rainer Günther</a> and <a href="https://www.researchgate.net/profile/Ulrich_Manthey">Ulrich Manthey</a> recognized it as unique and asked esteemed herpetologists Van Wallach and Bob Inger to compare it to specimens at Harvard and Chicago. Inger recognized similarity between Schäfer's specimen and his own, and sent both specimens back to Berlin to be described as new species. The dissimilarity between the two new specimens and all other known snakes was so great that <a href="https://www.researchgate.net/profile/Ulrich_Manthey/publication/233570693_Xenophidion_a_new_genus_with_two_new_species_of_snakes_from_Malaysia_Serpentes_Colubridae/links/5602a86308ae3b544e3511e2.pdf">they chose to establish a new genus</a>, which they tentatively placed into the family Colubridae (<a href="http://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">which at the time was much more inclusive</a>). The genus was elevated into a new family after the dissection of the <i>X. acanthognathus </i>specimen <a href="http://booksandjournals.brillonline.com/content/journals/10.1163/156853898x00052">by Wallach and Günther in 1998</a> failed to reveal an obvious affinity with any existing family.<br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgrubtB4QF-nXCZvyDUzsniIAmf4WKY75-5okiSt1sI5qxzxeOtdZRMYvx26qaSd6DIpM1rpXiGh4aOQP4nLu4wP2WfoLm7QAK6gjs-ckBRQUAwWBre0YVso_YolM1agw_DqFT6OmG4aTyL/s1600/Xenophidion+schaeferi+Gunther+%2526+Manthey+1995+Figs+2+and+3.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="376" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgrubtB4QF-nXCZvyDUzsniIAmf4WKY75-5okiSt1sI5qxzxeOtdZRMYvx26qaSd6DIpM1rpXiGh4aOQP4nLu4wP2WfoLm7QAK6gjs-ckBRQUAwWBre0YVso_YolM1agw_DqFT6OmG4aTyL/s400/Xenophidion+schaeferi+Gunther+%2526+Manthey+1995+Figs+2+and+3.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Drawing and photograph of the jaw spine of <i>X. schaeferi </i>(labeled 'Pp')<br />
From <a href="https://www.researchgate.net/profile/Ulrich_Manthey/publication/233570693_Xenophidion_a_new_genus_with_two_new_species_of_snakes_from_Malaysia_Serpentes_Colubridae/links/5602a86308ae3b544e3511e2.pdf">Günther & Manthey 1995</a></td></tr>
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The two specimens share a number of unique features that distinguish them from all other living snakes. Their head scales, especially those along their lips, bear numerous sensory papillae. Their prefrontal scales are much larger than those of other snakes, taking up most of the top of the head in front of the eyes, and the space between their eyes is slightly concave. Their upper jaw bears a long, spiny palatine process, after which <i>X. </i><i>acanthognathus </i>("spine jaw" in Greek)<i> </i>is named. Their small eyes, short tail, and wedge-shaped head all suggest a mostly fossorial lifestyle. Like many <a href="http://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">"henophidian"</a> snakes, <a href="http://snakesarelong.blogspot.com/2012/11/identifying-snake-sheds-part-iii.html">their ventral scales are only slightly wider than their dorsal scales</a>. But, unlike so many henophidians, both species of <i>Xenophidion</i> lack any vestiges of a pelvic girdle, left lung, or coronoid bone, suggesting that they are more closely related to caenophidian snakes. Wallach and Günther noted several similarities among the visceral characteristics of <i>Xenophidion</i> and <a href="http://snakesarelong.blogspot.com/2015/01/dwarf-boas.html">tropidophiids</a>, including a tracheal lung and unlobed kidneys., although we now know that tropidophiids are most closely related to <a href="http://snakesarelong.blogspot.com/2015/02/anilius-pipesnake-that-wasnt.html">aniliids</a>. They also suggested that <i>Xenophidion</i> and another enigmatic snake family, <a href="http://snakesarelong.blogspot.com/2012/04/round-island-splitjaw-snakes.html">bolyeriids</a>, might be related.<br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgXmzcDoKbQe9XgF1SmHY7rtp2TOUT60pPmpaNLMCyIda9fUgCLSRdOZCwEkyyUPOvPhvKDWU88MzFLBFHPz6Rk-zq7h6xQ5Q5wt89mjpoIuQhAWzmYpb1jtA2AcZL-hgWdfYsfCBFhSOer/s1600/Xenophidion+schaeferi+W+Grossman+Fig4+Gunther+%2526+Manthey+1995.jpg" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="285" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgXmzcDoKbQe9XgF1SmHY7rtp2TOUT60pPmpaNLMCyIda9fUgCLSRdOZCwEkyyUPOvPhvKDWU88MzFLBFHPz6Rk-zq7h6xQ5Q5wt89mjpoIuQhAWzmYpb1jtA2AcZL-hgWdfYsfCBFhSOer/s400/Xenophidion+schaeferi+W+Grossman+Fig4+Gunther+%2526+Manthey+1995.jpg" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">The only photograph of a living <i>Xenophidion schaeferi </i>(FMNH 235170),<br />
taken by W. Grossmann. From <a href="https://www.researchgate.net/profile/Ulrich_Manthey/publication/233570693_Xenophidion_a_new_genus_with_two_new_species_of_snakes_from_Malaysia_Serpentes_Colubridae/links/5602a86308ae3b544e3511e2.pdf">Günther & Manthey 1995</a></td><td class="tr-caption"><br /></td><td class="tr-caption"><br /></td></tr>
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In 2004, the sequence of the cytochrome <i>b</i> gene of <i>X. schaeferi</i> was sequenced. <a href="http://www.ncbi.nlm.nih.gov/nuccore/AY574279.1">This is still the only gene</a> we have from either species of <i>Xenophidion</i>, and it has suggested a sister relationship between <i>Xenophidion</i> and <a href="http://snakesarelong.blogspot.com/2012/04/round-island-splitjaw-snakes.html">Bolyeriidae</a> and a distant relationship between <i>Xenophidion</i> and Tropidophiidae in several studies. Evidently, unpublished CT scans of the skull of <i>Xenophidion</i> show that these snakes also have a joint in the maxilla, a characteristic unique to bolyeriids. We know almost nothing about the diet of <i>Xenophidion</i>, but thankfully the stomach of the <i>X. acanthognathus</i> specimen contains a <i>Sphenomorphus</i> skink. Skinks are also eaten by bolyeriids, which use their hinged upper jaws to grasp their hard-bodied, relatively non-deformable prey. It's not inconceivable that <i>Xenophidion </i>might do this as well. The current geographic distribution of Bolyeriidae is limited to Round Island in the Indian Ocean, which <a href="https://www.researchgate.net/profile/Frank_Burbrink/publication/231890539_A_molecular_approach_to_discerning_the_phylogenetic_placement_of_the_enigmatic_snake_Xenophidion_schaeferi_among_the_Alethinophidia/links/5523dcf50cf223eed3800080.pdf">suggests that the common ancestor of these two families</a> was probably ancient and widespread across Gondwanaland.<br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgT-FFG5C_pqBaOa1eNCCtC3jUnxvX_Dp8OWlNWezxLi48bxcttejnnNbRLeWi6ysg_Y0aSS4iPnL_Jh3Y3etZu4Q3VDPGf5thhmPInFVZAzE62B-haC8d9zqxZYzgbl638Le3TeXz0Y3S7/s1600/Xenophidion+acanthognathus+Fig1+Gunther+%2526+Manthey+1995.jpg" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="305" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgT-FFG5C_pqBaOa1eNCCtC3jUnxvX_Dp8OWlNWezxLi48bxcttejnnNbRLeWi6ysg_Y0aSS4iPnL_Jh3Y3etZu4Q3VDPGf5thhmPInFVZAzE62B-haC8d9zqxZYzgbl638Le3TeXz0Y3S7/s400/Xenophidion+acanthognathus+Fig1+Gunther+%2526+Manthey+1995.jpg" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Ventral view of the sole specimen of<br />
<i>Xenophidion acanthognathus </i>(ZMB 50534)<br />
From <a href="https://www.researchgate.net/profile/Ulrich_Manthey/publication/233570693_Xenophidion_a_new_genus_with_two_new_species_of_snakes_from_Malaysia_Serpentes_Colubridae/links/5602a86308ae3b544e3511e2.pdf">Günther & Manthey 1995</a></td></tr>
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There are numerous differences between the two species of <i>Xenophidion</i>. Both have 23 dorsal scale rows at midbody, but the dorsal scales of <i>X. acanthognathus</i> are more heavily keeled than those of <i>X. schaeferi. </i>They have a similar number of ventral scales (181 vs, 178), but <i>X. acanthognathus</i> has 51 subcaudals, 8 more than <i>X. schaeferi</i>. <i>Xenophidion schaeferi</i> also has more teeth on the palatine (10 vs. 8), pterygoid (16 vs. 13), and especially the dentary bone (19 vs. 12) than <i>X. acanthognathus</i>. Finally, <i>X. acanthognathus</i> has a large yellow-white patch on its neck. Because both of the specimens are females, the <a href="http://snakesarelong.blogspot.com/2014/03/why-do-snakes-have-two-penises.html">hemipenes</a>, which contain many taxonomically useful characters, have not been described. But, conveniently, the oviduct of the <i>X. acanthognathus</i> specimen contains two eggs, so at least we know the reproductive mode of these snakes.<br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiIDUqCR8kAPehvC60vRneVaari6I3Mqw3aGNdOzdSf_MJO5qLCMT9T9wGtVD9Por9ArHN_5_PZXrflxBsSsRzicFuESPmHwpDrZdF6k2dZDY-SBog3k18LeuLnOto3FRAVAUxHnER7BO-t/s1600/journal.pone.0161070.g001.PNG" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="400" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiIDUqCR8kAPehvC60vRneVaari6I3Mqw3aGNdOzdSf_MJO5qLCMT9T9wGtVD9Por9ArHN_5_PZXrflxBsSsRzicFuESPmHwpDrZdF6k2dZDY-SBog3k18LeuLnOto3FRAVAUxHnER7BO-t/s400/journal.pone.0161070.g001.PNG" width="307" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Snake family tree from <a href="http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0161070">Figueroa et al. 2016</a>, <span style="font-size: 12.8px;">showing</span><br />
Xenophidiidae + Bolyeriidae as sister to Caenophidia<br />
<a href="http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0161070#">Click</a> for a larger version</td></tr>
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<a href="http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0161070">Some phylogenetic studies</a> suggest that Xenophidiidae and Bolyeriidae might be sister to Caenophidia, leading some to call these two families "proto-colubroids". However, <a href="http://download.springer.com/static/pdf/292/art%253A10.1186%252F1471-2148-13-93.pdf?originUrl=http%3A%2F%2Fbmcevolbiol.biomedcentral.com%2Farticle%2F10.1186%2F1471-2148-13-93&token2=exp=1475028617~acl=%2Fstatic%2Fpdf%2F292%2Fart%25253A10.1186%25252F1471-2148-13-93.pdf*~hmac=d28fba58560302a3bab49227ef5d4f266fd195bc0f4204deb3df26e065f211ff">other genetic analyses</a> group them with boas, pythons, and other "henophidian" snakes instead. Hopefully further gene sequencing will sort this out, and of course fresh <i>Xenophidion</i> specimens wouldn't hurt. The forestry station where Inger collected <i>X. acanthognathus</i> is still operational and <a href="https://www.researchgate.net/publication/305326733_Nocturnal_Terrestrial_Mammals_Population_in_Sabah_Forest_Industries_SFI_Forest_Plantation_Sipitang/figures">researchers continue to work there</a>—I hope they know to keep their eyes open for small, strange snakes. Unfortunately, the primary forest where <i>X. schaeferi</i> was collected was cleared two years later and is now a banana plantation. <a href="http://www.sciencedirect.com/science/article/pii/S0959378000000595">Both peninsular Malaysia and Borneo are losing their forests to timber harvesting and oil palm plantations at an alarming rate</a>. People get upset when they learn that deforestation endangers charismatic species such as <a href="http://www.orangutan.com/threats-to-orangutans/">orangutans</a>, leading to <a href="http://www.rspo.org/">efforts to make palm oil production more sustainable</a>. This is really challenging because palm oil is used in all kinds of delicious things, such as <a href="https://www.girlscouts.org/en/cookies/all-about-cookies/FAQs.html">Girl Scout Cookies</a>, and high-profile controversy over its sustainability has been fueled by people's love for orangutans. I'm here to suggest that the many mysteries of <i>Xenophidion</i>—including what its penis looks like—may never be solved if the rain forests of southeast Asia are lost, and that <i>Xenophidion</i> is at least as valuable and interesting as orangutans.<br />
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<i><u>UPDATE:</u> Additional specimens have since been discovered in <a href="https://www.researchgate.net/profile/Perry_Wood_Jr/publication/326019319_The_rediscovery_of_Schaefer's_Spine-jawed_Snake_Xenophidion_schaeferi_Gunther_Manthey_1995_Serpentes_Xenophidiidae_from_Peninsular_Malaysia_with_notes_on_its_variation_and_the_first_record_of_the_genu/links/5c112cbc92851c39ebe7dd5c/The-rediscovery-of-Schaefers-Spine-jawed-Snake-Xenophidion-schaeferi-Guenther-Manthey-1995-Serpentes-Xenophidiidae-from-Peninsular-Malaysia-with-notes-on-its-variation-and-the-first-record-of-the.pdf">Malaysia, Sumatra</a>, and <a href="https://lkcnhm.nus.edu.sg/app/uploads/2020/01/RBZ-2020-0018.pdf">Borneo</a> including at least one male, bringing the total number of individuals known to six, potentially representing three species if the Sumatra specimen proves to be distinct. Some photos of one of the new specimens <a href="https://www.flickr.com/photos/orionmystery/32570401458/in/photolist-RC8PBQ-PZPRqZ">here</a>.</i></div>
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<a href="https://www.blogger.com/null" name="1"><b>1 </b></a><a href="http://www.iucnredlist.org/details/full/178498/0">The IUCN page for </a><i><a href="http://www.iucnredlist.org/details/full/178498/0">Xenophidion acanthognathus</a> </i>mentions a second specimen from Kinabalu, but I couldn't find any other references to this specimen. Instead, the IUCN references page pointed me, through a couple of intermediates, to a paper (published before the discovery of <i>Xenophidion</i>) that included a reference to the type specimen of <i><a href="http://snakesarelong.blogspot.com/2016/02/dragonsnakes-and-filesnakes-revisited.html">Stoliczkia borneensis</a></i>, which was collected on Mount Kinabalu. Since <i>Stoliczkia borneensis</i> is in the family Xenodermidae, I suspect there may have been some confusion around the somewhat similar family names. <a href="http://portal.vertnet.org/search?q=Xenophidion">VertNet lists only the single Sipitang specimen of <i>X. acanthognathus</i></a>, as does <a href="https://www.crcpress.com/rsc/downloads/JLN44_K21592_SAMPLE.pdf">Wallach et al.'s 2014 edition of <i>Snakes of the World</i></a></span><span style="font-size: x-small;">. Both species of </span><i style="font-size: small;">Xenophidion </i><span style="font-size: x-small;">are listed as Data Deficient by the IUCN.
<a href="#top1"><sup>↩</sup></a></span>
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ACKNOWLEDGMENTS<br />
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Thanks to <a href="http://kokay.hu/eng_index.htm">Szabolcs Kókay</a>, who painted the only color image of <i>Xenophidion</i> for <a href="https://books.google.com/books?id=LRRpBwAAQBAJ&dq=A+field+guide+to+the+reptiles+of+South-East+Asia&lr=&source=gbs_navlinks_s">A Field Guide to the Reptiles of South-east Asia</a>.</div>
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REFERENCES</div>
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<span style="font-size: x-small;">Chan-ard, T., Grossmann, W., Gumprecht, A. & Schulz, K.D. 1999. </span><span style="font-size: x-small; text-indent: -0.5in;">Amphibians and reptiles of Peninsular Malaysia and Thailand: an illustrated checklist. </span><span style="font-size: x-small; text-indent: -0.5in;">Bushmaster Publishing, Wuerselen, 240 pp. <<a href="http://www.nhbs.com/title/94675/amphibians-and-reptiles-of-peninsular-malaysia-and-thailand-an-illustrated-checklist-amphibien-und-reptilien-der-halbinsel-malaysia-und-thailands-eine-illustrierte-checkliste">link</a>></span></div>
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<span style="font-size: x-small;"><br /></span></div>
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<span style="font-size: x-small; text-indent: -0.5in;">Das, I. 2010. A field guide to the reptiles of South-East Asia. New </span><span style="font-size: x-small; text-indent: -0.5in;">Holland Publishers, London, 376 pp. <<a href="https://books.google.com/books?id=LRRpBwAAQBAJ&dq=A+field+guide+to+the+reptiles+of+South-East+Asia&lr=&source=gbs_navlinks_s">link</a>></span></div>
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<span style="font-size: x-small; text-indent: -0.5in;">Das, I. 2012. A naturalist’s guide to the snakes of South-East Asia</span><span style="font-size: x-small; text-indent: -0.5in;">. John Beaufoy Publishing, Oxford, 176 pp. <<a href="http://ir.unimas.my/422/1/snakes_of_south_east_Asia.pdf">excerpt</a>/<a href="http://johnbeaufoy.com/a-naturalists-guide-to-the-%E2%80%A8snakes-of-south-east-asia%E2%80%A8/">link</a>></span></div>
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<span style="font-size: x-small; text-indent: -0.5in;">Figueroa, A., A. D. McKelvy, L. L. Grismer, C. D. Bell, and S. P. Lailvaux. 2016. A species-level phylogeny of extant snakes with description of a new colubrid subfamily and genus. PLoS ONE 11:e0161070 <</span><a href="http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0161070" style="font-size: small; text-indent: -0.5in;">link</a><span style="font-size: x-small; text-indent: -0.5in;">></span></div>
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<div style="text-align: justify;">
<span style="font-size: x-small; text-indent: -0.5in;">Günther, R. & U. Manthey. 1995. </span><i style="font-size: x-small; text-indent: -0.5in;">Xenophidion</i><span style="font-size: x-small; text-indent: -0.5in;">, a new genus with two new
species of snakes from Malaysia (Serpentes, Colubridae). Amphibia-Reptilia</span><b style="font-size: small; font-style: italic; text-indent: -0.5in;"> </b><span style="font-size: x-small; text-indent: -0.5in;">16:229-240 <</span><a href="https://www.researchgate.net/profile/Ulrich_Manthey/publication/233570693_Xenophidion_a_new_genus_with_two_new_species_of_snakes_from_Malaysia_Serpentes_Colubridae/links/5602a86308ae3b544e3511e2.pdf" style="font-size: small; text-indent: -0.5in;">link</a><span style="font-size: x-small; text-indent: -0.5in;">></span></div>
<br />
<div style="text-align: justify;">
<span style="font-size: x-small;">Lawson, R., J. B. Slowinski & F. T. Burbrink. 2004. A molecular approach to discerning the phylogenetic placement of the
enigmatic snake <i>Xenophidion schaeferi</i>
among the Alethinophidia. Journal of
Zoology<i> </i>263:285-294 <<a href="https://www.researchgate.net/profile/Frank_Burbrink/publication/231890539_A_molecular_approach_to_discerning_the_phylogenetic_placement_of_the_enigmatic_snake_Xenophidion_schaeferi_among_the_Alethinophidia/links/5523dcf50cf223eed3800080.pdf">link</a>></span></div>
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<div style="text-align: justify;">
<span style="font-size: x-small;">Pyron, R. A., F. Burbrink, and J. J. Wiens. 2013. A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology 13:93 <<a href="http://bmcevolbiol.biomedcentral.com/articles/10.1186/1471-2148-13-93">link</a>></span></div>
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<span style="font-size: x-small;">Wallach, V. & R. Günther. 1998. Visceral anatomy
of the Malaysian snake genus <i>Xenophidion</i>,
including a cladistic analysis and allocation to a new family (Serpentes:
Xenophidiidae). Amphibia-Reptilia<i> </i>19:385-405 <<a href="http://booksandjournals.brillonline.com/content/journals/10.1163/156853898x00052">link</a>><o:p></o:p></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Wallach, V. W., Kenneth J. and J. Boundy. 2014. Snakes of the World: A Catalogue of Living and Extinct Species. CRC Press, Boca Raton, Florida, USA <<a href="https://books.google.com/books?id=zZ3NBQAAQBAJ&dq=Wallach,+V.+W.,+Kenneth+J.+and+J.+Boundy.+2014.+Snakes+of+the+World:+A+Catalogue+of+Living+and+Extinct+Species.+CRC+Press,+Boca+Raton,+Florida,+USA.&lr=&source=gbs_navlinks_s">link</a>/<a href="https://www.crcpress.com/rsc/downloads/JLN44_K21592_SAMPLE.pdf">sample</a>></span><br />
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<a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license"><span style="font-size: x-small;"><img alt="Creative Commons License" src="https://i.creativecommons.org/l/by-nc-nd/3.0/80x15.png" style="border-width: 0px;"></span></a></div>
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="https://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
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Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com8Raleigh, NC, USA35.7795897 -78.63817870000002635.3677027 -79.28362570000003 36.1914767 -77.992731700000022tag:blogger.com,1999:blog-7443075087825368900.post-7788706182424390122016-08-17T09:32:00.002-06:002017-04-19T00:22:09.480-06:00Why snakes are long (and other recent updates)<div style="text-align: justify;">
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<span style="font-size: x-small;">This post will soon be available in Spanish</span> </div>
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjYF1xyakn6YhL1mg96v6WNUggBEyRJHWLUL1otNvJoQjfjcvFL8nwAP1xYt8lBUV-PixdpD02XwNKuXV4PBlafberpE0-pOqK-KoGVFGEgrTDntLiCGy8PNZMdm8GsxORCTltNmJqjEfCI/s1600/iverson_hognest.jpg" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="195" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjYF1xyakn6YhL1mg96v6WNUggBEyRJHWLUL1otNvJoQjfjcvFL8nwAP1xYt8lBUV-PixdpD02XwNKuXV4PBlafberpE0-pOqK-KoGVFGEgrTDntLiCGy8PNZMdm8GsxORCTltNmJqjEfCI/s320/iverson_hognest.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">A Western Hog-nosed Snake (<i>Heterodon nasicus</i>)<br />
depredating a turtle nest in Nebraska</td></tr>
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As I did in <a href="http://snakesarelong.blogspot.com/2016/03/state-snakes-linnaean-names-and-other.html">March</a> and <a href="http://snakesarelong.blogspot.com/2016/06/virgin-birth-color-of-fossil-snakes-and.html">June</a>, I wanted to highlight some recent and exciting updates to some of my earlier articles. There is <i>so</i> much recent snake news this month, which is lucky for me because I've been writing my dissertation, which I'll be defending next month, so I haven't had much time to write. anything else! I hope to return to longer-form content in October.<br />
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Also, although I rarely promote my own research on this blog, I'm very excited to have just published the second chapter of my masters thesis, <a href="https://www.researchgate.net/publication/305653224_Ontogenetic_shifts_in_the_diet_of_plains_hog-nosed_snakes_Heterodon_nasicus_revealed_by_stable_isotope_analysis">documenting the diet of Western Hog-nosed Snakes (<i>Heterodon nasicus</i>) in Illinois</a> using stable isotopes to quadruple our sample size. We showed that the diet of juveniles was composed mostly of Six-lined Racerunners (<i>Aspidoscelis sexlineata</i>) and their eggs, whereas adults mostly feed on aquatic turtle eggs! Surprisingly, we found very little evidence that these snakes were eating amphibians, which are considered to be staples of their diet elsewhere.</div>
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<b><a href="http://snakesarelong.blogspot.com/2014/12/big-deal-snake-genomes.html">Snake Genomes</a>, <a href="http://snakesarelong.blogspot.com/2012/04/lizards-of-glass.html">Lizards of Glass</a>, and Why Snakes are Long</b><br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgsyzdInAUDFaZm7Y6nl7bwCT43yBAomtiLpW6LevOanUGa2SfgtDdEOR0HSc0wOhjRlTVJ0MNl5IKENMNCp-AXPEA9nreFhBW-kosIfcSr-CaJDQaYWSjdqwZu1KFYD4clWZtrdZJGCegj/s1600/Head+%2526+Polly+2015+Fig3.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="260" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgsyzdInAUDFaZm7Y6nl7bwCT43yBAomtiLpW6LevOanUGa2SfgtDdEOR0HSc0wOhjRlTVJ0MNl5IKENMNCp-AXPEA9nreFhBW-kosIfcSr-CaJDQaYWSjdqwZu1KFYD4clWZtrdZJGCegj/s400/Head+%2526+Polly+2015+Fig3.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Variation in the length of different vertebrate bodies,<br />
including a Rosy Boa (<i>Lichanura</i>)<br />
From <a href="http://s3.amazonaws.com/academia.edu.documents/40996608/Head_and_Polly__2015__Homoplasy_in_HOX.pdf?AWSAccessKeyId=AKIAJ56TQJRTWSMTNPEA&Expires=1471406746&Signature=K%2F%2FFovUe6TLHoURBlCiHeNfPYU4%3D&response-content-disposition=inline%3B%20filename%3DEvolution_of_the_snake_body_form_reveals.pdf">Head & Polly 2015</a></td></tr>
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Researchers from the <a href="http://www.igc.gulbenkian.pt/">Gulbenkian Institute of Science</a> in Portugal used data from <a href="http://snakesarelong.blogspot.com/2014/12/big-deal-snake-genomes.html">the king cobra and Burmese python genomes</a> as part of new research that's too meta for me not to write about. The developmental biology of snakes deserves a whole article (and, indeed, is <a href="http://www.cambridge.org/er/academic/subjects/life-sciences/evolutionary-biology/how-snake-lost-its-legs-curious-tales-frontier-evo-devo?format=PB">the subject of an entire book chapter</a>), but snake development is uniquely interesting from the perspective of understanding the evolution of both limblessness and body elongation. A very recent <a href="http://www.cell.com/developmental-cell/pdf/S1534-5807(16)30424-5.pdf">article in the journal <i>Developmental Cell</i></a> shed light on the genetic regulation of body length in vertebrates, which varies from a dozen or fewer in humans and most vertebrates to over 200 in many snakes and <a href="https://books.google.com/books?id=CzxVvKmrtIgC&pg=PA329&lpg=PA329&dq=how+many+vertebrae+do+caecilians+have&source=bl&ots=AYN2g7swkZ&sig=gvg1_XILaFRnpeNFm2IoQASCXp4&hl=en&sa=X&ved=0ahUKEwix_-PN4r_OAhVFZCYKHV4zBc0Q6AEISTAG#v=onepage&q=how%20many%20vertebrae%20do%20caecilians%20have&f=false">caecilians</a> and exceeds 400 in <a href="http://rsbl.royalsocietypublishing.org/content/3/3/296.short">some snake species</a>.<br />
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For a long time, scientists thought that <a href="http://www.nature.com/scitable/topicpage/hox-genes-in-development-the-hox-code-41402">Hox genes</a>, which control many aspects of body layout and development, probably controlled body length too. But, so far experiments modifying Hox genes have failed to produce differences in body length, and <a href="http://s3.amazonaws.com/academia.edu.documents/40996608/Head_and_Polly__2015__Homoplasy_in_HOX.pdf?AWSAccessKeyId=AKIAJ56TQJRTWSMTNPEA&Expires=1471449264&Signature=%2Fe%2FNNOxv6u09CK80PlN2aQjHeGg%3D&response-content-disposition=inline%3B%20filename%3DEvolution_of_the_snake_body_form_reveals.pdf">most snake Hox genes are not substantially different from those of other vertebrates</a>. Instead, the new study showed that the "junk DNA" surrounding a different gene, called Oct4, apparently influences body length in developing vertebrate embryos. Although the Oct4 gene itself was already known to play a role in regulating stem cell flexibility, the surrounding DNA <a href="http://www.sciencemag.org/news/2016/08/junk-dna-tells-mice-and-snakes-how-grow-backbone">was formerly considered to be "junk DNA" because it was not translated into RNA and seemed to have no purpose</a>. Measurements showed that Oct4 is active for longer in developing snake embryos than in mouse embryos, which is probably what causes their bodies to grow so long. And, just by copying snake "junk DNA" into mouse embryos, the researchers were able to artificially increase both the level of expression of Oct4 in mouse embryos as well as the length of their bodies. Comparing the genomes of <a href="http://snakesarelong.blogspot.com/2014/12/big-deal-snake-genomes.html">snakes</a>, lizards, and mammals showed that the Oct4 "junk DNA" of snakes differed from that of lizards and mammals. Interestingly, <a href="http://gooa.las.ac.cn/external/download/1172974/4175311/20150422040446140.pdf">glass lizard (<i>Ophisaurus</i>)</a> Oct4 "junk DNA" was similar to that of geckos and anoles, even though <a href="http://snakesarelong.blogspot.com/2012/04/lizards-of-glass.html">these limbless lizards share an elongated snake-like body form with snakes</a>.</div>
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<b><a href="http://snakesarelong.blogspot.com/2012/09/recent-conservation-successes-with.html">Conservation Successes with Indigo Snakes</a></b><br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgogT0Ics6HISSH07gRkmahVRegWDeAeGbuxRYB8BZdppJ_EJDTbbQVSyLCDPtT58Cl3JYNZvVpn091IA0Ee-3VdLAy77TkOOgjs2Z_bYQxZXHQ_aRSiTqKMVExiwx359NF3HaTWMPFA9W5/s1600/Bauder+et+al+2016+Fig1.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="302" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgogT0Ics6HISSH07gRkmahVRegWDeAeGbuxRYB8BZdppJ_EJDTbbQVSyLCDPtT58Cl3JYNZvVpn091IA0Ee-3VdLAy77TkOOgjs2Z_bYQxZXHQ_aRSiTqKMVExiwx359NF3HaTWMPFA9W5/s320/Bauder+et+al+2016+Fig1.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Seasonal variation in the probability of moving of female (top)<br />
& male (bottom) Eastern Indigo Snakes (<i>Drymarchon couperi</i>)<br />
in Florida. From <a href="http://www.hljournals.org/doi/abs/10.1655/Herpetologica-D-15-00039.1">Bauder et al. 2016</a></td></tr>
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Many snakes make seasonal movements to and from hibernacula, in search of food or mates, or for other reasons. Often, we think of these movements as driven by the changing of seasons, either <a href="http://www.jstor.org/stable/2265663?seq=1#page_scan_tab_contents">the wet and dry seasons</a> in the tropics or <a href="http://carlgans.org/bor-view/?borv=13&borpage=53">the four well-defined temperate seasons</a>. But, we don't know much about the seasonal movements of snakes that live in tropical and sub-tropical zones. For a species with such a small range, Eastern Indigo Snakes have fairly different requirements <a href="http://onlinelibrary.wiley.com/doi/10.1002/jwmg.645/full">in the northern part of their range</a>, where frosty Georgia winter nights force them to rely on deep, warm Gopher Tortoise (<i>Gopherus polyphemus</i>) burrows, and in sub-tropical peninsular Florida, where they are less reliant on such particular shelters. A recent study by a group of Florida scientists, including <a href="http://www.oriannesociety.org/">Orianne Society</a> staff, used radio-telemetry to document a seasonal pattern of movement in Florida Indigo Snakes that differs from their pattern in Georgia. In particular, male Florida Indigos are most likely to move in the late fall and early winter, when they are searching for mates, whereas both males and females stay put during the spring, for reasons yet unknown. In contrast, Indigos in the rest of their range maintain small winter home ranges on xeric sandhills but use much larger home ranges and a greater diversity of habitats during the rest of the year.<br />
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<b><a href="http://snakesarelong.blogspot.com/2013/02/malagasy-leaf-nosed-snakes.html">Malagasy Leaf-nosed Snakes</a></b><br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgBKoBVQHzWDZhvRWA5H9hxB4lMFfIlxYOqxJDaoWWd9NzpubB9T4wPdjTV7rVlRFhP-RTJX8U1HkhjwMq9OHWbFqXmtA2hGBOPqW2FzoDoIvrJegdWmQixeWCIRfgBSLAHPi8PC3Dg6ZFu/s1600/Langaha+pseudoalluaudi+iNat+2016.jpg" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="400" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgBKoBVQHzWDZhvRWA5H9hxB4lMFfIlxYOqxJDaoWWd9NzpubB9T4wPdjTV7rVlRFhP-RTJX8U1HkhjwMq9OHWbFqXmtA2hGBOPqW2FzoDoIvrJegdWmQixeWCIRfgBSLAHPi8PC3Dg6ZFu/s400/Langaha+pseudoalluaudi+iNat+2016.jpg" width="300" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">A female <i>Langaha pseudoalluaudi</i><br />
Photo from <a href="http://www.inaturalist.org/blog/6935-observation-of-the-week-8-10-16">iNaturalist</a></td></tr>
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Global all-taxa citizen science portal <a href="http://www.inaturalist.org/blog/6935-observation-of-the-week-8-10-16">iNaturalist's observation of the week</a> this week was a very rare snake indeed, a photograph of <i>Langaha pseudoalluaudi. </i>Less than two dozen other individuals of this species have ever been found by scientists. The first was collected in 1966 and described in 1988, and <a href="file:///C:/Users/Andrew/Documents/Articles/Kuchling-3-4-03.pdf">the second individual was </a><a href="http://www.salamandra-journal.com/index.php%3Foption%3Dcom_docman%26task%3Ddoc_download%26gid%3D215%26Itemid%3D71">photographed in 2003 by a Durrell Wildlife Conservation Trust biologist</a>. Since that time, a handful of other records have trickled in from the field in Madagascar, including <a href="https://www.flickr.com/photos/64231410@N00/7800989452/in/photolist-cTm7o9-e5UnEm-e5UnKL">a few photos on Flickr</a> and the only known photo of a male, published in a field guide in 2007. This individual was found by group of <a href="http://opwall.com/">Operation Wallacea</a> volunteers on a conservation research expedition, one of whom, Victoria Jackson, a student of Biological Sciences at the University of Exeter, posted it on iNaturalist. Of the three species of <i>Langaha</i>, none of which are particularly well-known, <i>L. pseudoalluaudi </i>is by far the rarest and most poorly-known. Perhaps the most fascinating aspect of <i>Langaha</i> biology is their obvious anatomical sexual dimorphism, a feature that is very rare among snakes. Females of all three <i>Langaha </i>species have a serrated snout that resembles a small flower that has not fully bloomed.<i> </i>Female <i>L. pseudoalluaudi</i> also have protruding horn-like scales above their eyes. Males have smooth, pointed snouts instead that <a href="http://snakesarelong.blogspot.com/2013/02/malagasy-leaf-nosed-snakes.html">resemble the seed pods of a Madagascan legume</a>. We have very little idea why these snakes might be sexually dimorphic—the nose ornaments could be shaped by sexual selection, or they might function to make the snakes more cryptic to predators or prey, if the sexes forage or hide in different environments or on different foods.</div>
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<b><a href="http://snakesarelong.blogspot.com/2015/08/do-snakes-sleep.html">Do Snakes Sleep?</a></b><br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi0oQBawHx8DRQPZ62Lt277Gur_0PNas4wwZ0w79dVxfWJJ5HLAjO6ZiC7vJPHfAkbrYI6h_W1wh0im-TgLRtykVRzuFaL27G-Vkgh6rreyUdhrLaogfNdqGC87HEqvNJ2O385gsd1GITo6/s1600/Shein-Idelson+et+al+2016+Fig1B.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi0oQBawHx8DRQPZ62Lt277Gur_0PNas4wwZ0w79dVxfWJJ5HLAjO6ZiC7vJPHfAkbrYI6h_W1wh0im-TgLRtykVRzuFaL27G-Vkgh6rreyUdhrLaogfNdqGC87HEqvNJ2O385gsd1GITo6/s320/Shein-Idelson+et+al+2016+Fig1B.png" width="289" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Sleep in Bearded Dragons<br />
From <a href="http://science.sciencemag.org/content/352/6285/590">Shine-Idelson et al. 2016</a></td></tr>
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There has only ever been one study of sleep in snakes. It was conducted in France in 1969 on an African Rock Python (<i>Python sebae</i>), which produced sleep-like brain waves almost 16 hours a day, increasing to over 20 hours following feeding. The data suggest that these brainwaves corresponded with slower breathing and heart rate, some muscle relaxation, and perhaps a lowered behavioral response threshold. There was no evidence for REM sleep in this snake. Evidence for REM sleep in other reptiles is mixed. <a href="http://science.sciencemag.org/content/352/6285/590">The April 29th issue of <i>Science</i></a> contained new data from the Max Planck Institute for Brain Research documenting slow-wave and rapid eye movement (REM) sleep in Bearded Dragons (<i>Pogona vitticeps</i>). <i>[Edit 4/19/2017: This research <a href="https://www.researchgate.net/profile/Niels_Rattenborg/publication/303770579_Sleep_Ecophysiology_Integrating_Neuroscience_and_Ecology/links/57555b9a08aec74acf57c743.pdf">has since been criticized</a> for failing to exclude the possibility that putative REM sleep was actually partial or full awakenings from sleep, which looks very similar.] </i>This is pretty cool because Bearded Dragons and snakes might be pretty close relatives (<a href="http://mfile.narotama.ac.id/files/Jurnal/Jurnal%202012-2013/Who%20speaks%20with%20a%20forked%20tongue.pdf">if phylogenetic trees using molecular data are to be believed</a>) and it suggests that not only do reptiles definitely sleep, they may also dream. Previously, scientists had hypothesized that slow-wave and REM sleep evolved independently in birds and mammals and, <a href="http://snakesarelong.blogspot.com/2015/11/snakes-that-are-good-parents.html">like parental care</a>, could be linked to endothermy. The unequivocal <i>[see above edit]</i> evidence for these sleep phases in reptiles suggest that REM sleep evolved much earlier and probably only once. The senior author on the study, neuroscientist Gilles Laurent <a href="http://www.scientificamerican.com/article/slumber-party-reptiles-like-us-have-rem-sleep-and-may-dream/">was quoted as saying</a> "If you forced me to speculate and to use a loose definition of dreaming, I'd speculate that [Bearded Dragon] dreams are about recent notable events: insects, maybe a place where there are good insects, an aggressive male in the next terrarium, et cetera. If I were an Australian dragon living in Frankfurt, I'd be dreaming of a warm day in the sun."</div>
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<b><a href="http://snakesarelong.blogspot.com/2014/11/the-9999th-reptile.html">The 9,999th Reptile</a></b><br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEggP-nDApblidCfJS2sR987WM7jnRxdhHybTCPJJkoVktKQj2XG9CRKGl5NFw17NJlpDQJ6i7W_PNZes9_poUxbgUU8lodWXnn_91VnAcW2L0Fl9uwIG2X-GjPIcai2eXgJxk1eGZ93VuUd/s1600/Geophis+lorcana.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="217" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEggP-nDApblidCfJS2sR987WM7jnRxdhHybTCPJJkoVktKQj2XG9CRKGl5NFw17NJlpDQJ6i7W_PNZes9_poUxbgUU8lodWXnn_91VnAcW2L0Fl9uwIG2X-GjPIcai2eXgJxk1eGZ93VuUd/s320/Geophis+lorcana.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Geophis lorcana</i><br />
From <a href="http://zookeys.pensoft.net/articles.php?id=8605&display_type=list&element_type=9">Canseco-Márquez et al. 2016</a></td></tr>
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<a href="http://www.reptile-database.org/db-info/news.html">Snake species number over 3,600 this month</a>, in part because of <a href="http://zookeys.pensoft.net/articles.php?id=8605&display_type=list&element_type=9">the description of a new species of <i>Geophis</i> from Mexico</a>. The beautiful <i>Geophis lorcana</i> is the 50th species in the genus <i>Geophis</i> and the 8th new species in that genus since the turn of the century. It was discovered in the cloud forests of the Sierra Zongolica and Sierra de Quimixtlán mountains by biologist Miguel Ángel de la Torre Loranca, in whose honor the new species is named. Like other <i>Geophis</i>, this snake is fossorial and secretive, and has a small geographic range. Further exploration of this region combined with molecular and anatomical data is likely to yield additional new species, although the habitats in which they are likely to be found are vulnerable to a variety of threats. Other new snake species from the past year include <a href="http://www.mesoamericanherpetology.com/uploads/3/4/7/9/34798824/wallach-epictia_paper.pdf">10 new blindsnakes in the genus <i>Epictia </i>from Central and South America</a>, <a href="https://www.researchgate.net/profile/Robert_Reynolds11/publication/303504961_Discovery_of_a_Remarkable_New_Boa_from_the_Conception_Island_Bank_Bahamas/links/57588fdc08ae9a9c954a7aa1.pdf">a new boa from the Bahamas</a> that sheds light on the island biogeography of the Caribbean, and <a href="http://www.hljournals.org/doi/abs/10.1655/HERPETOLOGICA-D-15-00059">a new species of Neotropical watersnake</a> (genus <i>Helicops</i>) <a href="http://reptile-database.reptarium.cz/species?genus=Helicops&species=nentur&search_param=%28%28genus%3D%27Helicops%27%29%29">whose specific epithet is taken from one of Tolkien's Elvish languages</a>.</div>
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<b><a href="http://snakesarelong.blogspot.com/2012/04/identifying-snake-sheds.html">Identifying Snake Sheds</a></b><br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg2aiLwaKNu_s-oeLi-Rs9xlr-drbcZdLwFYQmD0dzUXYNyCLtJHX02VxE0ZNPMC2PNRivz7_nU6ZKTlPmNb29sUnMdSWZLS8hyphenhyphene11HDB0Oe6oLXbnUhGXP6AGwJykNwYB1sxCCnYaao6FK/s1600/Antaresia+stimsoni+in+shed+ASRC+video.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="214" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg2aiLwaKNu_s-oeLi-Rs9xlr-drbcZdLwFYQmD0dzUXYNyCLtJHX02VxE0ZNPMC2PNRivz7_nU6ZKTlPmNb29sUnMdSWZLS8hyphenhyphene11HDB0Oe6oLXbnUhGXP6AGwJykNwYB1sxCCnYaao6FK/s320/Antaresia+stimsoni+in+shed+ASRC+video.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Antaresia stimsoni</i> inside its shed skin<br />
Video still from <a href="http://www.reptilecentre.com.au/">Alice Springs Reptile Centre</a></td></tr>
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It's not peer-reviewed research, but a recent video from the <a href="http://www.reptilecentre.com.au/">Alice Springs Reptile Centre</a> in Alice Springs, Australia showed an unusual occurrence—a shedding Stimson's Python (<i>Antaresia stimsoni</i>) that seemed to have gotten stuck inside of an endless loop of its own shed skin. The snake must have crawled into the mouth orifice of the shed skin before it finished shedding the skin from the posterior part of its body. According to <a href="https://www.facebook.com/alicespringsreptilecentre/videos/1225923104085501/">a Facebook post</a>, the Alice Springs Reptile Centre staff reported that they had not observed this phenomenon before and that the python was able to free itself after about three hours of crawling in a circle by making a small, tidy exit hole in the shed. The video was featured on the popular <a href="http://www.iflscience.com/plants-and-animals/snake-gets-trapped-in-an-endless-loop-of-its-own-skin-after-shedding/">IFLS</a> science fan site.</div>
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ACKNOWLEDGMENTS<br />
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Thanks to John Iverson for the use of his photo.</div>
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REFERENCES</div>
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhWgPys_7-lx6PmkO30rg2XbDM5hDgTPZ2R8FIznWQV4WdHaAZa3D4h6UNgKrKZBHYGfMEAQMuaPsFBnuJIO7DokzQZjyy3BNUNzmK8qCEUj-U9HwnqNdkEPGECYKHLUtGdKRc8ysTSbVw_/s1600/Langaha+pseudoalluaudi+Domergue+1988.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="400" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhWgPys_7-lx6PmkO30rg2XbDM5hDgTPZ2R8FIznWQV4WdHaAZa3D4h6UNgKrKZBHYGfMEAQMuaPsFBnuJIO7DokzQZjyy3BNUNzmK8qCEUj-U9HwnqNdkEPGECYKHLUtGdKRc8ysTSbVw_/s400/Langaha+pseudoalluaudi+Domergue+1988.jpg" width="280" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Female <i>Langaha </i><i style="font-size: small; text-align: justify;">pseudoalluaudi</i><span style="font-size: x-small; text-align: justify;"><br />From its original description<br />in <a href="http://bionames.org/references/5e90b8ee26199f42be24bb9aad4d49b0">Domergue 1988</a></span></td></tr>
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<span style="font-size: x-small;">Aires, R., Jurberg, Arnon D., Leal, F., Nóvoa, A., Cohn, Martin J. & Mallo, M. (2016) Oct4 is a key regulator of vertebrate trunk length diversity. Developmental Cell, 38, 262-274 <<a href="http://www.cell.com/developmental-cell/pdf/S1534-5807(16)30424-5.pdf">link</a>></span><br />
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<span style="font-size: x-small;">Aulsebrook, A. E., T. M. Jones, N. C. Rattenborg, T. C. Roth, II, and J. A. Lesku. Sleep Ecophysiology: Integrating Neuroscience and Ecology. Trends in Ecology & Evolution 31:590-599 <<a href="https://www.researchgate.net/profile/Niels_Rattenborg/publication/303770579_Sleep_Ecophysiology_Integrating_Neuroscience_and_Ecology/links/57555b9a08aec74acf57c743.pdf">link</a>></span></div>
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<span style="font-size: x-small;">Bauder, J.M., Breininger, D.R., Bolt, M.R., Legare, M.L., Jenkins, C.L., Rothermel, B.B. & McGarigal, K. (2016) Seasonal variation in Eastern Indigo Snake (<i>Drymarchon couperi</i>) movement patterns and space use in peninsular Florida at multiple temporal scales. Herpetologica, 72, 214-226 <<a href="http://www.hljournals.org/doi/abs/10.1655/Herpetologica-D-15-00039.1">link</a>></span><br />
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<span style="font-size: x-small;">Canseco-Márquez, L., C. J. Pavón-Vázquez, M. A. Lòpez-Luna, and A. Nieto-Montes de Oca. 2016. A new species of earth snake (Dipsadidae, <i>Geophis</i>) from Mexico. ZooKeys 610:131-145 <<a href="http://zookeys.pensoft.net/articles.php?id=8605&display_type=list&element_type=9">link</a>></span><br />
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<span style="font-size: x-small;">Costa, H. C., D. J. Santana, F. Leal, R. Koroiva, and P. C. A. Garcia. 2016. A New Species of <i>Helicops</i> (Serpentes: Dipsadidae: Hydropsini) from Southeastern Brazil. Herpetologica 72:157-166 <<a href="http://www.hljournals.org/doi/abs/10.1655/HERPETOLOGICA-D-15-00059">link</a>></span><br />
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<span style="font-size: x-small;">Domergue, C. A. 1988. Notes sur les serpents de la région malgache. VIII: Colubridae nouveaux. Bulletin du Muséum national d'histoire naturelle. Section A, Zoologie, biologie et écologie animales 10:135-146 <<a href="http://bionames.org/references/5e90b8ee26199f42be24bb9aad4d49b0">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Durso, A. M. and S. J. Mullin. 2016. Ontogenetic shifts in the diets of Plains Hog-nosed Snakes (Colubridae: <i>Heterodon</i>) revealed by stable isotope analysis. Zoology DOI:10.1016/j.zool.2016.07.004 <<a href="https://www.researchgate.net/publication/305653224_Ontogenetic_shifts_in_the_diet_of_plains_hog-nosed_snakes_Heterodon_nasicus_revealed_by_stable_isotope_analysis">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Head, J. J. and P. D. Polly. 2015. Evolution of the snake body form reveals homoplasy in amniote Hox gene function. Nature 520:86-89 <<a href="http://s3.amazonaws.com/academia.edu.documents/40996608/Head_and_Polly__2015__Homoplasy_in_HOX.pdf?AWSAccessKeyId=AKIAJ56TQJRTWSMTNPEA&Expires=1471406746&Signature=K%2F%2FFovUe6TLHoURBlCiHeNfPYU4%3D&response-content-disposition=inline%3B%20filename%3DEvolution_of_the_snake_body_form_reveals.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Held Jr., L. I. 2014. The snake. Pages 75-94 in L. I. Held Jr., editor. How the Snake Lost its Legs. Cambridge University Press, Cambridge <<a href="http://www.cambridge.org/er/academic/subjects/life-sciences/evolutionary-biology/how-snake-lost-its-legs-curious-tales-frontier-evo-devo?format=PB">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Kuchling, G. (2003) New record, range extension, and colouration in life of <i>Langaha pseudoalluaudi</i> (Reptilia: Colubridae) in north-western Madagascar. Salamandra, 39, 235-240 <<a href="http://www.salamandra-journal.com/index.php%3Foption%3Dcom_docman%26task%3Ddoc_download%26gid%3D215%26Itemid%3D71">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Reynolds, R. G., A. R. Puente-Rolón, A. J. Geneva, K. J. Aviles-Rodriguez, and N. C. Herrmann. 2016. Discovery of a Remarkable New Boa from the Conception Island Bank, Bahamas. Breviora 549:1-19 <<a href="https://www.researchgate.net/profile/Robert_Reynolds11/publication/303504961_Discovery_of_a_Remarkable_New_Boa_from_the_Conception_Island_Bank_Bahamas/links/57588fdc08ae9a9c954a7aa1.pdf">link</a>></span></div>
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<span style="font-size: x-small;">Shein-Idelson, M., Ondracek, J.M., Liaw, H.-P., Reiter, S. & Laurent, G. (2016) Slow waves, sharp waves, ripples, and REM in sleeping dragons. Science, 352, 590-595 <<a href="http://science.sciencemag.org/content/352/6285/590">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Wallach, V. 2016. Morphological review and taxonomic status of the <i>Epictia phenops</i> species group of Mesoamerica, with description of six new species and discussion of South American <i>Epictia albifrons</i>, <i>E. goudotii</i>, and <i>E. tenella </i>(Serpentes: Leptotyphlopidae: Epictinae). Mesoamerican Herpetology 3:216-374 <<a href="http://www.mesoamericanherpetology.com/uploads/3/4/7/9/34798824/wallach-epictia_paper.pdf">link</a>></span></div>
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="https://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com0Raleigh, NC, USA35.7795897 -78.63817870000002635.3677027 -79.28362570000003 36.1914767 -77.992731700000022tag:blogger.com,1999:blog-7443075087825368900.post-47182696615693991162016-07-01T07:00:00.000-06:002017-03-20T10:36:22.778-06:00What the Provincial Snakes of Canada Should Be<div class="separator" style="clear: both; text-align: center;">
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<span style="font-size: x-small;">This post will soon be available in Spanish!</span></div>
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<tr><td style="text-align: center;"><a href="https://upload.wikimedia.org/wikipedia/commons/thumb/1/14/Political_map_of_Canada.png/650px-Political_map_of_Canada.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto; text-align: justify;"><img border="0" height="345" src="https://upload.wikimedia.org/wikipedia/commons/thumb/1/14/Political_map_of_Canada.png/650px-Political_map_of_Canada.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">In case, <a href="http://www.buzzfeed.com/tanyachen/americans-fail-canada-again#.dad9j5wvA">like many Americans</a>, you need a map</td></tr>
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Happy Canada Day! And indeed there is a lot to celebrate, in particular <a href="http://www.sciencemag.org/news/2016/03/canadian-scientists-smile-liberals-deliver-d-j-vu-budget">Canada's new liberal government and the positive effects it has had on science and the environment</a>. Three summers ago, I wrote in two parts (<a href="http://snakesarelong.blogspot.com/2013/06/what-state-snakes-should-be-part-i.html">I</a> and <a href="http://snakesarelong.blogspot.com/2013/08/what-state-snakes-should-be-part-ii.html">II</a>) about what the symbolic snakes of each of the US states should be, inspired by the witty and spot-on post '<a href="http://www.thebirdist.com/2013/04/state-birds-what-they-should-be.html">The State Birds: What They SHOULD Be</a>' from <a href="http://www.thebirdist.com/">thebirdist.com</a>. In response to a tweet from <a href="https://twitter.com/CanFieldNat/status/385254776432111616">Canadian Field Naturalist</a>, a <a href="http://www.canadianfieldnaturalist.ca/index.php/cfn">journal</a> that publishes ecology, behaviour, taxonomy, conservation, and other topics relevant to Canadian natural history, and because <a href="https://en.wikipedia.org/wiki/List_of_Canadian_provincial_and_territorial_symbols">Canadian provinces also have various representative symbols</a> (none reptilian, except for the feathered kind, which I might add are somewhat <a href="http://www.thebirdist.com/2013/04/state-birds-what-they-should-be.html">better chosen</a> than those of the US states), this summer I decided to cover the US's northern neighbor as well. Does Canada even have any snakes, you might ask? In fact, <a href="http://www.carcnet.ca/english/herps.php#snake">Canada is home to 27 species of snake</a>, which might surprise those of us who have grown up in regions farther south. That's enough for every province and territory to have two provincial snakes, with one left over, although the uneven geographic distribution of species precludes that, as we'll see. I followed the same "no duplication" rule as I did for the State Snakes, but I allowed snakes that had been used as U.S. State Snakes to be used again, because almost all of the species found in Canada had also been used for a U.S. state. Feel free to chime in with your opinion about what your favorite province's snake should be, if it differs from my choice.<br />
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<b>1. Alberta. Prairie Rattlesnake (<i>Crotalus viridis</i>)</b><br />
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<tr><td class="tr-caption" style="text-align: center;">Prairie Rattlesnake (<i>Crotalus viridis</i>)</td></tr>
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Alberta, well-known for its dinosaurs, also harbors a fairly substantial diversity of modern reptiles for a place with such long winters. Seven species of snake can be found in the province, but perhaps the most quintessential are Prairie Rattlesnakes. Prairie Rattlesnakes in Alberta occur in shortgrass prairies, dry grasslands, and sagebrush in the southeastern part of the province. At the northwestern edge of their range, Prairie Rattlesnakes in Alberta take 5-8 years to reach sexual maturity, and give birth to 4-12 live young, which are quite large (~11" long; compared to ~9" in the more southerly parts of their range). <a href="http://snakesarelong.blogspot.com/2015/11/snakes-that-are-good-parents.html">Females may remain with their young for up to 10 days after giving birth</a>. Historically, Prairie Rattlesnakes were found as far west as Calgary and almost as far north as Red Deer, but <a href="http://aep.alberta.ca/fish-wildlife/species-at-risk/species-at-risk-publications-web-resources/reptiles/documents/SAR-StatusPrairieRattlesnakeAlberta-2012Update-Dec2012.pdf">the species has declined in many areas</a> due to persecution and habitat loss. Venomous snakes are rarely very popular, but provincial symbol-hood might help establish rattlesnakes <a href="http://snakesarelong.blogspot.com/2016/05/rattlesnake-roundups-revisited.html">as wildlife to be valued rather than pests to be exterminated</a> (and Alberta <a href="http://lethbridgeherald.com/news/local-news/2016/04/21/all-snakes-in-alberta-are-protected-by-the-law/">is already quite progressive about protecting its snakes</a>).</div>
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<b>2. British Columbia. Sharp-tailed Snake (<i>Contia tenuis</i>)</b><br />
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<tr><td class="tr-caption" style="text-align: center;">Sharp-tailed Snake (<i>Contia tenuis</i>)</td></tr>
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BC might be my favorite province, principally because of the <a href="http://www.nanaimo.ca/EN/main/visitors/NanaimoBars.html">Nanaimo Bar</a>, a three-layer no-bake dessert created in the eponymous coastal city of Nanaimo. I chose the Sharp-tailed Snake to represent BC because in some ways it resembles a reversed Nanaimo Bar—the dorsal coloration is similar to the graham-cracker-and-almond base, the color of the sides to the vanilla custard center (sort of), and the belly to the delectable chocolate-and-coconut topping. These snakes are found on Vancouver Island, the nearby Gulf Islands, and possibly on the adjacent mainland. These cute little snakes eat slugs, including the infamous banana slugs, which I bet don't taste anywhere near as good as Nanaimo Bars. <a href="http://www.biodiversitylibrary.org/page/1779710#page/196/mode/1up">Descriptions of Sharp-tailed Snakes were first published in 1852</a> (by herpetologists Spencer Fullerton Baird & Charles Frédéric Girard, who received <a href="https://en.wikipedia.org/wiki/United_States_Exploring_Expedition">collections made the decade before</a> in the Puget Sound area), exactly 100 years before the first printed recipes featuring Nanaimo bar ingredients <a href="https://en.wikipedia.org/wiki/Nanaimo_bar">were published in the Women's Auxiliary to the Nanaimo Hospital Cookbook</a> (although I'll admit that's a pretty <i>tenuis</i> connection).</div>
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<b>3. Manitoba. Western Hog-nosed Snake (<i>Heterodon nasicus</i>)</b><br />
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<tr><td class="tr-caption" style="text-align: center;">Western Hog-nosed Snake (<i>Heterodon nasicus</i>)</td></tr>
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Even though Manitoba is very well-known for its <a href="http://www.naturenorth.com/spring/creature/garter/Narcisse_Snake_Dens.html">Narcisse Gartersnake Dens</a>, it has greater snake diversity than several of the other provinces, for which the gartersnake must be reserved. Some of Manitoba's most interesting snakes are Western Hog-nosed Snakes, which are found in sandy areas in the southwestern part of the province. As with other snakes at the northern limits of their range, they have a short activity season—they mate in May and lay 5-12 eggs in late June or early July, which then hatch by August. A study of Western Hog-nosed Snakes in <a href="https://www.gov.mb.ca/conservation/parks/popular_parks/western/spruce.html">Spruce Woods Provincial Heritage Park</a>, Manitoba, found that they emerge from their burrows on any day when they could achieve a body temperature of at least 29°C (84°F). Like gartersnakes (though not quite to the same extent), these snakes can achieve fairly high densities in certain areas, so I think they could be good candidates for expanding our knowledge of snake ecology and behavior in the wild into phylogenetically-uncharted territory, challenging <a href="https://books.google.com/books?id=n2rW7E8_uJoC&pg=PA55&lpg=PA55&dq=%22It%27s+a+good+thing+you+Yanks+have+garter+snakes,+or+you+wouldn%27t+have+anything+to+study.%22&source=bl&ots=KcatLcnzc7&sig=k4P2bYqz8LRkvXFV57NZn5QDqjo&hl=en&sa=X&ved=0ahUKEwjsrvGO6prNAhUBsBQKHejiCAQQ6AEIHDAA#v=onepage&q=%22It's%20a%20good%20thing%20you%20Yanks%20have%20garter%20snakes%2C%20or%20you%20wouldn't%20have%20anything%20to%20study.%22&f=false">the statement made by Rick Shine in 1987</a> that <i>"It's a good thing you Yanks have garter snakes, or you wouldn't have anything to study."</i><br />
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<b>4. Newfoundland & Labrador. Maritime Gartersnake (<i>Thamnophis sirtalis pallidulus</i>)</b><br />
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<tr><td class="tr-caption" style="text-align: center;">Maritime Gartersnake (<i>Thamnophis sirtalis pallidulus</i>)</td></tr>
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Newfoundland and Labrador is the only Canadian province without any native snakes. However, in recent years southwestern Newfoundland in the vicinity of St. David's <a href="http://www.thetelegram.com/News/Local/2012-10-09/article-3094976/Province-appears-snake-bitten/1">has apparently been colonized by Maritime Gartersnakes</a>, a beautiful subspecies of Common Gartersnake. Although no genetic analyses have been performed, it's likely that this population was founded by individuals shipped across the Gulf of St. Lawrence in hay bales or other cargo from Québec, New Brunswick, Nova Scotia, or Prince Edward Island. <a href="http://www.cbc.ca/news/canada/newfoundland-labrador/snakes-found-breeding-in-western-nfld-1.931515">A poll by the CBC</a> revealed that 12% of respondents thought that the recent colonization was "actually kind of cool", whereas a discouraging 49% of respondents were "not happy about it at all". <a href="http://www.reptilescanada.com/archive/index.php/t-63675.html">It's rumored that</a> gartersnakes were purposefully but unsuccessfully released in the St. John's area in eastern Newfoundland decades ago, either by farmers hoping to control rat populations or by someone who brought them back from the mainland hoping to sell them as pets (though both scenarios are likely more urban legend than fact). A string of recent mild winters may have allowed the gartersnakes in western Newfoundland to persist, but the extent to which climate change will enable a <a href="http://link.springer.com/article/10.1007%2Fs10530-008-9228-z">Florida-pythons scenario</a> writ-small in Newfoundland remains to be seen. At the very least, this could be a golden opportunity for snake biologists to study what happens when snakes enter an ecosystem from which they have been absent for thousands of years, a rare event even in <a href="http://snakesarelong.blogspot.com/2014/01/the-first-invasive-snake.html">an age of snake invasions</a>.</div>
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<b>5. New Brunswick. </b><b>Smooth Greensnake (<i>Opheodrys vernalis</i>)</b><br />
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<tr><td class="tr-caption" style="text-align: center;">Smooth Greensnake (<i>Opheodrys vernalis</i>)</td></tr>
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Soctsman <a href="https://en.wikipedia.org/wiki/Andrew_Leith_Adams">Andrew Leith Adams</a> was an army physician who served in India, Egypt, and Canada during the 1800s. He spent his spare time studying the natural history of these countries, about which he later wrote several books, including his 1873 <a href="http://www.biodiversitylibrary.org/item/129608#page/224/mode/1up" style="font-style: italic;">Field and forest rambles, with notes and observations on the natural history of eastern Canada</a>. In it, he wrote<i> "The Reptiles of New Brunswick are neither numerous nor </i><i>formidable."</i>, which, compared with the snake fauna he doubtless experienced in Egypt and India, was certainly true. He mentioned several snake species, in particular noting that "<i>One of our most common fangless snakes is the active little </i><i>green species (C. vernalis)"</i>, using the C. to abbreviate the genus <i>Coluber</i>, <a href="http://snakesarelong.blogspot.com/2015/06/the-linnaean-snakes-part-ii.html">which Linnaeus had used for practically all snakes</a> except boas and rattlesnakes. This handsome species has also frequently gone by the binomial <i>Liochlorophis vernalis</i>, among <a href="http://reptile-database.reptarium.cz/species?genus=Opheodrys&species=vernalis">a half-dozen other genera</a> into which it has been placed over the years.</div>
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<b>6. Northwest Territories. Red-sided Gartersnake (<i>Thamnophis sirtalis parietalis</i>)</b></div>
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<tr><td class="tr-caption" style="text-align: center;">Mating ball of <i>Thamnophis sirtalis parietalis</i></td></tr>
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Red-sided Gartersnakes are the only snakes found in the Northwest Territories, where they achieve high densities near Fort Smith between the southern shore of the Great Slave Lake and <a href="http://www.pc.gc.ca/eng/pn-np/nt/woodbuffalo/index.aspx">Wood Buffalo National Park</a>. Because there are few suitable hibernacula, thousands of individuals share the same den. Long winters and short, cool summers have resulted in a mating system that is unusual among snakes, although it is also possibly the most well-known <a href="http://www.jstor.org/stable/2426514">because it is easily studied</a>. Upon emergence from the in mid-April, snakes spend 2-3 weeks hanging around the entrance, during which time males compete fiercely to mate with females, forming colossal "mating balls". They then migrate over 2.3 miles (3.75 km) to their summer marshland habitat, where they remain until late August, giving birth to litters of young that are relatively <a href="ftp://dnrftp.dnr.state.md.us/Public/Wild/MARA%20Species%20Account%20Info/2_Reptiles/3_Snakes/Eastern%20Gartersnake/Larsen_1993_EAGA.pdf">small in number</a> (~12 vs. ~19 in Manitboa) and large in body size (191 mm SVL vs. 154 mm in Manitoba). Females in the NWT rarely give birth in two successive years, instead saving up energy from one year in order to reproduce the next. They also mature at larger body sizes (570 mm SVL vs. 527 mm in Manitboa) than snakes further south. I bent the rules a little here since both Newfoundland and the NWT have only <i>T. sirtalis </i>(they have different subspecies, and this species might be split up fairly soon). </div>
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<b>7. Nova Scotia. </b><b>Ring-necked Snake (<i>Diadophis punctatus</i>)</b><br />
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<tr><td class="tr-caption" style="text-align: center;">Brown-morph and normal <i>Diadophis punctatus</i> from Nova Scotia<br />
From <a href="http://www.canadianfieldnaturalist.ca/index.php/cfn/article/download/1131/1135">Gilhen 2011</a></td></tr>
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Ring-necked Snakes are cute little snakes that mostly eat invertebrates, although they have been known to snack on the occasional salamander. In Nova Scotia, they can be found almost throughout the province, and <a href="http://www.canadianfieldnaturalist.ca/index.php/cfn/article/download/1131/1135">an unusual brown morph</a> occurs, particularly on Big Tancook Island in Mahone Bay along the east coast. According to the notebooks of <a href="https://en.wikipedia.org/wiki/Harry_Piers">Harry Piers</a>, an early 20th century naturalist, museum curator, and historian, ringnecks were known to the native Mi'kmaq People as “the worst snake, Um-taa-kum (k)”, although it's not clear why. One communal nest found under a boulder near McCabe Lake in Halifax County contained 117 eggs, which must have been laid by at last 15, and probably many more, females (clutch size ranges from one to eight).</div>
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<b>8. Nunavut. Ellesmere Island erycine (Eocene boa)</b><br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg9_uJiloa6A3FOORVXqI9I3DM4x0Syb11PAEqvqGfkPBTmw73D-a1cvPoiNeWACGHVpL5FbRSEMLDroWIQBHB1B_jL-JY1i2TP30xGCZlJX1JLplwrl2pFPmn6A90G_yKy0dbgexgGvp6J/s1600/Ellesmere+Island+erycine+Estes+%2526+Hutchinson+1980_modified.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="129" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg9_uJiloa6A3FOORVXqI9I3DM4x0Syb11PAEqvqGfkPBTmw73D-a1cvPoiNeWACGHVpL5FbRSEMLDroWIQBHB1B_jL-JY1i2TP30xGCZlJX1JLplwrl2pFPmn6A90G_yKy0dbgexgGvp6J/s320/Ellesmere+Island+erycine+Estes+%2526+Hutchinson+1980_modified.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Drawing of Ellesmere Island erycine vertebra<br />
Dotted lines show best-guesses at broken-off parts<br />
A. Dorsal and B. right lateral view<br />
From Estes & Hutchison 1980</td></tr>
</tbody></table>
Unfortunately, there are no living wild snakes in Nunavut. Initially I was going to get around this by writing only about the true provinces, but then I found evidence that a 50-million-year-old fossil snake vertebrae was found on Ellesmere Island, above the Arctic Circle at about 78.5° north (find it <a href="https://paleobiodb.org/navigator/">here at the awesome new Paleobiology Database Navigator</a>). This vertebra belonged to an undescribed species of boid snake probably related to <a href="http://snakesarelong.blogspot.de/2012/04/utahs-boa.html">rubber boas</a>, and it was found in <a href="http://paleodb.org/bridge.pl?a=basicCollectionSearch&collection_no=15583">an Eocene fossil deposit</a> that used to be a lush river delta and floodplain, with abundant swamps, alongside pike, bowfin, and gar, mud & softshell turtles, alligators, monitor lizards, giant salamanders, and even primates. The single bone is part of the collection of the Canadian Museum of Nature (<a href="http://collections.nature.ca/en/Search/Search">specimen number 32403</a>) and hasn't been assigned to a species or even a genus because it's broken. Paleontologists are fairly confident that it is an erycine boid based on comparisons made with a half-dozen other extinct genera that probably belong in this group. <a href="http://home.gwu.edu/~rpyron/publications/Pyron_et_al_2014a.pdf">Recent phylogenies of booids</a> elevate Erycinae to a family, but do not include extinct taxa, so it's difficult to say for sure how these snakes were related to each other and to living species.</div>
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<b>9. Ontario. Eastern Foxsnake (<i>Pantherophis vulpinus</i>)</b><br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjiXz5MbJy-G2TPtt2siulpISalWdiID-MGe6tC62RkBNhk62MNQuhE6Hsr_LUlkNBaVy-p1eZY1wqvn0yTeNdD0HL7Vu7zThze-t0mi2vDgd8bQf_ldNjRpxpwXBJmOM_zzpvZGPCkea7A/s1600/Pantherophis+vulpinus+Nick+Scobel.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="213" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjiXz5MbJy-G2TPtt2siulpISalWdiID-MGe6tC62RkBNhk62MNQuhE6Hsr_LUlkNBaVy-p1eZY1wqvn0yTeNdD0HL7Vu7zThze-t0mi2vDgd8bQf_ldNjRpxpwXBJmOM_zzpvZGPCkea7A/s320/Pantherophis+vulpinus+Nick+Scobel.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Eastern Foxsnake (<i>Pantherophis vulpinus</i>)</td></tr>
</tbody></table>
<div style="text-align: justify;">
<b></b>Ontario has more snake species to choose from than any other province, including seven that are found nowhere else in Canada. At the JMIH meeting in Reno last summer, I posed the question of which one best represented Ontario to herpetologist <a href="http://laurentian.ca/faculty/jlitzgus">Jacqueline Litzgus</a>, a native of Ontario and a professor at Laurentian University. She was unhesitant in recommending the Eastern Foxsnake, the only species of snake whose range is mostly in Canada (which perhaps makes it sort of a national snake as well, although the common gartersnake is found in more provinces). Foxsnakes are large constrictors that are closely related to cornsnakes and (slightly less closely) to ratsnakes. They probably recolonized northern North America more quickly after the retreat of the glaciers than most snakes because of their mobility and the flat terrain left behind in the midwest. We once thought that the two species had a disjunct range, with the western foxsnake (formerly <i>P. vulpinus</i>) being found in the USA between the Missouri River and Lake Michigan, separated by a foxsnake-less area in northeastern Indiana and the lower peninsula of Michigan from the eastern foxsnake (formerly <i>P. gloydi</i>), which was found south and east of Lake Huron in Ontario, Michigan, and Ohio. However, <a href="http://www.southeastern.edu/acad_research/depts/biol/faculty/pdf/crother2011.pdf">a 2011 study</a> used evidence from a single mitochondrial gene to suggest that the Mississippi River seemed to be a more significant genetic barrier and that western foxsnakes east of the Big Muddy in Wisconsin and Illinois were more closely related to eastern foxsnakes than they were to western foxsnakes in Iowa and Minnesota. Because the type specimens for both former foxsnake species were within the eastern lineage, this species became <i>P. vulpinus</i> (the older name), <i>P. gloydi</i> disappeared, and the "new" western foxsnake was named <i>P. ramspotti. Runner up:</i> Massasauga (<i>Sistrurus catenatus</i>), because of the town of Missisauga, Ontario.</div>
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<b>10. Prince Edward Island. Red-bellied Snake (<i>Storeria occipitomaculata</i>)</b><br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiTCb1kxOaFeAEOYxiL6ReQ19sOxQibkFfidCkhV0KqPhhoBWpvbiBW8pD_Bwv_Il0HQrFnHuIPhhaPSaumyvWtUA6P-2zl-nxInEgad5MZIMFwGujj_nvH0-QcgMTcnYbJFbt71e5y67VJ/s1600/3973.jpg" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="212" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiTCb1kxOaFeAEOYxiL6ReQ19sOxQibkFfidCkhV0KqPhhoBWpvbiBW8pD_Bwv_Il0HQrFnHuIPhhaPSaumyvWtUA6P-2zl-nxInEgad5MZIMFwGujj_nvH0-QcgMTcnYbJFbt71e5y67VJ/s320/3973.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Red-bellied Snake (<i>Storeria occipitomaculata</i>)</td></tr>
</tbody></table>
<div style="text-align: justify;">
Located in the Gulf of St. Lawrence, Prince Edward Island was formed as a sandstone peninsula 250-300 million years ago. The end of the ice age 15,000 years ago and the retreat of the glaciers laid down glacial till and increased the sea level, disconnecting PEI from the mainland. <a href="http://www.carcnet.ca/english/reptiles/tour/province/repPEI.php">PEI only has three species of snakes</a>, all of which colonized the island within the last 15,000 years. Despite the fact that no lizards or turtles have been able to make the same crossing, PEI is still way ahead of Québec's similarly-sized <a href="http://www.chaireanticosti.ulaval.ca/en/ile_danticosti/">Île d'Anticosti</a>, which lies ~190 miles (~300 km) to the north and has no native species of amphibians or reptiles. Of the tiny red-bellied snake, PEI naturalist <a href="http://dalspace.library.dal.ca/bitstream/handle/10222/11251/Pt2Art7.pdf?sequence=1&isAllowed=y">John Mellish</a> wrote in the 1870s <i>"This variety is numerous, is smaller in size, and seems to be less courageous than some of the other species"</i>. Although Mellish got this much right, <a href="http://vre2.upei.ca/islandmagazine/fedora/repository/vre:islemag-batch2-369/OBJ/05_Things_that_slither_p_12-14.pdf">he was as prone to exaggeration as many modern observers</a>, interspersing his species accounts with tales of snakes charming their prey, swallowing their young, and attacking people. In reality, red-bellied snakes mostly attack slugs, and <a href="https://www.researchgate.net/publication/249488146_Lip-curling_in_redbelly_snakes_%28Storeria_occipitomaculata%29_functional_morphology_and_ecological_significance">their peculiar lip-curling display</a> is hardly threatening to a human.</div>
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<b> 11. Québec. Milksnake (<i>Lampropeltis triangulum</i>)</b><br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgeU2-K_JZ9Eziq6f8DiwRWeCqOe_2itB1GIlWDz6SG2OSX-u2XHtd42sKVUX8Fz4-zpbloWLK5FM4Bnw3fx9QlSe-igfLkQ6ipZg588kC3JVJHOQkrTIyoKVb_SCEz_hQHO42SoqH4ndIV/s1600/Lampropeltis+triangulum+Michael.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgeU2-K_JZ9Eziq6f8DiwRWeCqOe_2itB1GIlWDz6SG2OSX-u2XHtd42sKVUX8Fz4-zpbloWLK5FM4Bnw3fx9QlSe-igfLkQ6ipZg588kC3JVJHOQkrTIyoKVb_SCEz_hQHO42SoqH4ndIV/s320/Lampropeltis+triangulum+Michael.jpg" width="264" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Milksnake (<i>Lampropeltis triangulum</i>)</td></tr>
</tbody></table>
<div style="text-align: justify;">
Québec is best emblematized by the Milksnake, which was first described by a French herpetologist, <a href="https://en.wikipedia.org/wiki/Bernard_Germain_de_Lac%C3%A9p%C3%A8de">Bernard Germain de Lacépède</a>, in 1789. Lacépède's two-volume masterpiece, <i>Histoire Naturelle</i>, is a classic work in herpetology. Although Lacépède mostly used French vernacular names, (<a href="http://biodiversitylibrary.org/item/54321#page/457/mode/1up">"le triangle" for the milksnake</a>, after the double triangles on top of its head), he used <a href="http://snakesarelong.blogspot.com/2015/05/linnaean-snakes-part-i.html">Linnaeus's Latin binomial system</a> about 65% of the time in <a href="http://www.biodiversitylibrary.org/item/24722#page/150/mode/1up">a 59-page table</a> in the third section of the second volume, which covered legless amphibians and reptiles. However, because he was not consistent in his use of Latin binomials, the taxonomic community <a href="http://biodiversitylibrary.org/page/12229636#page/293/mode/1up">decided in 1987</a> that the names in volume two were not valid (volume one, which covers turtles, lizards, and amphibians, contains a 3.5' x 1.75' fold-out table that was consistently binomial, so these names remain valid). Four snake names, including <i>Lampropeltis triangulum</i>, were rescued because of their long history of use. The other three (<i>Agkistrodon piscivorus</i>, <i>Langaha madagascarensis</i>, and <i>Python reticulatus</i>) were much longer-used than <i>L. triangulum</i>, which probably wouldn't have made the cut if not for <a href="http://biodiversitylibrary.org/page/12223169#page/36/mode/1up">an earlier decision by the ICZN</a> as part of <a href="http://biodiversitylibrary.org/page/12222253#page/430/mode/1up">a case involving the mistaken identity of Linnaeus's scarletsnake</a> (<i>Cemophora coccinea</i>) specimen and the name he gave it, <i>Coluber doliatus</i>, which was mistakenly used for the milksnake for over 150 years. The 1967 case invalidated <i>doliatus</i> and fixed <i>triangulum</i> as the specific epithet of the milksnake, which prevented it from later being invalidated with the rest of Lacépède's snake names. In this way the species is somewhat rebellious (in a nomenclatural sense), which I think would please many Québécois.</div>
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<b>12. Saskatchewan. Gophersnake (<i>Pituophis catenifer</i>)</b><br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj_VrZtwvRx-LbNZOAP-8bC3ad2doOiEExQ-ajGH7A0DrnL5QNlpi42If2j71IRZU4i_VIQoYxD43rMiP6o00lRqy91ETAPOt-HTqoVFu66XkupcF9kfnDTOsimSoqy-ZLMHtqpPZMGa0Q8/s1600/Pituophis+catenifer+AMD.jpg" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj_VrZtwvRx-LbNZOAP-8bC3ad2doOiEExQ-ajGH7A0DrnL5QNlpi42If2j71IRZU4i_VIQoYxD43rMiP6o00lRqy91ETAPOt-HTqoVFu66XkupcF9kfnDTOsimSoqy-ZLMHtqpPZMGa0Q8/s320/Pituophis+catenifer+AMD.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Gophersnake (<i>Pituophis catenifer</i>)</td></tr>
</tbody></table>
<div style="text-align: justify;">
On the first page of one of my favorite novels, Farley Mowat's <i><a href="http://www.amazon.com/Owls-Family-Farley-Mowat/dp/0440413613">Owls in the Family</a></i>, the author describes growing up in Saskatoon, Saskatchewan: <i>"When you stepped off the end of the Railroad Bridge you stepped right onto the prairie and there you were—free as the gophers. Gophers were the commonest thing on the prairie. The little mounds of yellow dirt around their burrows were so thick, sometimes, it looked as if the fields had yellow measles." </i>Although I like owls, these days I more often have another gopher predator in mind—the eponymous gophersnake (<i>Pituophis catenifer</i>), also less-aptly known as the bullsnake. These harmless creatures are often mistaken for rattlesnakes, because they have a superficially similar pattern (and they do rattle their tails, although they have no specialized noise-making structure). Confusion over the common name led Edward Abbey or one of his editors to include the scientific name of the <a href="http://snakesarelong.blogspot.com/2012/09/recent-conservation-successes-with.html">eastern indigo snake</a> (aka the blue gophersnake), <i>Drymarchon corais couperi</i>, for the bullsnake in the essay 'The Serpents of Paradise' in <a href="https://books.google.com/books?id=Na5l5WZ5mQwC&pg=PA107&lpg=PA107&dq=Edward+Abbey+Drymarchon+corais+couperi+serpents+of+paradise&source=bl&ots=G9PiQY9ssX&sig=l2yO98Q-9vGQcToh24vuj7nIhEQ&hl=en&sa=X&ved=0CCQQ6AEwAWoVChMItNT98YKsxwIVRQmSCh1dtAce#v=onepage&q=Drymarchon%20corais%20couperi&f=false">the 1968 edition</a> of <i>Desert Solitaire</i> (although <a href="https://books.google.com/books?id=lkhMtksYyhYC&pg=PA20&dq=edward+abbey+desert+solitaire+pituophis&hl=en&sa=X&ved=0CB4Q6AEwAGoVChMI7JPn3arbxgIVSxmSCh33WQOk#v=onepage&q=edward%20abbey%20desert%20solitaire%20pituophis&f=false">it is correct in 1988 edition</a>).</div>
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<b>13. Yukon. ?</b><br />
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<div style="text-align: justify;">
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh3HNdI7A6-IMPRPHKv7RlMEHBNdzHQhQR7yG0molGr1izpeB8vtiTfOU8h-HkR3jP5fYe1BtkYZD3FzruIA_ITVqB6O5JZIgt2stYT3LADlK9bRscaoPSDmjs3A6mzQarxg3WmOi7IrQQH/s1600/Yukon_Ho.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="200" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh3HNdI7A6-IMPRPHKv7RlMEHBNdzHQhQR7yG0molGr1izpeB8vtiTfOU8h-HkR3jP5fYe1BtkYZD3FzruIA_ITVqB6O5JZIgt2stYT3LADlK9bRscaoPSDmjs3A6mzQarxg3WmOi7IrQQH/s200/Yukon_Ho.jpg" width="188" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">I hope they find a snake</td></tr>
</tbody></table>
The Yukon Territory has no living snakes and no snake fossils (yet). This is actually quite ironic, because most living North American snakes crossed into our continent from Asia over the Bering Land Bridge, and some of them almost certainly slithered through what is today the Yukon. It is possible that somewhere in the southern Yukon exists a population of gartersnakes, which are found in the southern NWT and <a href="http://aknhp.uaa.alaska.edu/herps/enigma.htm">also possibly in the Alaskan panhandle</a>. <a href="http://snakesarelong.blogspot.com/2013/06/what-state-snakes-should-be-part-i.html">Three reliable sight records</a> and one specimen (now lost) from remote areas along Taku & Stikine Rivers in Alaska give us hope, although unfortunately neither basin enters the Yukon. Other snake sightings of snakes from Alaska include odd <i>T. sirtalis </i>and <i>T. ordinoides</i> specimens from more urban areas, which almost certainly represent translocations (<a href="http://www.public.iastate.edu/~fjanzen/pdf/11JHerpetol.pdf">genetic evidence supports this in at least one case</a>). <i>T. sirtalis</i> are found just 200 miles (320 km) south of the Yukon border in BC. It isn't completely crazy to imagine snakes living at such northerly latitudes; European Adders (<i>Vipera berus</i>) are found above the Arctic Circle in Scandinavia. If nothing else, gartersnakes from British Columbia will probably disperse there eventually if climate change keeps up with predictions.</div>
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<div style="text-align: center;">
ACKNOWLEDGMENTS</div>
<div style="text-align: center;">
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Thanks to <a href="https://www.flickr.com/photos/squamatologist/">Ben Lowe</a>, <a href="http://www.flickr.com/people/33102730@N02/">David O'Connor</a>, <a href="https://www.flickr.com/people/57809070@N03/">JD Willson</a>, <a href="https://www.flickr.com/photos/twpierson">Todd Pierson</a>, <a href="https://www.flickr.com/photos/63265212@N03/">Andy Teucher</a>, <a href="https://www.flickr.com/people/28113115@N00/">Michael</a>, <a href="http://www.californiaherps.com/info/photouse.html">Gary Nafis</a>, and <a href="https://www.flickr.com/photos/25268283@N06/">Nick Scobel</a> for the use of their photos, to Jackie Litzgus for helping me make the decision about Ontario, and to Gareth Hopkins for introducing me to Nanaimo bars.</div>
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<div style="text-align: center;">
REFERENCES</div>
<div style="text-align: center;">
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhwCKlR3oPG4K8rzjK9rKT5HAxjq1txytLDAir3ujr4wNmrmMOPHgaIrs863BEO4D7pxxLwLL5B-7MEjmOZiNIeMp1eErnSZttryhud3U4w2LLOsJkUTi1BbKp-yxgj-rNXTVd0w9cgfXuz/s1600/Thamnophis+Manitoba+U-Haul.jpg" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="219" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhwCKlR3oPG4K8rzjK9rKT5HAxjq1txytLDAir3ujr4wNmrmMOPHgaIrs863BEO4D7pxxLwLL5B-7MEjmOZiNIeMp1eErnSZttryhud3U4w2LLOsJkUTi1BbKp-yxgj-rNXTVd0w9cgfXuz/s320/Thamnophis+Manitoba+U-Haul.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Manitoba <i>Thamnophis</i> on the side of a U-Haul truck</td></tr>
</tbody></table>
<span style="font-size: x-small;">Anonymous. 1987. Opinion 1463. De Lacépède, 1788-1789, Histoire Naturelle des Serpens and later editions: rejected as a non-binominal work. Bulletin of Zoological Nomenclature 44:265-267 <<a href="http://biodiversitylibrary.org/page/12229636#page/293/mode/1up">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Baird, S.F. and C. Girard. 1852. Descriptions of new species of reptiles, collected by the U.S. exploring expedition under the command of Capt. Charles Wilkes, U.S.N. First part. - Including the species from the Western coast of America. Proceedings of the Academy of Natural Sciences of Philadelphia 6:174-177 <<a href="http://www.biodiversitylibrary.org/page/1779710#page/196/mode/1up">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Brongersma, L.D. 1972. On the “Histoire naturelle des Serpens” by de la Cépède, 1789 and 1790, with a request to reject this work as a whole, and with proposals to place seven names of snakes, being nomina oblita, on the Official index of rejected and invalid names in zoology, and to place three names of snakes on the Official list of specific names in zoology (Class Reptilia). Bulletin of Zoological Nomenclature 29:44-61 <<a href="http://biodiversitylibrary.org/page/12224790#page/62/mode/1up">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Crother, B.I., M.E. White, J.M. Savage, M.E. Eckstut, M.R. Graham, and D.W. Gardner. 2011. A reevaluation of the status of the Foxsnakes <i>Pantherophis gloydi</i> Conant and<i> P. vulpinus</i> Baird and Girard (Lepidosauria). ISRN Zoology 2011 <<a href="http://www.southeastern.edu/acad_research/depts/biol/faculty/pdf/crother2011.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Estes R, Howard Hutchison J, 1980. Eocene lower vertebrates from Ellesmere Island, Canadian Arctic Archipelago. Palaeogeography, Palaeoclimatology, Palaeoecology 30:325-347 <<a href="http://www.sciencedirect.com/science/article/pii/0031018280900644">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Gilhen, J. 2011. The Brown Morph of the Northern Ringneck Snake, <i>Diadophis punctatus edwardsii</i>, on Big Tancook Island, Mahone Bay, Nova Scotia. The Canadian Field-Naturalist 125:69-71 <<a href="http://www.canadianfieldnaturalist.ca/index.php/cfn/article/download/1131/1135">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Hodge, R.P. 1976. Amphibians and Reptiles in Alaska, the Yukon, and Northwest Territories. Alaska Northwest Pub. Co.</span></div>
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<span style="font-size: x-small;">Larsen KW, Gregory PT, Antoniak R, 1993. Reproductive ecology of the Common Garter Snake <i>Thamnophis sirtalis</i> at the northern limit of its range. American Midland Naturalist 129:336-345 <<a href="http://www.jstor.org/stable/2426514">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Leavesley, L.K. 1987. Natural history and thermal relations of the Western Hognose Snake (<i>Heterodon nasicus nasicus</i>) in southwestern Manitoba. MS thesis. University of Manitoba, Winnipeg, Manitoba.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Rossman, D.A., N.B. Ford, and R.A. Seigel. 1996. The Garter Snakes: Evolution and Ecology. University of Oklahoma Press, Norman, Oklahoma. (Shine quote opens <a href="https://books.google.com/books?id=n2rW7E8_uJoC&pg=PA55&lpg=PA55&dq=%22It%27s+a+good+thing+you+Yanks+have+garter+snakes,+or+you+wouldn%27t+have+anything+to+study.%22&source=bl&ots=KcatLcnzc7&sig=k4P2bYqz8LRkvXFV57NZn5QDqjo&hl=en&sa=X&ved=0ahUKEwjsrvGO6prNAhUBsBQKHejiCAQQ6AEIHDAA#v=onepage&q=%22It's%20a%20good%20thing%20you%20Yanks%20have%20garter%20snakes%2C%20or%20you%20wouldn't%20have%20anything%20to%20study.%22&f=false">chapter 4, page 55</a>)</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">West, R.M., M.R. Dawson, and J.H. Hutchison. 1977. Fossils from the Paleogene Eureka Sound Formation, N.W.T., Canada; occurrence, climatic and paleogeographic implications. Milwaukee Public Museum Contributions in Biology and Geology 2:77-93.</span><br />
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="http://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
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Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com11Logan, UT, USA41.7369803 -111.833835941.6422018 -111.9951974 41.831758799999996 -111.6724744tag:blogger.com,1999:blog-7443075087825368900.post-80429849632586124102016-06-08T02:00:00.000-06:002016-07-25T17:49:07.976-06:00Virgin Birth, the Color of Fossil Snakes, and More Recent Updates<div>
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<span style="font-size: x-small;">This post will soon be available in Spanish</span></div>
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As I <a href="http://snakesarelong.blogspot.com/2016/03/state-snakes-linnaean-names-and-other.html">did in March</a>, I wanted to highlight some recent and exciting updates to some of my older articles.<br />
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<a href="http://snakesarelong.blogspot.com/2014/02/snakes-that-give-virgin-birth.html"><b>Snakes That Give Virgin Birth</b></a><br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj9vfmJKS8nqNw2mSk8ZGf1bn3ebjZU8vRG5Svz_8c1Nu2t87fx_Pnbu776JwcThAlFoPRgELA17zzq60b2rqY3R3ru182pSETivL5GQG2HL-dScypJzc89o-1GxiY9SI4tpJacJKLpgNSc/s1600/Booth+%2526+Schuett+2016+Fig2+phylogenetic+pattern+of+parthenogenesis+in+snakes.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="267" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj9vfmJKS8nqNw2mSk8ZGf1bn3ebjZU8vRG5Svz_8c1Nu2t87fx_Pnbu776JwcThAlFoPRgELA17zzq60b2rqY3R3ru182pSETivL5GQG2HL-dScypJzc89o-1GxiY9SI4tpJacJKLpgNSc/s400/Booth+%2526+Schuett+2016+Fig2+phylogenetic+pattern+of+parthenogenesis+in+snakes.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Phylogenetic pattern of parthenogenesis in snakes<br />
Molecular tree on left, morphological tree on right<br />
From <a href="http://eherp.com/pdf/88512.pdf">Booth & Schuett 2016</a></td></tr>
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When I wrote about asexual reproduction in snakes in February 2014, new records of this phenomenon were rapidly accumulating, from snakes as <a href="http://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">distantly related</a> as cottonmouths and boa constrictors. In <a href="http://eherp.com/pdf/88512.pdf">a new paper</a>, <a href="http://www.booth-lab.org/">Warren Booth</a> and <a href="https://www.researchgate.net/profile/Gordon_Schuett">Gordon Schuett</a> review the knowns and unknowns of "virgin birth" in snakes, a subject which has become their specialty (it even has <a href="https://www.facebook.com/groups/463481383787022/">its own Facebook group</a>). Although obligate parthenogenesis is still known only from <a href="http://snakesarelong.blogspot.com/2014/04/the-most-widespread-snake-in-world.html">Brahminy Blindsnakes (<i>Indotyphlops braminus</i>)</a>, the new summary reports that facultative parthenogenesis has now been documented in 20 species of alethinophidian<a href="#1" name="top1"><sup>1</sup></a> snakes, and this list is anticipated to grow, although so far confirmed cases are limited to <a href="http://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">five lineages</a>: boids, pythonids, <i><a href="http://snakesarelong.blogspot.com/2014/08/filesnakes-wartsnakes-or-elephant.html">Acrochordus</a></i>, Crotalinae, and Natricinae. This new synthesis formalizes one of the trends that I wrote about in 2014, namely distinguishing between "Type A" facultative parthenogenesis, in which the offspring produced are large clutches of viable females that seem to have <a href="http://snakesarelong.blogspot.com/2016/02/dragonsnakes-and-filesnakes-revisited.html">a strange "WW" sex chromosome arrangement</a> (apparently typical of boas and pythons), and "Type B" facultative parthenogenesis, which is where all the offspring are male and few are born alive, many with extreme developmental abnormalities (apparently typical of colubroids).<br />
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Most intriguing is the hypothesis laid out for explaining this dichotomy: that boas and pythons (and possibly other basal alethinophidian snakes) might have an XY sex determination system rather than a ZW one like most snakes. Changes from ZW to XY or vice versa (and between genetic and temperature-dependent sex determination) <a href="http://mbe.oxfordjournals.org/content/early/2015/03/08/molbev.msv023.full">have been documented in geckos</a> and turtles, and could have been overlooked in boas and pythons due to their <a href="http://snakesarelong.blogspot.com/2016/02/dragonsnakes-and-filesnakes-revisited.html">similar-looking sex chromosomes</a> (tests are currently underway to falsify or verify this hypothesis). If true, this would explain the production of all-female offspring by facultative parthenogenesis; instead of WW, those females would be XX, just like humans!<br />
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<b><a href="http://snakesarelong.blogspot.com/2012/11/identifying-snake-sheds-part-iii.html">Identifying Snake Sheds</a></b><br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjAn1yGtCxEATKG_wvkSkv8HvE2KTXGCPRo-ss6CxqpoKw0-emoQBrsYihRicDTque9xMdzeNFOml1P6gZvuGysJHh3bpWBk7aUiDK_vypc46Nj0oRtoLZULsw214iTgSyREG2NpAxpBa7L/s1600/McNamara+et+al+2015+Fig4b.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="267" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjAn1yGtCxEATKG_wvkSkv8HvE2KTXGCPRo-ss6CxqpoKw0-emoQBrsYihRicDTque9xMdzeNFOml1P6gZvuGysJHh3bpWBk7aUiDK_vypc46Nj0oRtoLZULsw214iTgSyREG2NpAxpBa7L/s320/McNamara+et+al+2015+Fig4b.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">True-color representation of the fossil snake<br />
(MNCN 66503) in <a href="http://www.cell.com/current-biology/pdf/S0960-9822(16)30120-8.pdf">McNamara et al. 2016</a>.<br />
The dentition looks too <a href="http://snakesarelong.blogspot.com/2013/09/basics-of-snake-fangs.html">solenoglyphous</a> for a<br />
colubrid, although the 10-million year old specimen,<br />
which is missing its head, has not and<br />
probably can not be identified to species.</td></tr>
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Ever since the <a href="https://www.researchgate.net/profile/Liliana_DAlba/publication/41408373_Plumage_color_patterns_of_an_extinct_dinosaur/links/0fcfd50be126120e25000000.pdf">first reports of color from the skin and feathers of dinosaur fossils were published in <i>Science</i> in 2010</a>, I've been fascinated by the ability of paleontologists to see in color when they look into the past. <a href="http://www.cell.com/current-biology/pdf/S0960-9822(16)30120-8.pdf">A new paper in the journal <i>Current Biology</i></a> reveals the color of a fossil snake, determined from using scanning electron microscopy (SEM) to examine microfossils of certain types of skin cells, called chromatophores. So far, only melanin-based chromatophores (melanosomes, which are responsible for brown and black color) have been detected in fossilized skin and feathers, probably <a href="http://rsbl.royalsocietypublishing.org/content/9/3/20130184.short">because they are the most resistant to the decomposition process</a>. But, this study was also able to detect and measure other types of chromatophores from fossilized skin, including xanthophores (responsible for yellow, orange, or red color, derived from carotenoids or pteridines) and iridophores (responsible for iridescence). By comparing the fossil's chromatophore abundance and position to that of living reptiles, they were able to reconstruct the original color and pattern of the fossil snake's skin. For example, in the skin of living snakes, xanthophores with many more pteridine granules than carotenoid granules produce a red hue, whereas xanthophores with equal amounts of carotenoid granules and pteridine granules—as in the fossil—produce yellowish hues. Skin regions with abundant iridophores and xanthophores, but relatively few melanophores, are associated with green hues <a href="https://www.researchgate.net/profile/Masami_Hasegawa2/publication/6752991_Ultrastructure_of_the_dermal_chromatophores_in_a_lizard_(Scincidae_Plestiodon_latiscutatus)_with_conspicuous_body_and_tail_coloration/links/00b4953185beda85e0000000.pdf">in some living skinks</a>, whereas skin regions with many melanophores, a few xanthophores, and no iridophores suggest correspond to dark brown or black tones. As you can see in the depiction, this snake seems to have had a pale, creamy venter and a green back and sides, with areas of brown/black and yellow/green, perhaps not unlike modern Green Watersnakes (<i>Nerodia floridana</i>) or Boomslangs (<i>Dispholidus typus</i>).<br />
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<b><a href="http://snakesarelong.blogspot.com/2012/07/snakes-flying-without-planes.html">Snakes Flying Without Planes</a></b><br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgESqU01Hcils9am7mzS-7Tgj00dlZnLPMjs3c8crlEiAIBXiMmZVY5ZB3D2089ZvWHoUah18jUDt1Xwsn4FmKPnN-m7SVotOxsLTW7nY_Ortkt21akaHJENZvuNDn9_zMvkcMxambgobXg/s1600/Chrysopelea+paradisi+mating+Kaiser+et+al+2016.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="197" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgESqU01Hcils9am7mzS-7Tgj00dlZnLPMjs3c8crlEiAIBXiMmZVY5ZB3D2089ZvWHoUah18jUDt1Xwsn4FmKPnN-m7SVotOxsLTW7nY_Ortkt21akaHJENZvuNDn9_zMvkcMxambgobXg/s400/Chrysopelea+paradisi+mating+Kaiser+et+al+2016.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Photo and diagram of courtship behavior of <i>Chrysopelea paradisi</i><br />
Taken at the Sepilok Jungle Resort in Sabah, Malaysia<br />
Female shown in gray, males in blue, green, and orange<br />
From <a href="https://www.researchgate.net/publication/295912393_Tangled_skeins_a_first_report_of_non-captive_mating_behavior_in_the_Southeast_Asian_Paradise_Flying_Snake_Reptilia_Squamata_Colubridae_Chrysopelea_paradisi">Kaiser et al. 2016</a></td></tr>
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<a href="https://www.researchgate.net/publication/295912393_Tangled_skeins_a_first_report_of_non-captive_mating_behavior_in_the_Southeast_Asian_Paradise_Flying_Snake_Reptilia_Squamata_Colubridae_Chrysopelea_paradisi">A new report on the mating behavior</a> of Paradise Flying Snakes (<i>Chrysopelea paradisi</i>) showed that their courtship can involve multiple males. Although <a href="http://www.flyingsnake.org/">several experiments</a> have been performed on the gliding behavior of these snakes, almost nothing is known about their natural history in the wild. Males of many species of snakes court females <i>en masse</i> by rubbing their chins along their bodies, a behavior which allows them to sense her sex pheromones and <a href="http://snakesarelong.blogspot.com/2014/03/why-do-snakes-have-two-penises.html">jockey for position</a>. The role played by the female in choosing a male is unclear in most snake species; although conventional biological wisdom suggests that females should be the choosy sex, male-male competition seems to dominate courtship behavior in several species of snakes. Multi-male courtship behavior precedes mating in some well-studied temperate snakes (<i>e.g.,</i> gartersnakes emerging from hibernation), as well as in some tropical species (<i>e.g.,</i> anacondas, some other southeast Asian colubrids, such as <i>Boiga irregularis </i>and <i>Dryophiops rubescens</i>). Of course, it seems that most female snakes can store sperm for long periods of time, and they may have some control over which male's sperm to use to fertilize their eggs, so the genetic contribution of a female snake's male partners may not follow from their courtship or mating success. Unlike the terrestrial or aquatic mating balls documented for other snakes, the flyingsnakes in this observation were able to move as a unit for almost 50 feet through complex habitat—under a porch, up a tree—an adaptation that seems to fit their active, arboreal lifestyle and might help reduce the likelihood of a predatory attack during what must otherwise be a vulnerable time.</div>
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<a href="https://www.blogger.com/null" name="1"><b>1 </b></a>In a few places, the authors use "<a href="http://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">alethinophidian</a>" to refer to boas, pythons, and their relatives but not caenophidians, when instead they should have either used "henophidian" or "basal alethinophidian" (they mostly use the latter term throughout). Many people don't like the term "henophidian" because it is a paraphyletic group, but it is a convenient way to refer to non-scolecophidian, non-caenophidian snakes. In my mind it's essentially synonymous with "basal/stem alethinophidian". Alethinophidians are all snakes except for blindsnakes (scolecophidians), and Caenophidia is a subset of Alethinophidia. There are also at least three references to "Caenophidia + Colubroidea", which is confusing because Colubroidea is a subgroup of Caenophidia, and Caenophidia = Colubroidea + <i>Acrochordus</i>, which is perhaps what they meant.<a href="#top1"><sup>↩</sup></a><br />
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ACKNOWLEDGMENTS</div>
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Thanks to Gordon Schuett for clearing up some of the details of his recent paper.</div>
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REFERENCES</div>
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<span style="font-size: x-small;">Booth W, Schuett GW (2016) The emerging phylogenetic pattern of parthenogenesis in snakes. Biological Journal of the Linnaean Society 118:172-186 <<a href="http://eherp.com/pdf/88512.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Gamble, T., J. Coryell, T. Ezaz, J. Lynch, D. Scantlebury, and D. Zarkower. 2015. Restriction site-associated DNA sequencing (RAD-seq) reveals an extraordinary number of transitions among gecko sex-determining systems. Molecular Biology and Evolution 32:1296-1309 <<a href="http://mbe.oxfordjournals.org/content/early/2015/03/08/molbev.msv023.full">link</a>></span><br />
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<span style="font-size: x-small;">Kaiser H, Lim J, Worth H, O’Shea M (2016) Tangled skeins: a first report of non-captive mating behavior in the Southeast Asian Paradise Flying Snake (Reptilia: Squamata: Colubridae: <i>Chrysopelea paradisi</i>). Journal of Threatened Taxa 8:8488–8494 <<a href="https://www.researchgate.net/publication/295912393_Tangled_skeins_a_first_report_of_non-captive_mating_behavior_in_the_Southeast_Asian_Paradise_Flying_Snake_Reptilia_Squamata_Colubridae_Chrysopelea_paradisi">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Kuriyama, T., K. Miyaji, M. Sugimoto, and M. Hasegawa. 2006. Ultrastructure of the Dermal Chromatophores in a Lizard (Scincidae: Plestiodon latiscutatus) with Conspicuous Body and Tail Coloration. Zoological Science 23:793-799 <<a href="https://www.researchgate.net/profile/Masami_Hasegawa2/publication/6752991_Ultrastructure_of_the_dermal_chromatophores_in_a_lizard_(Scincidae_Plestiodon_latiscutatus)_with_conspicuous_body_and_tail_coloration/links/00b4953185beda85e0000000.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Li, Q., K. Q. Gao, J. Vinther, M. D. Shawkey, J. A. Clarke, L. D’Alba, Q. Meng, D. E. G. Briggs, and R. O. Prum. 2010. Plumage color patterns of an extinct dinosaur. Science 327:1369 <<a href="https://www.researchgate.net/profile/Liliana_DAlba/publication/41408373_Plumage_color_patterns_of_an_extinct_dinosaur/links/0fcfd50be126120e25000000.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">McNamara, Maria E., Patrick J. Orr, Stuart L. Kearns, L. Alcalá, P. Anadón, and E. Peñalver. 2016. Reconstructing Carotenoid-Based and Structural Coloration in Fossil Skin. Current Biology <<a href="http://www.cell.com/current-biology/pdf/S0960-9822(16)30120-8.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">McNamara, M. E., D. E. G. Briggs, P. J. Orr, D. J. Field, and Z. Wang. 2013. Experimental maturation of feathers: implications for reconstructions of fossil feather colour. Biology Letters 9 <<a href="http://rsbl.royalsocietypublishing.org/content/9/3/20130184.short">link</a>></span></div>
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="https://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
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Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com0tag:blogger.com,1999:blog-7443075087825368900.post-89077764901880765362016-05-28T14:16:00.000-06:002016-08-16T20:47:11.068-06:00Rattlesnake Roundups Revisited<div style="text-align: center;">
<span style="font-size: x-small;">This article will soon be available in Spanish</span></div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgl-apIrKvxb_B7JNzbi2QywH7E7CvXmYCt64Hv-uJC_Fnou2hTwMtuWuyUtHNscNvExlcxunxpDOJM7PiYvI-798vAAOz6R8s-MS2f_1YZxNq8S4Lu6_dSAuDHehnJr4fY8_hiHIWczA9i/s1600/2016SwRR_board.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgl-apIrKvxb_B7JNzbi2QywH7E7CvXmYCt64Hv-uJC_Fnou2hTwMtuWuyUtHNscNvExlcxunxpDOJM7PiYvI-798vAAOz6R8s-MS2f_1YZxNq8S4Lu6_dSAuDHehnJr4fY8_hiHIWczA9i/s320/2016SwRR_board.jpg" width="277" /></a></td></tr>
<tr><td class="tr-caption" style="font-size: 12.8px; text-align: center;">A chalkboard at the 2016 Sweetwater Rattlesnake Roundup,<br />
showing that a record number of pounds of snake had<br />
already been bought and sold by the second day, and that<br />
commerce was suspended on the third and fourth days of the<br />
event due to the massive surplus.<br />
Photo source unknown.</td></tr>
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At the 58th annual Sweetwater Rattlesnake Roundup this March, <a href="http://texashillcountry.com/sweetwaters-record-rattlesnake-roundup/">a record 24,481 pounds of rattlesnakes</a> (about 21,000 individuals), primarily Western Diamond-backed Rattlesnakes (<i>Crotalus atrox</i>), were slaughtered. That's over four times the all-time average and about five times the recent average, breaking from <a href="http://tpwd.texas.gov/huntwild/wild/wildlife_diversity/nongame/media/TPWD-SHWG-References.pdf">a trajectory of slow decline</a> at the few remaining rattlesnake roundups. The Sweetwater Jaycees attribute this year’s record catch to <a href="http://www.npr.org/sections/thetwo-way/2015/05/25/409474791/more-heavy-rain-predicted-for-texas-oklahoma">heavy rains</a>, an explanation which might hold some water, but another probable contributing factor is <a href="http://www.npr.org/2016/05/14/477694584/amid-much-hissing-texas-could-impose-ban-to-rattle-snake-hunters">the possibility of an impending Texas Parks & Wildlife ban</a> on using gasoline fumes to collect rattlesnakes, which was discussed this week at a meeting in Austin on May 25th, 2016. The Texas Parks and Wildlife Commission <a href="http://www.lampasasdispatchrecord.com/news/2016-06-28/Front_Page/Talks_resume_about_rattlesnake_rules.html">decided to begin developing language for a new rule either prohibiting or further regulating this practice in the state</a>. The rule is still far from going into effect, and would include a two-year delay on the effective date. It won't be reviewed again until November 2016 (at which time, watch this space for a link to an opportunity for a public comment, if available). <a href="http://tpwd.texas.gov/faq/huntwild/gassing.phtml">TPWD's Snake Harvest Working Group recommended earlier this year</a> that Texas join 29 other states in banning this environmentally-harmful practice, which <a href="http://tpwd.texas.gov/huntwild/wild/wildlife_diversity/nongame/media/TPWD-SHWG-References.pdf">has been shown to kill numerous non-target species</a> and has been compared with other unsportsmanlike methods of hunting, such as shooting at an out-of-range bird or fishing with dynamite. The state wildlife agency has been moving slowly but steadily to regulate rattlesnake collection in Texas because of the economic importance of rattlesnake roundups to towns like Sweetwater (<i>e.g., </i><a href="http://sweetwatertexas.org/wp-content/uploads/2015/12/Sweetwater%20Rattlesnake%20Roundup_EIA_report-20154_FINAL.pdf">over 25,000 people contributed over $8 million to the local economy in 2015</a>, although the <a href="http://tpwd.texas.gov/huntwild/wild/wildlife_diversity/nongame/media/TPWD-SHWG-Report.pdf">TPWD report</a> found that the weather and the diversity of other events had stronger associations with profits than the number of rattlesnakes at an event).</div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgtbEHJTsVrQjl-3LyTMdLxP9z1_Sy-Lui1NCKxG4npyBSFEEfAhc31tznKqEpHOBUmzCVcMiLwQQJZLgSEJVr8BFscbcevpb3OF8Vuxx9YAOHdElnhCgCDu-gm4_Edz3H4FLzk0xT4YN7z/s1600/remaining+RRs+TWPD+RefDoc5Fig2.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgtbEHJTsVrQjl-3LyTMdLxP9z1_Sy-Lui1NCKxG4npyBSFEEfAhc31tznKqEpHOBUmzCVcMiLwQQJZLgSEJVr8BFscbcevpb3OF8Vuxx9YAOHdElnhCgCDu-gm4_Edz3H4FLzk0xT4YN7z/s320/remaining+RRs+TWPD+RefDoc5Fig2.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Locations of the remaining rattlesnake roundups,<br />
including non-lethal festivals.<br />
From <a href="http://tpwd.texas.gov/huntwild/wild/wildlife_diversity/nongame/media/TPWD-SHWG-References.pdf">TPWD Report Reference Document (p. 22)</a></td></tr>
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Ironically, this year's surplus of snakes drove the price of rattlesnake down so much (historically as high as $10.00 per pound, this year the price fell below $0.50/lb. despite efforts to maintain higher prices) that only about a quarter of the rattlesnakes collected were purchased for their meat, rattles, and skins before all demand had been exhausted. <a href="http://tpwd.texas.gov/huntwild/wild/wildlife_diversity/nongame/media/TPWD-SHWG-References.pdf">Rattlesnakes collected using gassing are no longer purchased by the antivenom industry</a>, because of their short lifespan and poor health (as well as a <a href="http://www.genetics.org/content/199/1/165">more nuanced understanding of the importance of geographic variation in venom composition</a>, emphasizing the necessity of knowing the geographic origin of each snake used in venom research). The fate of the rattlesnakes left unsold after Sweetwater (<a href="http://www.reporternews.com/news/big-country/no-market-for-rattlesnakes-after-record-roundup-2e566fd6-cf31-11a7-e053-0100007f77a0-372621081.html">which some have speculated as being up to 75,000</a>) has not been made public, although <a href="http://texashillcountry.com/sweetwaters-record-rattlesnake-roundup/">reports suggest that prices are also down at other roundups in Texas and Oklahoma</a>, possibly as a result of vendors trying to sell their snakes there. Anyone who has gone to great expense to collect snakes in this manner and now cannot find a buyer is at risk of losing their investment. <a href="http://texashillcountry.com/sweetwaters-record-rattlesnake-roundup/">Claims about the impacts on snakebites to humans and livestock</a> if these snakes were to be released are unsubstantiatable and untrue, considering that the survival of wild snakes captured and released elsewhere <a href="https://www.jstor.org/stable/1565542?seq=1#page_scan_tab_contents">is greatly reduced</a> (not to mention the <a href="http://snakesarelong.blogspot.com/2013/11/the-truth-about-snakebite.html">dubiousness of the link between rattlesnake abundance and snakebite frequency in the first place</a>).<br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEho49nXni7dFTCLkDi4ygL8gQK2EySFZdvDnLF-n_ejdMqKuwBqanF9OQEjEAg4naDeWtnHuV6VGd758MN8GyQT6Pkbmfq3yxXmkuGL2I0aikrxKjtgsxxAV04UbFn32wekTmTmzwb_lic7/s1600/money+%2526+snakes+over+time+TWPD+RefDoc5Fig2.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="290" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEho49nXni7dFTCLkDi4ygL8gQK2EySFZdvDnLF-n_ejdMqKuwBqanF9OQEjEAg4naDeWtnHuV6VGd758MN8GyQT6Pkbmfq3yxXmkuGL2I0aikrxKjtgsxxAV04UbFn32wekTmTmzwb_lic7/s400/money+%2526+snakes+over+time+TWPD+RefDoc5Fig2.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Trajectory of profit (red, blue), number of snakes (purple), and<br />
weather conditions (green) at the Sweetwater Roundup over the last decade.<br />
Chart prepared by Rob Denkhaus, TPWD Wildlife Diversity Advisory Committee<br />
and presented in <a href="http://tpwd.texas.gov/huntwild/wild/wildlife_diversity/nongame/media/TPWD-SHWG-References.pdf" style="font-size: 12.8px;">TPWD Report Reference Document (p. 64)</a></td></tr>
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I am hopeful that eventually all stakeholders can <a href="http://www.npr.org/2016/05/14/477694584/amid-much-hissing-texas-could-impose-ban-to-rattle-snake-hunters">overcome the cognitive dissonance</a> between the flawed concept of predator population control (which was the original impetus behind rattlesnake roundups) and the implicit economic reasons behind their persistence. Although rattlesnake roundups are inarguably sensational and exploitative, claims about the sustainability of the wild rattlesnake harvest cannot currently be independently evaluated (I encourage anyone interested in the subject to <a href="http://snakesarelong.blogspot.com/2015/03/rattlesnake-roundups.html">read my previous article</a> and check out <a href="http://www.amazon.com/gp/product/1603440356/">this well-researched book</a>). But, increasing oversight by Texas wildlife agencies could allow them or others to monitor the effect of the harvest on rattlesnakes, which could lead to valuable insights into snake biology and help prevent <a href="http://www.bigcountryhomepage.com/news/sweetwater-jaycees-run-out-of-room-to-transport-snakes">economic and environmental disasters like this year's Sweetwater roundup</a>. This week's decision <a href="http://www.blogtalkradio.com/urbanjunglesradio/2016/05/23/ujr-episode-218--jackie-bibby-hacking-away-at-rattlesnake-deaths">inches us towards the hopeful possibility of a sustainable snake harvest</a> that could, over time, change the relationship between humans and western diamondbacks into a positive one, similar to our view of white-tailed deer, bobwhite quail, or largemouth bass. It's a non-traditional model for snake conservation, to be sure, but the efforts of the TPWD Snake Harvest Working Group combined with actions being taken by some <a href="http://www.blogtalkradio.com/urbanjunglesradio/2016/05/23/ujr-episode-218--jackie-bibby-hacking-away-at-rattlesnake-deaths">unlikely allies, such as roundup organizer Jackie Bibby</a>, will hopefully continue to move us towards a common goal of respectfully managing rattlesnakes as either game or non-game wildlife and not as pests. <i>[Edit: <a href="http://www.jstor.org/stable/4617307">An analysis from 2000</a> showed that </i><i>imposing size restrictions on rattlesnake harvests to individuals >90 cm in SVL (the size at maturity for most females) would earn hunters 19% more money.</i><i>]</i> The best part: we can help people in the process (<i>e.g., </i>by<i> </i>providing healthier products with stable prices, such as rattlesnake meat untainted with gasoline).</div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhJqUxE0I8jUA50d974hvr3BoXhllLEFnRa6GAeNXZMSm9TZGY-nOxwFHiMkShQiP2h31H-sPcYOJt5VRZmslXQhNQHJVgX5CfXw2DC5uUfm8hHh6RUq82y4GZH20T9nBpuwmfmZlRvtrW-/s1600/Dorcas+%2526+Willson+2013+Fig5.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="193" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhJqUxE0I8jUA50d974hvr3BoXhllLEFnRa6GAeNXZMSm9TZGY-nOxwFHiMkShQiP2h31H-sPcYOJt5VRZmslXQhNQHJVgX5CfXw2DC5uUfm8hHh6RUq82y4GZH20T9nBpuwmfmZlRvtrW-/s400/Dorcas+%2526+Willson+2013+Fig5.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Percentage of time radio-tracked Burmese Pythons spent<br />
fully concealed (black), partly visible (gray), and mostly visible (white).<br />
In nineteen 30-minute searches of a 30 x 25 m enclosure containing<br />
ten pythons, only two pythons were detected out of<br />
190 possible detection opportunities.<br />
From <a href="http://comp.uark.edu/~jwillson/publications/2013%20-%20Dorcas%20and%20Willson%20-%202013%20-%20%20Hidden%20Giants%20-%20Reptiles%20in%20Research.pdf">Dorcas & Willson 2013</a></td></tr>
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And—as if the irony weren't already thick enough—compare the above totals with the <a href="http://myfwc.com/news/news-releases/2016/february/27/python-awards/">~2000 lbs. of Burmese Python (106 snakes)</a> collected in Florida this year as part of an Florida Fish and Wildlife Conservation Commission-sponsored contest to control a snake whose populations actually do need to be "controlled" (despite the near-total impossibility of doing so). Among the several reasons for the difference include the lack of cultural inertia promoting snake hunting in Florida, <a href="http://comp.uark.edu/~jwillson/publications/2013%20-%20Dorcas%20and%20Willson%20-%202013%20-%20%20Hidden%20Giants%20-%20Reptiles%20in%20Research.pdf">the challenging habitat of the Everglades</a>, and the snakes' biology—pythons don't aggregate the way rattlesnakes do. If gassing is banned in Texas, <a href="http://www.smithsonianmag.com/smart-news/controversial-texas-rattlesnake-roundup-nets-largest-catch-date-180958575/">flushing rattlesnakes out of their hibernacula <i>en masse</i> will no longer be a legal hunting strategy.</a> Does this mean that rattlesnake roundup totals will become more like those of the Python Challenge? Not necessarily—<a href="http://tpwd.texas.gov/huntwild/wild/wildlife_diversity/nongame/media/TPWD-SHWG-References.pdf">the TPWD report references</a> alternative strategies already in use in other parts of the country that can still yield hundreds of pounds of rattlesnakes. Would a change in the hunting methods allowed have positive effects on snakes and other wildlife? Almost certainly. What would be the impacts on the roundup? I think it's worth pointing out that many former roundups, such as the Claxton Rattlesnake Festival in Claxton, Georgia, hosted by the <a href="http://www.evanscountywildlifeclub.com/">Evans County Wildlife Club</a>, and the <a href="http://www.wildchickenfestival.com/">Fitzgerald Wild Chicken Festival</a> in Fitzgerald, Georgia, still generate economic opportunity for their towns without collecting and killing wild snakes. I think it's quite likely that events like the Sweetwater Rattlesnake Roundup could continue to bring benefits to their communities without using gas to extract rattlesnakes from their dens.<br />
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ACKNOWLEDGMENTS</div>
<br />
Thanks to Ray Autry and Dale Burton from the <a href="https://www.facebook.com/groups/2799875358/">Rise Against Rattlesnake Roundups Facebook group</a> for pointing me to some resources about the 2016 Sweetwater Roundup.<br />
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REFERENCES</div>
<br />
<span style="font-size: x-small;">Adams, C.E. and J.K. Thomas. 2008. Texas Rattlesnake Roundups. Texas A&M University Press, College Station, Texas <<a href="http://www.amazon.com/gp/product/1603440356/">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Arena, P. C., C. Warwick, and D. Duvall. 1995. Rattlesnake Round-ups. Pages 313-324 in R. L. Knight and K. Gutzwiller, editors. Wildlife and Recreationists. Island Press, Washington, DC <<a href="https://books.google.de/books?id=BRbBAvLwQlAC&pg=PA313&lpg=PA313&dq=Wildlife+and+Recreationists:+Coexistence+Through+Management+And+Research+Rattlesnake+Round-ups&source=bl&ots=tMyVRTCzCG&sig=xc9PqfJCF5FbtzTV37bXf_KeZfA&hl=en&sa=X&ved=0ahUKEwjBtfudxP3MAhUGBSwKHRsHBOMQ6AEIKTAB#v=onepage&q=Wildlife%20and%20Recreationists%3A%20Coexistence%20Through%20Management%20And%20Research%20Rattlesnake%20Round-ups&f=false">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Campbell, J. A., D. R. Formanowicz Jr, and E. D. Brodie Jr. 1989. Potential impact of rattlesnake roundups on natural populations. Texas Journal of Science 41:301-317.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Dorcas, M. E., and J. D. Willson. 2013. Hidden giants: problems associated with studying secretive invasive pythons. Pages 367-385 in W. I. Lutterschmidt, editor. Reptiles in Research. Nova Biomedical, New York, New York <<a href="http://comp.uark.edu/~jwillson/publications/2013%20-%20Dorcas%20and%20Willson%20-%202013%20-%20%20Hidden%20Giants%20-%20Reptiles%20in%20Research.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Elliott, W. R. 2000. Conservation of the North American cave and karst biota. Pages 665-689 in H. Wilkens, D. Culver, and W. Humphreys, editors. Subterranean Ecosystems. Elsevier, Amsterdam.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Fitzgerald, L.A. and C.W. Painter. 2000. Rattlesnake commercialization: Long-term trends, issues, and implications for conservation. Wildlife Society Bulletin 28:235-253 <<a href="http://www.jstor.org/stable/4617307">link</a>> </span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Jackley, A. M. 1939. Rattlesnake Control and Conservation. South Dakota Conservation Digest 6:11.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Margres, M. J., J. J. McGivern, M. Seavy, K. P. Wray, J. Facente, and D. R. Rokyta. 2015. Contrasting modes and tempos of venom expression evolution in two snake species. Genetics 199:165-176 <<a href="http://www.genetics.org/content/199/1/165">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Reinert, H., and R. Rupert. 1999. Impacts of translocation on behavior and survival of Timber Rattlesnakes, <i>Crotalus horridus</i>. Journal of Herpetology 33:45-61 <<a href="https://www.jstor.org/stable/1565542?seq=1#page_scan_tab_contents">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Texas Parks and Wildlife Department. 2016. Snake Harvest Working Group Final Report <<a href="http://tpwd.texas.gov/huntwild/wild/wildlife_diversity/nongame/media/TPWD-SHWG-Report.pdf">link</a>> <<a href="http://tpwd.texas.gov/huntwild/wild/wildlife_diversity/nongame/media/TPWD-SHWG-References.pdf">references</a>> <<a href="http://tpwd.texas.gov/huntwild/wild/wildlife_diversity/nongame/media/TPWD-SHWG-Executive-Summary.pdf">summary</a>></span><br />
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="http://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
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Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com2tag:blogger.com,1999:blog-7443075087825368900.post-7115491221075668692016-04-26T08:00:00.000-06:002016-04-26T08:00:21.607-06:00Even snakes have their charismatic megafauna<div style="text-align: center;">
<span style="font-size: x-small;">This post will soon become available in Spanish.</span></div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjC9drNDHfrEaqeQ94O_1pYMjTJ5U-XX4plFbLOsX_CiKzmT6G0EDjFxOqVJsce67zx_PmBQUVa_HMbWR0GT7m6_l88n3phLLbFjdju4iraE-SGbGcHHqZ2O29fpChp3UlD6PZ2a0llZd_-/s1600/Bitis+harenna.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjC9drNDHfrEaqeQ94O_1pYMjTJ5U-XX4plFbLOsX_CiKzmT6G0EDjFxOqVJsce67zx_PmBQUVa_HMbWR0GT7m6_l88n3phLLbFjdju4iraE-SGbGcHHqZ2O29fpChp3UlD6PZ2a0llZd_-/s320/Bitis+harenna.jpg" width="296" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Bitis harenna</i><br />
From <a href="http://mapress.com/j/zt/article/view/zootaxa.4093.1.3">Gower et al. 2016</a></td></tr>
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Last year, I wrote about <a href="http://snakesarelong.blogspot.com/2014/11/the-9999th-reptile.html">the 10,000th reptile and the 3,500th snake</a> species to be described by scientists. The pace has not slowed down—as of <a href="http://www.reptile-database.org/db-info/news.html">its most recent update last week</a>, The Reptile Database currently lists <a href="http://reptile-database.reptarium.cz/advanced_search?taxon=serpentes&submit=Search">3,596 species of snakes</a> out of a total of <a href="http://reptile-database.reptarium.cz/advanced_search?submit=Search">10,391 species of (non-avian) reptiles</a>. A few weeks ago, the March 21st issue of the frequently-published journal <a href="http://www.mapress.com/j/zt/issue/archive"><i>Zootaxa </i>(volume 4093, issue 1)</a> included descriptions of three of these new snake species. What's interesting is that I initially looked this issue up because I saw one of them being shared a lot on social media—a new large species of viper. The other two, a pipesnake and a blindsnake, hadn't received as much attention. <a href="https://twitter.com/Zootaxa">Zootaxa tweets</a> all of their new species, and an examination of their feed shows that the <a href="https://twitter.com/Zootaxa/status/711709094042673152">viper tweet</a> received 4 retweets and 2 likes, whereas the <a href="https://twitter.com/Zootaxa/status/711708926350262272">pipesnake</a> and the <a href="https://twitter.com/Zootaxa/status/711709623980437504">blindsnake</a> received 2 retweets and one like each (even though the pipesnake had a photo<a href="#1" name="top1"><sup>1</sup></a> and was on the cover). Even though that's a small sample size, I think it's telling that even snakes have their <a href="https://en.wikipedia.org/wiki/Charismatic_megafauna">charismatic megafauna</a>.<br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjCFQRhwRZp5PHebaXCGnxDE9BamurI0Zh31kGP5VppUMuvS-vOQdZPQl84tWTV7EiuJB-V1b3AYVxHgoVk-qIrAWL_1Nm_1QQHqVOcFiDHdybhOwEwqe4Boa1SMoRrj4DtPdq1zY2Jxp51/s1600/bongo%2526tiger.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="400" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjCFQRhwRZp5PHebaXCGnxDE9BamurI0Zh31kGP5VppUMuvS-vOQdZPQl84tWTV7EiuJB-V1b3AYVxHgoVk-qIrAWL_1Nm_1QQHqVOcFiDHdybhOwEwqe4Boa1SMoRrj4DtPdq1zY2Jxp51/s400/bongo%2526tiger.png" width="272" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">A bongo (<i>Tragelaphus eurycerus</i>, top)<br />
and a tiger (<i>Panthera tigris</i>, bottom).<br />
You only needed a caption for one</td></tr>
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It seems backwards, in a way, that the dangerously venomous viper should be more popular than the innocuous pipesnake. <a href="https://conservationbytes.com/2015/10/13/only-thing-worse-than-being-labelled-deadly-is-not-being-called-anything-at-all/">One conservation blogger, Corey Bradshaw, put it nicely</a> by saying that "the only thing worse than being labelled deadly is not being called anything at all". Bradshaw pointed out that drawing attention to the potential for a species to cause harm to humans is not necessarily bad for the species in question. Even though snake biologists often decry these claims as exaggerated (usually because they are), Bradshaw wondered whether they are really very harmful. He suggested that people are generally more fascinated with animals that could kill us (even if they rarely do) than they are with entire groups of benign species, such as skinks or plethodontid salamanders, which are often considered boring (if a person is even aware of their existence). Compare tigers with, say, bongos. Both are critically endangered, inarguably gorgeous animals from exotic places. Tigers sometimes kill and eat people. Everyone knows a tiger. Most people think a bongo is a drum. Or, if you want a snake example, take rattlesnakes. <a href="http://www.greatseal.com/symbols/rattlesnake.html">Rattlesnakes are the Bald Eagles of snakes</a>. They are distinctly North American. Everybody in North America knows them. <a href="http://snakesarelong.blogspot.com/2013/11/how-snakes-see-through-closed-eyes.html">One was on our flag</a>. In contrast, the USA has never had a Smooth Greensnake (<i>Opheodrys vernalis</i>) on its flag, even though they are beautiful and North American and eat spiders. Perhaps the idea that any publicity is good publicity applies to conservation as well. Then again, perhaps not—<a href="http://www.bostonmagazine.com/news/blog/2016/02/23/rattlesnakes-in-the-quabbin-reservoir/">many residents of Massachusetts are needlessly worried about a Timber Rattlesnake reintroduction plan</a> on an island in the Quabbin Reservoir, probably in part because of the bad PR that rattlesnakes get on a regular basis. If the Massachusetts Division of Fisheries & Wildlife were reintroducing Smooth Greensnakes, I doubt that most people would care (and it certainly wouldn't have been the subject of <a href="http://pressblog.uchicago.edu/2016/03/30/ted-levin-on.html">such venomous debate in the media</a>). Indeed, Illinois's Lincoln Park Zoo is <a href="http://www.lpzoo.org/conservation-science/projects/restoring-smooth-green-snake">reintroducing Smooth Greensnakes in Chicago</a>, and nobody is writing letters to the editor about it (and, in a way, that's a shame, because it's an interesting and worthwhile effort).<br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi1vSmcMCuKfnnu2_EzSnduTVNzEkAjKL3AHzOh7outNIpxYpD32G9N06Sbf7p8c01J6vVQ5X_jTSYJYXc8xm9P6xdIljEfUQw3QZ5dh2wdOiGflLK6pyE7vOJvV1JrPqBnaHc4MJhuf2VY/s1600/Letheobia+mbeerensis+Malonza+et+al+2016.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi1vSmcMCuKfnnu2_EzSnduTVNzEkAjKL3AHzOh7outNIpxYpD32G9N06Sbf7p8c01J6vVQ5X_jTSYJYXc8xm9P6xdIljEfUQw3QZ5dh2wdOiGflLK6pyE7vOJvV1JrPqBnaHc4MJhuf2VY/s320/Letheobia+mbeerensis+Malonza+et+al+2016.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Letheobia mbeerensis</i><br />
From <a href="http://mapress.com/j/zt/article/view/zootaxa.4093.1.10">Malonza et al. 2016</a></td></tr>
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Anyway, I wanted to give some well-deserved press to the two less-publicized new snakes. The blindsnake, <i>Letheobia mbeerensis</i>, is pink with tiny, barely visible eyes. It was described from a single specimen collected southeast of Mt. Kenya in April of 2014 by a local farmer, who found it while tilling his fields. This person, whose name was not known to the scientists who wrote the article, made a considerable effort to get the snake identified—he traveled 125 miles from Siakago to Nairobi, where he gave the specimen to the Nairobi Snake Park, who forwarded it to herpetologists at the National Museums of Kenya. It is unique in having a relatively long tail (for a blindsnake), and in being found in a moist inland savanna. The other two Kenyan species of <i>Letheobia, </i>one of which was just described in 2007, are found in coastal lowlands with sandy soils. It is the 24th species of blindsnake known from Kenya, but I can guarantee that it won't be the last.</div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi0HeqJjeWL-_b1tBhGbTbC1GwmWkBjv5yBbs9_NfEMaK4DbEhLHDvmMs8HrJhTXfql-tYBJTBQ0wsF6TUsLh_KlCMwoi3s2pAK6rD-jQZIptRIJaWcaD_QrajlpBJ3mHtFUQAXnLU7I_Wn/s1600/Cylindrophis+ruffus+sensu+historico+Historical+drawings+Kieckbusch+et+al+2016+Zootaxa.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="640" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi0HeqJjeWL-_b1tBhGbTbC1GwmWkBjv5yBbs9_NfEMaK4DbEhLHDvmMs8HrJhTXfql-tYBJTBQ0wsF6TUsLh_KlCMwoi3s2pAK6rD-jQZIptRIJaWcaD_QrajlpBJ3mHtFUQAXnLU7I_Wn/s640/Cylindrophis+ruffus+sensu+historico+Historical+drawings+Kieckbusch+et+al+2016+Zootaxa.png" width="403" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><span style="font-size: 12.8px; text-align: left;">Historical drawings of <i>Cylindrophis ruffus</i><br />Illustrations A-C from <a href="https://books.google.com/books?id=E3vLU5r5HikC&printsec=frontcover&source=gbs_ge_summary_r&cad=0#v=onepage&q&f=false">Scheuchzer 1735</a><br />D-E from </span><span style="font-size: 12.8px; text-align: left;"><a href="http://www.biodiversitylibrary.org/page/13956829#page/319/mode/1up">Seba 1735</a><br />From <a href="http://mapress.com/j/zt/article/view/zootaxa.4093.1.1">Kieckbusch et al. 2016</a></span></td></tr>
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The story of the new pipesnake is even more interesting, and I suspect the paper in which it is described will ultimately be the most read and most cited of the three snake papers in this issue. This is because, in addition to describing the new species, it contains "an overview of the tangled taxonomic history of <i>C[ylindrophis] ruffus</i>", a widespread species <a href="http://reptile-database.reptarium.cz/species?genus=Cylindrophis&species=ruffus">commonly known as the Red-tailed or Common Pipe or Cylinder Snake</a>. The fourteen species of Asian Pipesnakes (<a href="http://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">family Cylindrophiidae</a>) are secretive and semifossorial snakes with small eyes, bodies that barely taper at all, and <a href="http://snakesarelong.blogspot.com/2012/11/identifying-snake-sheds-part-iii.html">ventral scales</a> only slightly larger than or equal in size to their dorsal scales. Many have contrasting light and dark ventral blotching with conspicuous bright coloration on the underside of their short tail, which they expose when threatened. Scientific knowledge of these snakes predates modern biological nomenclature. One is pictured in <a href="http://snakesarelong.blogspot.com/2015/05/linnaean-snakes-part-i.html">Albertus Seba's <i>Thesaurus</i>, which was one of Linnaeus's main sources</a>, although Linnaeus didn't include <i>C. ruffus</i> in either the 1758 or the 1766 edition of his <i>Systema Naturae—</i>instead, its first post-Linnaean description was written by Laurenti in 1768. Compared with other <i>Cylindrophis</i>, <i>C. ruffus </i>has a much larger distribution than any other species of Asian pipesnake. It's one of those species that is really a species complex—a group of closely related species that are very similar in appearance, to the point that the boundaries between them are often unclear. Other well-known examples include <a href="https://www.researchgate.net/profile/Martin_Villet/publication/49626436_Molecular_systematics_of_the_African_snake_family_Lamprophiidae_Fitzinger_1843_Serpentes_Elapoidea_with_particular_focus_on_the_genera_Lamprophis_Fitzinger_1843_and_Mehelya_Csiki_1903/links/00b7d532147848afc0000000.pdf">African House Snakes</a> (<i>Boaedon fuliginosus</i>, formerly <i>Lamprophis fuliginosus</i>) and <a href="https://www.researchgate.net/profile/Robert_Bryson_Jr/publication/259319878_Coalescent_Species_Delimitation_in_Milksnakes_(Genus_Lampropeltis)_and_Impacts_on_Phylogenetic_Comparative_Analyses/links/553817f10cf226723ab615b4.pdf">American Milksnakes</a> (<i>Lampropeltis triangulum</i>). Often unusual populations of these species are described as separate species, but without extensive rangewide sampling it's easy to miss more subtle, clinal variation, especially when that variation is genetic rather than morphological. <a href="https://www.researchgate.net/profile/Robert_Bryson_Jr/publication/259319878_Coalescent_Species_Delimitation_in_Milksnakes_(Genus_Lampropeltis)_and_Impacts_on_Phylogenetic_Comparative_Analyses/links/553817f10cf226723ab615b4.pdf">A recent revision of milksnakes</a> split this wide-ranging species into several, and researchers have been working on African House Snakes as well. But no one has really examined Red-tailed Pipesnakes. Last year, <a href="http://www.amphibian-reptile-conservation.org/pdfs/Volume/Vol_9_no_1/ARC_9_1_[General_Section]_34-51_e98_high_res.pdf">a group of European and Indonesian researchers examined a large number of <i>Cylindrophis </i>museum specimens</a> and discovered several specimens which did not fit any recognized species. But many of these specimens are old and some of their locations are uncertain. We don't have a lot of molecular data, and we have no specimens at all from many areas. And, no one has yet carried out a totally comprehensive review of the species complex (which really should encompass the entire genus, since the milksnake researchers found that some "milksnakes" were actually more closely related to mountain kingsnakes than they were to other milksnakes).<br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjuId7gG1jkE1zbcXdfmRxjKIR9UDDWhPNZFR1-enq7i6VRXbpr-BfLEBolTcSoQi2MZTmhngl1IVLgzwp9vAZl0BtV20m882YFnF4BxlayqrYyrDE-TFpeSHlRwNALdXZewN25h3lOSMCs/s1600/Cylindrophis_ruffus_02.JPG" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjuId7gG1jkE1zbcXdfmRxjKIR9UDDWhPNZFR1-enq7i6VRXbpr-BfLEBolTcSoQi2MZTmhngl1IVLgzwp9vAZl0BtV20m882YFnF4BxlayqrYyrDE-TFpeSHlRwNALdXZewN25h3lOSMCs/s320/Cylindrophis_ruffus_02.JPG" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Cylindrophis ruffus</i> raising its tail "flag"</td></tr>
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Despite its <a href="http://www.amphibian-reptile-conservation.org/pdfs/Volume/Vol_9_no_1/ARC_9_1_[General_Section]_34-51_e98_high_res.pdf">re-description in 2015</a>, <i>Cylindrophis ruffus</i> is still a species complex that suffers from a lot of complexity. Its morphology is highly variable. Its geographic range limits are unsettled. There is no <a href="https://en.wikipedia.org/wiki/Type_(biology)">type specimen</a>. The original type locality (“Surinami”) is a hemisphere away, obviously an error, which complicates decisions about which populations of <i>C. ruffus</i> should get to keep that name and which should change. The 2015 paper, as the authors of this month's paper delicately put it, "contain[s] some inaccuracies, including descriptive errors, which unfortunately increase the complexity of an already intricate taxonomic situation". The researchers state that they are currently undertaking the kind of comprehensive review that I called for above, but that in the process they discovered a morphologically distinct population from central Java, which they describe as <i>Cylindrophis subocularis</i> in this paper. But the real value of this paper, in my mind, is the step-by-step description of the history of this snake, starting with its first depiction in 1735 and continuing to present day. I'll leave the gory details for those who are really interested (the full-text is available <a href="https://www.researchgate.net/profile/Max_Kieckbusch/publication/299204673_An_inconspicuous_conspicuous_new_species_of_Asian_pipesnake_genus_Cylindrophis_Reptilia_Squamata_Cylindrophiidae_from_the_south_coast_of_Jawa_Tengah_Java_Indonesia_and_an_overview_of_the_tangled_taxon/links/56efb4f708ae4b8b5e757883.pdf">here</a>), but suffice it to say that the story of <i>Cylindrophis ruffus</i> is much more interesting than I ever knew (it took almost 100 years to get the geography right), and far from over.<br />
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<hr width="80%"><p><span class="Apple-style-span" style="font-size: x-small;"><br />
<a name="1"><b>1 </b></a>Granted, it was a photo of a preserved specimen.<a href="#top1"><sup>↩</sup></a><br />
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ACKNOWLEDGMENTS</div>
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Thanks to <a href="https://www.flickr.com/photos/mamuin/">M. A. Muin</a>, <a href="https://www.flickr.com/photos/zooeurope/9691133235/in/photolist-fLnAh4-7wCCxt-51hC7c-7mVa4w-bFSzme-5WcHjN-nmSDza-4RKtqy-6cY9tE-rnCGLE-nmSEdJ-7xPjL3-4jLck-6hLDs7-7mRgNc-75EgWL-6p61n6-bBCuPd-2418Av-2f4xxs-72cQd7-89jdhV-75Eg2G-fLpXs4-89nCFw-Amyu-6waj1U-6djoYM-89nrXj-9yPUM2-ecCoVq-4J1w11-Gn6ka-bQxbdv-58pq9z-fLEbv1-6w66Pe-5w6xjb-Gn2B6-asdvbE-EzYEt-Gn53X-mQAQ67-GnatT-ecwLEt-rGm1J-GmYAy-5xvJ5K-bWzWEZ-kzLez">Nigel Swales</a> and <a href="https://www.flickr.com/photos/marcusmeissner/5543057449/in/photolist-9rPBS2-7oivEY-8UGT3t-8iuh5V-8PxPkJ-8gRKrX-5rB7vW-dxew6o-qmzFxT-4HgkbG-8bnXFS-4xzJLY-6tYCzN-4yheCD-rbzGEG-pfo5h2-eXH6nK-6DQ83-abZ4St-eeBZgY-dx97Wv-4DYCGE-3wqLvJ-8MqmBm-n1spQ-PDwsf-8uPntX-7dphPX-6q6B35-4iXi1-8Rp3rv-7dtaMf-eHqCfN-4DYCB3-uYMUA-8TRwLu-7AttvU-9rPCL6-6dMqVH-af4FW4-KpkFJ-at1ik5-8wUbEr-asXExF-9rPCiP-6jjQtN-mr7DP-FRJpw-5gbonT-7rr1mv">Marcus Meissner</a> for the use of their photos.</div>
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REFERENCES</div>
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<span style="font-size: x-small;">Amarasinghe, A. A. T., P. D. Campbell, J. Hallermann, I. Sidik, J. Supriatna, and I. Ineich. 2015. Two new species of the genus <i>Cylindrophis </i>Wagler, 1828 (Squamata: Cylindrophiidae) from Southeast Asia. Amphibian and Reptile Conservation 9:34-51 <<a href="http://www.amphibian-reptile-conservation.org/pdfs/Volume/Vol_9_no_1/ARC_9_1_[General_Section]_34-51_e98_high_res.pdf">link</a>></span><br />
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<span style="font-size: x-small;">Gower, D. J., E. O. Z. Wade, S. Spawls, W. Böhme, E. R. Buechley, D. Sykes, and T. J. Colston. 2016. A new large species of <i>Bitis </i>Gray, 1842 (Serpentes: Viperidae) from the Bale Mountains of Ethiopia. Zootaxa 4093:41-63 <<a href="http://mapress.com/j/zt/article/view/zootaxa.4093.1.3">link</a>></span></div>
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<span style="font-size: x-small;">Kieckbusch, M., S. Mecke, L. Hartmann, L. Ehrmantraut, M. O’Shea, and H. Kaiser. 2016. An inconspicuous, conspicuous new species of Asian pipesnake, genus <i>Cylindrophis </i>(Reptilia: Squamata: Cylindrophiidae), from the south coast of Jawa Tengah, Java, Indonesia, and an overview of the tangled taxonomic history of <i>C. ruffus</i> (Laurenti, 1768). Zootaxa 4093:1-25 <<a href="http://mapress.com/j/zt/article/view/zootaxa.4093.1.1">link</a>></span></div>
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<span style="font-size: x-small;">Malonza, P. K., A. M. Bauer, and J. M. Ngwava. 2016. A new species of <i>Letheobia </i>(Serpentes: Typhlopidae) from central Kenya. Zootaxa 4093:143-150 <<a href="http://mapress.com/j/zt/article/view/zootaxa.4093.1.10">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Scheuchzer, J. J. 1735. Physica Sacra Iconibus Anaeis Illustrata, Procurante & Sumtus Suppeditante. Tomus IV. Augustae Vindelicorum et Ulmae, Ulm <<a href="https://books.google.com/books?id=E3vLU5r5HikC&printsec=frontcover&source=gbs_ge_summary_r&cad=0#v=onepage&q&f=false">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Seba, A. 1734-1765. Locupletissimi rerum naturalium thesauri accurata descriptio, et iconibus artificiosissimis expressio, per universam physices historiam :opus, cui, in hoc rerum genere, nullum par exstitit. Apud Janssonio-Waesbergios & J. Wetstenium & Gul. Smith, Amstelaedami <<a href="http://www.biodiversitylibrary.org/page/13956829#page/319/mode/1up">link</a>></span></div>
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<a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license"><span style="font-size: x-small;"><img alt="Creative Commons License" src="http://i.creativecommons.org/l/by-nc-nd/3.0/80x15.png" style="border-width: 0px;" /></span></a></div>
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="http://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com12tag:blogger.com,1999:blog-7443075087825368900.post-56254212470822876372016-03-28T22:22:00.000-06:002017-03-18T10:33:33.252-06:00State Snakes, Linnaean Names, and Other Recent Updates<div style="text-align: justify;">
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<span style="font-size: x-small;"><a href="https://drive.google.com/file/d/0B2jxDXUJh_X7XzZ4SEJJWjlGNmM/view?usp=sharing">Click here to read in Spanish</a></span></div>
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<span style="font-size: x-small;"><a href="https://drive.google.com/file/d/0B2jxDXUJh_X7XzZ4SEJJWjlGNmM/view?usp=sharing">Haga clic aquí para leer en español</a></span></div>
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<a href="http://snakesarelong.blogspot.com/2015/12/life-is-short-but-snakes-are-long-2015.html">As I wrote in December</a>, the demands of completing my dissertation (and my new position as <a href="http://upr.org/term/andrew-durso">a science reporter with Utah Public Radio</a>) haven't left me enough time to write <a href="http://snakesarelong.blogspot.com/2014/10/how-to-teach-yourself-about-obscure.html">the more in-depth long-form content</a> that I (and readers, it seems) like so much. If all goes according to plan, I should return to those more elaborate articles towards the end of 2016, but in the meantime I wanted to highlight some recent and exciting updates to some of my older articles.</div>
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<b style="text-align: justify;"><a href="http://snakesarelong.blogspot.com/2013/08/what-state-snakes-should-be-part-ii.html">What the State Snakes Should Be</a></b><br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgp690v85_atErUuOhrWnpLqThLWlxPyK_8KjeIqRlNcP-VpUaHNu0FdYOB_7OVZ8W4h3lmtMh7TMaZGrfU1Ot7IrEebWmC_dJ_f592dCIzxUKYjxOP5XejThpNH9mRLnzADxVUo2t53CI6/s1600/Thamnophis+sirtalis+eating+Anaxyrus+woodhousei.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgp690v85_atErUuOhrWnpLqThLWlxPyK_8KjeIqRlNcP-VpUaHNu0FdYOB_7OVZ8W4h3lmtMh7TMaZGrfU1Ot7IrEebWmC_dJ_f592dCIzxUKYjxOP5XejThpNH9mRLnzADxVUo2t53CI6/s1600/Thamnophis+sirtalis+eating+Anaxyrus+woodhousei.jpg" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">A Common Gartersnake (<i>Thamnophis sirtalis</i>)<br />
eats a Woodhouse's Toad (<i>Anaxyrus woodhousei</i>)</td></tr>
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In February the state of Virginia <a href="https://www.washingtonpost.com/local/virginia-politics/stinky-and-wimpy-but-its-ours-virginia-legislature-picks-a-state-snake/2016/02/25/767237e4-dbf6-11e5-891a-4ed04f4213e8_story.html?utm_content=bufferb8572&utm_medium=social&utm_source=facebook.com&utm_campaign=buffer">became the first state to officially designate a state snake</a>. They chose the Common Gartersnake (<i>Thamnophis sirtalis</i>), despite being literally one of only two states in the nation to share their name with a genus of snake! If they had read <a href="http://snakesarelong.blogspot.com/2013/08/what-state-snakes-should-be-part-ii.html">my 2013 article</a>, they might have gone with my pick of <i>Virginia valeriae</i>, the widespread Smooth Earthsnake, instead. But, perhaps there was already enough controversy: the gartersnake was proposed by 11-year-old Aiden Coleman of Williamsburg, but was put down by senators for being too wimpy. A couple of senators preferred the Timber Rattlesnake (<i>Crotalus horridus</i>), but the gartersnake was reinstated after Coleman asked one of them "just how much like West Virginia do you want us to be?"—unlike the legislators, Coleman already knew that the Timber Rattlesnake is the (very well-chosen) state reptile of West Virginia. The bill is now with the governor, whom some have suggested is the real snake.<br />
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<b><a href="http://snakesarelong.blogspot.com/2015/06/the-linnaean-snakes-part-ii.html">The Linnaean Snakes</a></b><br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjCP-3lbYQE3drI7HFGMltNU7nvZHk16awB7pECXGKI6tew9EtYAi6WvRJgk6hbSVOeUR5UpYGDmJN1vydEisu7yT85lwQTH4Wjq3hiyfsGSayPMoKXuPxd07b3vZ_LZZPu0W3vfMh2CMzi/s1600/4095.jpg" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="213" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjCP-3lbYQE3drI7HFGMltNU7nvZHk16awB7pECXGKI6tew9EtYAi6WvRJgk6hbSVOeUR5UpYGDmJN1vydEisu7yT85lwQTH4Wjq3hiyfsGSayPMoKXuPxd07b3vZ_LZZPu0W3vfMh2CMzi/s320/4095.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">An Eastern Ribbonsnake from the panhandle of Florida</td></tr>
</tbody></table>
Heads up, taxonomy buffs—the scientific name of the Eastern Ribbonsnake (currently <i>Thamnophis sauritus</i>) is probably about to change to <i>Thamnophis saurita</i>, for some fairly technical linguistic reasons. Linnaeus named both this species and <i>Thamnophis sirtalis</i>, but because Linnaeus's description for <i>sirtalis</i> better matched <i>sauritus</i>, the two names were for decades confusingly interchanged. All seemed to be settled by a 1956 ICZN ruling, but in March a new paper in the journal <i>Herpetological Review </i>pointed out that <i>Saurita</i>, the original spelling used by Linnaeus, was capitalized and that its –<i>a </i>ending did not match the masculine gender of his genus <i>Coluber</i>. According to the grammatical rules of species naming that Linnaeus followed and which we still follow, this means that he meant "<i>Saurita</i>" to be a noun, rather than an adjective, and so the ending should not change to match the gender of the genus. The common assertion that <a href="https://repositories.lib.utexas.edu/bitstream/handle/2152/45026/0099_Thamnophis_sauritus.pdf?sequence=1&isAllowed=y">"The specific name <i>sauritus </i>is New Latin, meaning lizardlike"</a> is incorrect: <i>sauros </i>is Greek, not Latin, and the suffix <i>–ita</i> does not mean "like", but "little" (in Spanish). An obscure 5th-Century Greek dictionary by the lexicographer Hesychius, which is famous for being the only remaining source for a lot of ancient Greek words and would have been available to Linnaeus, lists "<i>Saurita</i>" as "a kind of serpent", settling the issue.<br />
<br />
<b><a href="http://snakesarelong.blogspot.com/2013/11/the-truth-about-snakebite.html">The Truth About Snakebite</a></b><br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh_MtSO9JFAK_W5rJ5BQKnwoccIdPHaS7tozuJRlhJ9MEfGlqusSDsW5QI9e9F0BV6I9pu9a48FaJJMXFD5QSop1OS_0UtuMj8zivSgbfYO40fo5jbh25paCbQv4FF8_ZvAcix1Ric6-jwu/s1600/snakebite_infographic_crop.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="273" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEh_MtSO9JFAK_W5rJ5BQKnwoccIdPHaS7tozuJRlhJ9MEfGlqusSDsW5QI9e9F0BV6I9pu9a48FaJJMXFD5QSop1OS_0UtuMj8zivSgbfYO40fo5jbh25paCbQv4FF8_ZvAcix1Ric6-jwu/s320/snakebite_infographic_crop.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Close-up of part of <a href="https://www.etsy.com/listing/253283392/the-truth-about-venomous-snakebites-in?ref=shop_home_active_1">Liz Nixon's infographic</a></td></tr>
</tbody></table>
Fear of snakes <a href="http://www.nytimes.com/2016/03/24/opinion/terrorists-bathtubs-and-snakes.html">made the <i>New York Times</i> op-ed section this week</a> in an insightful article about the way humans assess the relative risks of terrorism and climate change. Although I completely agree with the article's point, in my opinion the author missed an opportunity to emphasize how our fear of snakes, like our fear of terrorism, is way beyond the risk posed by either (especially in the USA). It was a bit frustrating for me to read an article that came so close to making the analogy that we fear snakes even though they are unlikely to do us harm, but instead used fear of snakes as an example of an urgent fear distracting us from more gradual, but ultimately more dangerous threats. It's a tricky subject, but I did like the comparison between the number of deaths in the USA from falling in the bathtub (464/year) vs. from a terrorist attack (17/year)—both more likely than death from venomous snakebite (5/year). Also, if you haven't seen it, check out <a href="https://www.etsy.com/listing/253283392/the-truth-about-venomous-snakebites-in?ref=shop_home_active_1">the awesome infographic</a> that <a href="http://www.nixonmedicalmedia.com/portfolio/">scientific illustrator Liz Nixon</a> made using some of the data in my snakebite post.</div>
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<br />
<b><a href="http://snakesarelong.blogspot.com/2014/07/tetrodotoxin-resistant-snakes.html">Tetrodotoxin-resistant Snakes</a></b><br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhTlkwbiNvxShUCVTp51M4rbqDUH2nJ8aagoyvnOIbBNURM6g8Dij9KPa70lxc250roEe_6LWg1iyMqetaTtqxtY6KuPO3-fdGFXylxKDNvGr-DHJgf5Z_sV_XBBwDu6GMuxwAvogJadazI/s1600/Heterodon+platirhinos+eating+toad+JT+Sandow.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="189" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhTlkwbiNvxShUCVTp51M4rbqDUH2nJ8aagoyvnOIbBNURM6g8Dij9KPa70lxc250roEe_6LWg1iyMqetaTtqxtY6KuPO3-fdGFXylxKDNvGr-DHJgf5Z_sV_XBBwDu6GMuxwAvogJadazI/s320/Heterodon+platirhinos+eating+toad+JT+Sandow.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">An Eastern Hog-nosed Snake eats a toad</td></tr>
</tbody></table>
I rarely reference my own research on this blog, but last year I collaborated with Dr. Butch Brodie and members of his lab to <a href="https://www.researchgate.net/publication/281827008_Is_there_more_than_one_way_to_skin_a_newt_Convergent_toxin_resistance_in_snakes_is_not_due_to_a_common_genetic_mechanism">publish some data on tetrodotoxin resistance in hog-nosed snakes</a> (genus <i>Heterodon</i>). These snakes are well-known toad-eaters, but the few records of them eating newts were scattered until I brought them together in our new paper. Combined with molecular and whole-body resistance data, we showed that Eastern Hog-nosed Snakes from parts of upstate New York are more resistant to tetrodotoxin (TTX) than even the most resistant gartersnakes. But, Eastern Hog-nosed Snakes elsewhere are not as TTX-resistant, and Western Hog-nosed Snakes do not appear to be TTX-resistant at all. Most interesting, the mechanism of resistance appears to be something quite distinct from <a href="http://www.pnas.org/content/109/12/4556.full">the conserved mutations in gartersnakes and other newt-eating snakes</a>, and so far unknown.</div>
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ACKNOWLEDGMENTS</div>
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Thanks to <a href="http://ecosym.ecology.colostate.edu/gallery_contest2006.php">David Herasimtschuk</a>, <a href="https://www.flickr.com/photos/wildliferesourcesdivision/3919904830">Patti and Jack Sandow</a>, and <a href="https://www.flickr.com/photos/nclarkii/sets/72157630638940662/">Pierson Hill</a> for the use of their photos.</div>
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REFERENCES</div>
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<span style="font-size: x-small;">Feldman, C. R., E. D. Brodie, and M. E. Pfrender. 2012. Constraint shapes convergence in tetrodotoxin-resistant sodium channels of snakes. Proceedings of the National Academy of Sciences 106:13415-13420 <<a href="http://www.pnas.org/content/109/12/4556.full">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Feldman, C. R., A. M. Durso, C. T. Hanifin, M. E. Pfrender, P. K. Ducey, A. N. Stokes, K. E. Barnett, E. Brodie III, and E. Brodie Jr. 2016. Is there more than one way to skin a newt? North American snakes with convergent feeding adaptations do not share a common genetic mechanism. Heredity 116:84-91 <<a href="https://www.researchgate.net/publication/281827008_Is_there_more_than_one_way_to_skin_a_newt_Convergent_toxin_resistance_in_snakes_is_not_due_to_a_common_genetic_mechanism">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Kraus, F. and H. D. Cameron. 2016. A note on the proper nomenclature for the snake currently known as <i>Thamnophis sauritus</i>. Herpetological Review 47:74-75</span></div>
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<a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license"><span style="font-size: x-small;"><img alt="Creative Commons License" src="http://i.creativecommons.org/l/by-nc-nd/3.0/80x15.png" style="border-width: 0px;" /></span></a></div>
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="http://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com0tag:blogger.com,1999:blog-7443075087825368900.post-79882644764170409602016-02-23T08:00:00.000-07:002018-06-04T09:53:38.579-06:00Dragonsnakes and Filesnakes Revisited<div style="text-align: center;">
<span style="font-size: x-small;"><a href="https://drive.google.com/file/d/0B2jxDXUJh_X7T2JUZXVxZjhnWkk/view">Click here to read this post in Spanish</a></span><br />
<span style="font-size: x-small;"><a href="https://drive.google.com/file/d/0B2jxDXUJh_X7T2JUZXVxZjhnWkk/view">Haga clic aquí para leer este mensaje en español</a></span><br />
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I've written about both <a href="http://snakesarelong.blogspot.com/2014/08/filesnakes-wartsnakes-or-elephant.html">filesnakes</a> (family Acrochordidae) and <a href="http://snakesarelong.blogspot.com/2013/09/dragonsnakes.html">dragonsnakes</a> (part of the family Xenodermidae<a href="#1" name="top1"><sup>1</sup></a>) before. <a href="http://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">Traditional snake taxonomy</a> suggests that, although they <a href="http://www.timetree.org/search/pairwise/xenodermus/acrochordus">branch off from the main stem of the snake family tree at about the same time</a>, they're not very closely related. But, new evidence emphasizes the uniqueness of dragonsnakes and thickens the plot in the unfolding story of the evolution of snakes.<br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiNoY_0AuDcd-io1UC8CscMAYw1uQawd4nvXMGhmBhBegDUJ83tqhN5T_isrGFFP_LyxyfVNoasbbMZBRDC_hg5tjxrwmLcxse9k0WH4j7LFOpLi_5RT3jLJFwEYilzSA7bzAipdU6ejsnz/s1600/mt+vs+nuc+cladograms+Oguiura+et+al+2009.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="277" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiNoY_0AuDcd-io1UC8CscMAYw1uQawd4nvXMGhmBhBegDUJ83tqhN5T_isrGFFP_LyxyfVNoasbbMZBRDC_hg5tjxrwmLcxse9k0WH4j7LFOpLi_5RT3jLJFwEYilzSA7bzAipdU6ejsnz/s400/mt+vs+nuc+cladograms+Oguiura+et+al+2009.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Two hypotheses about the relationships of the major groups of snakes.<br />
Left: tree based on nuclear genes, showing Acrochordidae and Xenodermidae<br />
as successive outgroups to core Colubroidea<br />
Right: tree based on mitochondrial genes, showing a sister relationship<br />
between Acrochordidae and Xenodermidae<br />
From <a href="https://www.researchgate.net/profile/Nancy_Oguiura/publication/41848785_Cytogenetics_and_molecular_data_in_snakes_a_phylogenetic_approach/links/0fcfd5062ef29db787000000.pdf">Oguiura et al. 2009</a></td></tr>
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Most phylogenetic analyses are pretty consistent in classifying both filesnakes and dragonsnakes as caenophidians, or "advanced" snakes. But, they differ in their placement of dragonsnakes and other xenodermids, including the truly strange and obscure odd-scaled snakes (<i>Achalinus</i>), bearded snakes (<i>Fimbrios</i>), stream, earth, or red snakes (<i>Stoliczkia</i><a href="#2" name="top2"><sup>2</sup></a>), wood, mountain, or narrow-headed snakes (<i>Xylophis</i>), and a new genus, <a href="https://www.researchgate.net/profile/Minh_Le8/publication/273260157_A_new_genus_and_species_of_xenodermatid_snake_Squamata_Caenophidia_Xenodermatidae_from_northern_Lao_People's_Democratic_Republic/links/54fd1b4a0cf20700c5ea3f4c.pdf">just described in 2015</a> and still without a common name, <i>Parafimbrios. </i>Most analyses group xenodermids with the colubroids (<a href="http://snakesarelong.blogspot.com/2012/04/asymmetrical-snakes.html">pareids</a><a href="#1" name="top1"><sup>1</sup></a>, vipers, homalopsids, colubrids, lamprophiids, and elapids), albeit as the most basal branch. Many textbooks actually define Caenophidia as Colubroidea + Acrochordidae (aka Acrochordoidea), distinctly separating the colubroids from the filesnakes on the basis of shared, derived characteristics such as wide ventral scales, as well as features of the skull, hemipenes, and the muscles, cartilages, and arteries between the ribs. However, several recent trees based on DNA sequences suggest instead that filesnakes and dragonsnakes might be one another's closest living relatives.<br />
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<b><span style="font-size: 10.0pt;">Study<o:p></o:p></span></b></div>
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<td style="border-left: none; border: solid windowtext 1.0pt; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<b><span style="font-size: 10.0pt;">Acrochordid-Xenodermid Relationship<o:p></o:p></span></b></div>
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<td style="border-left: none; border: solid windowtext 1.0pt; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<b><span style="font-size: 10.0pt;">Support<o:p></o:p></span></b></div>
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<td style="border-left: none; border: solid windowtext 1.0pt; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<b><span style="font-size: 10.0pt;">How many species?<o:p></o:p></span></b></div>
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<td style="border-left: none; border: solid windowtext 1.0pt; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 115.8pt;" width="154"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<b><span style="font-size: 10.0pt;">What data were used?<o:p></o:p></span></b></div>
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<td style="border-top: none; border: solid windowtext 1.0pt; mso-border-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 107.75pt;" width="144"><div class="MsoNormal" style="margin-bottom: 0.0001pt;">
<span style="font-size: 10.0pt;"><a href="http://www.biodiversitylibrary.org/item/72743#page/190/mode/1up">Boulenger
1893</a><o:p></o:p></span></div>
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<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">X&A<o:p></o:p></span></div>
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<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">-<o:p></o:p></span></div>
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<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">-<o:p></o:p></span></div>
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<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 115.8pt;" width="154"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">Morphology<o:p></o:p></span></div>
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<td style="border-top: none; border: solid windowtext 1.0pt; mso-border-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 107.75pt;" width="144"><div class="MsoNormal" style="margin-bottom: 0.0001pt;">
<span style="font-size: 10.0pt;"><a href="https://www.researchgate.net/profile/Wesley_Brown3/publication/227763582_Phylogenetic_relationships_of_colubroid_snakes_based_on_mitochondrial_DNA_sequences/links/00463513f6af51ee2f000000.pdf">Kraus
& Brown 1998</a><o:p></o:p></span></div>
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<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">X+A<o:p></o:p></span></div>
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<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">8%<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">37<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 115.8pt;" width="154"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">ND4<o:p></o:p></span></div>
</td>
</tr>
<tr>
<td style="border-top: none; border: solid windowtext 1.0pt; mso-border-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 107.75pt;" width="144"><div class="MsoNormal" style="margin-bottom: 0.0001pt;">
<span style="font-size: 10.0pt;"><a href="http://www2.bren.ucsb.edu/~kendall/supertree/examples/Kelly2003SystBiol.pdf">Kelly
et al. 2003</a><o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">X+A<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">98%<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">98<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 115.8pt;" width="154"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">4
mitochondrial genes<o:p></o:p></span></div>
</td>
</tr>
<tr>
<td style="border-top: none; border: solid windowtext 1.0pt; mso-border-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 107.75pt;" width="144"><div class="MsoNormal" style="margin-bottom: 0.0001pt;">
<span style="font-size: 10.0pt;"><a href="https://www.infona.pl/resource/bwmeta1.element.elsevier-439e9ff0-fb59-358a-8586-b0410c04aa73">Vidal
et al. 2007</a><o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">A,X<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">not reported<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">25<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 115.8pt;" width="154"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">7
nuclear genes<o:p></o:p></span></div>
</td>
</tr>
<tr>
<td style="border-top: none; border: solid windowtext 1.0pt; mso-border-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 107.75pt;" width="144"><div class="MsoNormal" style="margin-bottom: 0.0001pt;">
<span style="font-size: 10.0pt;"><a href="http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.576.2548&rep=rep1&type=pdf">Wiens
et al. 2008</a><o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">A,X<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">>95%<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">50<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 115.8pt;" width="154"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">20
nuclear genes<o:p></o:p></span></div>
</td>
</tr>
<tr>
<td style="border-top: none; border: solid windowtext 1.0pt; mso-border-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 107.75pt;" width="144"><div class="MsoNormal" style="margin-bottom: 0.0001pt;">
<span style="font-size: 10.0pt;"><a href="https://www.researchgate.net/publication/225766738_The_complete_mitochondrial_DNA_sequence_and_the_phylogenetic_position_of_Achalinus_meiguensis_Reptilia_Squamata">Wang
et al. 2009</a><o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">A,X*<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">100%<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">30<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 115.8pt;" width="154"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">12
nuclear genes<o:p></o:p></span></div>
</td>
</tr>
<tr>
<td style="border-top: none; border: solid windowtext 1.0pt; mso-border-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 107.75pt;" width="144"><div class="MsoNormal" style="margin-bottom: 0.0001pt;">
<span style="font-size: 10.0pt;"><a href="http://www.scielo.br/pdf/paz/v49n11/a01v49n11.pdf">Zaher et al. 2009</a><o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">A,X<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">94%<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">131<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 115.8pt;" width="154"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">2
mitochondrial genes + 1 nuclear gene<o:p></o:p></span></div>
</td>
</tr>
<tr>
<td style="border-top: none; border: solid windowtext 1.0pt; mso-border-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 107.75pt;" width="144"><div class="MsoNormal" style="margin-bottom: 0.0001pt;">
<span style="font-size: 10.0pt;"><a href="https://www.researchgate.net/profile/Adrian_Nieto-Montes_de_Oca/publication/47790057_The_phylogeny_of_advanced_snakes_(Colubroidea)_with_discovery_of_a_new_subfamily_and_comparison_of_support_for_likelihood_trees/links/00b7d52a4709539ad7000000.pdf">Pyron
et al. 2011</a><o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">A,X<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">97-100%<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">761<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 115.8pt;" width="154"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">3
mitochondrial genes + 2 nuclear genes<o:p></o:p></span></div>
</td>
</tr>
<tr>
<td style="border-top: none; border: solid windowtext 1.0pt; mso-border-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 107.75pt;" width="144"><div class="MsoNormal" style="margin-bottom: 0.0001pt;">
<span style="font-size: 10.0pt;"><a href="http://people.earth.yale.edu/sites/default/files/files/Gauthier/Gauthier.pdf">Gauthier
et al. 2012</a><o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">X,A<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">95-100%<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">141
extant +<br />
51 extinct<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 115.8pt;" width="154"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">610
morphological characters<o:p></o:p></span></div>
</td>
</tr>
<tr>
<td style="border-top: none; border: solid windowtext 1.0pt; mso-border-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 107.75pt;" width="144"><div class="MsoNormal" style="margin-bottom: 0.0001pt;">
<span style="font-size: 10.0pt;"><a href="http://rsbl.royalsocietypublishing.org/content/8/6/1043.short">Wiens et
al. 2012</a><o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">A,X<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">100%<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">161<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 115.8pt;" width="154"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">44
nuclear genes<o:p></o:p></span></div>
</td>
</tr>
<tr>
<td style="border-top: none; border: solid windowtext 1.0pt; mso-border-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 107.75pt;" width="144"><div class="MsoNormal" style="margin-bottom: 0.0001pt;">
<span style="font-size: 10.0pt;"><a href="http://bmcevolbiol.biomedcentral.com/articles/10.1186/1471-2148-13-93">Pyron
et al. 2013</a><o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">X+A<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">95%<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">4,161<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 115.8pt;" width="154"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">5
mitochondrial genes + 7 nuclear genes<o:p></o:p></span></div>
</td>
</tr>
<tr>
<td style="border-top: none; border: solid windowtext 1.0pt; mso-border-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 107.75pt;" width="144"><div class="MsoNormal" style="margin-bottom: 0.0001pt;">
<span style="font-size: 10.0pt;"><a href="http://med.fsu.edu/userFiles/file/TSL/Genomics__Pyron_et_al,_Molecular_Phylogenetics_and_Evolution_(2014)_81_p221_231.pdf">Pyron
et al. 2014</a><o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">A,X*<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">99%<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">32<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 115.8pt;" width="154"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">333
nuclear loci<br />
with 100% coverage<o:p></o:p></span></div>
</td>
</tr>
<tr>
<td style="border-top: none; border: solid windowtext 1.0pt; mso-border-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 107.75pt;" width="144"><div class="MsoNormal" style="margin-bottom: 0.0001pt;">
<span style="font-size: 10.0pt;"><a href="http://www.wienslab.com/Publications_files/Zheng_Wiens_2015b_MPE.pdf">Zheng
& Wiens 2016</a><o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">A,X<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">91%<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 77.9pt;" width="104"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">4,162<o:p></o:p></span></div>
</td>
<td style="border-bottom: solid windowtext 1.0pt; border-left: none; border-right: solid windowtext 1.0pt; border-top: none; mso-border-alt: solid windowtext .5pt; mso-border-left-alt: solid windowtext .5pt; mso-border-top-alt: solid windowtext .5pt; padding: 0in 5.4pt 0in 5.4pt; width: 115.8pt;" width="154"><div align="center" class="MsoNormal" style="margin-bottom: 0.0001pt; text-align: center;">
<span style="font-size: 10.0pt;">5
mitochondrial genes + 47 nuclear genes<o:p></o:p></span></div>
</td>
</tr>
</tbody></table>
<div style="text-align: center;">
<span style="font-size: x-small;">A selection of studies that have examined the relationship between acrochordids and xenodermids.</span></div>
<div style="text-align: center;">
<span style="font-size: x-small;">X+A means that the two are each other's closest relatives; A,X means that acrochordids are more distantly</span><br />
<span style="font-size: x-small;">related to colubroids than xenodermids; X,A means that xenodermids are more distant</span><br />
<span style="font-size: x-small;">*Relationships differed depending on which methods were used</span></div>
<div style="text-align: center;">
<span style="text-align: left;"></span><br />
<span style="text-align: left;"></span>
<br />
<div style="text-align: justify;">
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiwQ_o_g9rBsqrnXAY8tAJ7V_T6i9lTPOGNUiSixGgbD0N8j0lOxpF_MzUoBLt9byTNwCCGkJkBxhxMPMwdN84n9mEAqS04VFT3juX_W72t3hyRSVyHYgUGgMc2hNP_svXgeMUv93Sz3lTo/s1600/Acrochordus+arafurae+Stephen+Zozaya.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="205" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiwQ_o_g9rBsqrnXAY8tAJ7V_T6i9lTPOGNUiSixGgbD0N8j0lOxpF_MzUoBLt9byTNwCCGkJkBxhxMPMwdN84n9mEAqS04VFT3juX_W72t3hyRSVyHYgUGgMc2hNP_svXgeMUv93Sz3lTo/s320/Acrochordus+arafurae+Stephen+Zozaya.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Arafura Filesnake (<i>Acrochordus arafurae</i>)</td></tr>
</tbody></table>
For example, <a href="https://www.researchgate.net/profile/Wesley_Brown3/publication/227763582_Phylogenetic_relationships_of_colubroid_snakes_based_on_mitochondrial_DNA_sequences/links/00463513f6af51ee2f000000.pdf">the first study to use DNA</a> to examine the relationships of these two groups of snakes found some support for each hypothesis, concluding that the "potential sister-taxon relationship of acrochordids and xenodermines [is] a reasonable hypothesis requiring future testing." In 2003, <a href="http://www2.bren.ucsb.edu/~kendall/supertree/examples/Kelly2003SystBiol.pdf">data from three more mitochondrial genes</a> resulted in the same relationship, causing the authors to suggest that xenodermids should be excluded from Colubroidea. However, since that time, numerous studies have not repeated this result. <span style="text-align: left;"><a href="http://www.scielo.br/pdf/paz/v49n11/a01v49n11.pdf">In 2009, one research group</a> predicted that "these differences...are due to taxonomic sampling issues", predicting that as DNA was collected from more species of snakes, the basal position of <i>Acrochordus </i>would be confirmed.</span><br />
<br />
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiZj4VwuxjtWg75dNOVbNZRyoEnF5XOGzRnt80jW_689O16YOyvdbzr759WWCqwywCZZpHDsghQFbeB1NgNIhbNhZuOjX76UEtNA8uig0vWrc3CWeKo9K34vzj6kbMnOvbd807VJWfuBDMZ/s1600/Xenodermus+javanicus+Konrad+Mebert+2.jpg" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="197" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiZj4VwuxjtWg75dNOVbNZRyoEnF5XOGzRnt80jW_689O16YOyvdbzr759WWCqwywCZZpHDsghQFbeB1NgNIhbNhZuOjX76UEtNA8uig0vWrc3CWeKo9K34vzj6kbMnOvbd807VJWfuBDMZ/s320/Xenodermus+javanicus+Konrad+Mebert+2.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Dragonsnake (<i>Xenodermus javanicus</i>)</td></tr>
</tbody></table>
<span style="text-align: left;">So, it was a real surprise when <a href="http://bmcevolbiol.biomedcentral.com/articles/10.1186/1471-2148-13-93">a 2013 analysis, the largest yet,</a> including samples from 80% of all snake genera, placed Acrochordidae and Xenodermidae as sister groups. Neither <a href="http://www.wienslab.com/Publications_files/Zheng_Wiens_2015b_MPE.pdf">a follow-up analysis</a> combining that dataset with one containing data from many more genes nor <a href="http://med.fsu.edu/userFiles/file/TSL/Genomics__Pyron_et_al,_Molecular_Phylogenetics_and_Evolution_(2014)_81_p221_231.pdf">an analysis using only the most complete data</a> have settled the issue. The latter study compared several methods for generating phylogenetic trees and found that the relationship between acrochordids and xenodermids depended a lot on which methods were used. Part of the problem is that, even if they are each others' closest relatives, they still diverged between 70 and 80 million years ago, making them susceptible to a problem in phylogenetics known as <a href="https://en.wikipedia.org/wiki/Long_branch_attraction">long-branch attraction</a>, which happens when the amount of evolutionary change within a lineage causes that lineage to appear similar (and thus closely related) to another long-branched lineage, solely because they have both undergone a lot of change, rather than because they are actually related.</span><br />
<br />
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhGyWvoGi6bnnZE5SbgfAmMWTFLN3Q21TQobyMEi4388keHuOitH_umQ0HquAHI4AkjHie4l2pnE_Py0MOOXNCRjqu5dWKtYOjh_SrBqe_gmJDUJ8CTZ-4z3alMnraXdXnpmYG4pLH6Vj5f/s1600/Fimbrios+klossi+Thomas+Calame.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="213" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhGyWvoGi6bnnZE5SbgfAmMWTFLN3Q21TQobyMEi4388keHuOitH_umQ0HquAHI4AkjHie4l2pnE_Py0MOOXNCRjqu5dWKtYOjh_SrBqe_gmJDUJ8CTZ-4z3alMnraXdXnpmYG4pLH6Vj5f/s320/Fimbrios+klossi+Thomas+Calame.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Bearded Snake (<i>Fimbrios klossi</i>)</td></tr>
</tbody></table>
The truth is that both acrochordids and xenodermids are obscure snakes, and we don't have that much data on either one of them. They are both found in areas of the world that are hard to get to. Morphologically, they appear superficially similar, and <span style="text-align: left;"><a href="http://www.biodiversitylibrary.org/item/72743#page/190/mode/1up">an association between them was first hypothesized in 1893</a>. </span>But, even <a href="http://people.earth.yale.edu/sites/default/files/files/Gauthier/Gauthier.pdf">the most comprehensive morphological trait database for snakes</a> is missing crucial data on their anatomy, such as whether or not their <a href="http://snakesarelong.blogspot.com/2014/03/why-do-snakes-have-two-penises.html">hemipenial spines</a> are mineralized. This would be helpful to know because the hemipenial spines of basal snakes such as boas and pythons are not mineralized, whereas those of definitive colubroids are heavily mineralized.<br />
<br /></div>
</div>
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj_JzL7wr7hCaTcdW3fa_Pj8w06slNxDM95wzLYLQha9ukqCFi1eTMhqDm_ihMaysSlM7s3hjwFCjkIo4c-hBjQf0yzn1iSLPe4UA9iHkmQei9QbvESmFJqDAR5XlSMAOUePZ5xYdgIfsTK/s1600/Parafimbrios+lao+Teynie+et+al+2015.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="236" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEj_JzL7wr7hCaTcdW3fa_Pj8w06slNxDM95wzLYLQha9ukqCFi1eTMhqDm_ihMaysSlM7s3hjwFCjkIo4c-hBjQf0yzn1iSLPe4UA9iHkmQei9QbvESmFJqDAR5XlSMAOUePZ5xYdgIfsTK/s320/Parafimbrios+lao+Teynie+et+al+2015.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Parafimbrios lao</i><br />
From <a href="https://www.researchgate.net/profile/Minh_Le8/publication/273260157_A_new_genus_and_species_of_xenodermatid_snake_Squamata_Caenophidia_Xenodermatidae_from_northern_Lao_People's_Democratic_Republic/links/54fd1b4a0cf20700c5ea3f4c.pdf">Teynié et al. 2015</a></td></tr>
</tbody></table>
Within the past year, two new studies on the chromosomes of dragonsnakes (<i>Xenodermus javanicus</i>) have been published. <a href="https://www.researchgate.net/profile/Michail_Rovatsos/publication/284157437_Differentiation_of_Sex_Chromosomes_and_Karyotype_Characterisation_in_the_Dragonsnake_Xenodermus_javanicus_(Squamata_Xenodermatidae)/links/564c94bd08ae4ae893ba78ef.pdf">In the first</a>, the karyotype (the number of chromosomes and their shape) of dragonsnakes was reported for the first time. In humans, each cell normally contains 23 pairs of chromosomes, for a total of 46. <a href="http://chromorep.univpm.it/">In most snakes, each cell normally contains 18 pairs of chromosomes, for a total of 36</a>. Usually, eight of these pairs are relatively large (called macrochromosomes), and the other ten are somewhat small (called microchromosomes). Dragonsnakes have 16 pairs of chromosomes, for a total of 32, of which seven are large and nine are small. The dragonsnake karyotype probably evolved by two fusion events, one of two macrochromosomes and the other between a macrochromosome with a microchromosome. There are some other exceptions to the 18-pair pattern; <a href="https://www.researchgate.net/profile/Nancy_Oguiura/publication/41848785_Cytogenetics_and_molecular_data_in_snakes_a_phylogenetic_approach/links/0fcfd5062ef29db787000000.pdf">some snakes have as few as 12 or as many as 25 pairs</a>, including the only other xenodermid to have been karyotyped, the Sichuan Odd-scaled Snake (<i>Achalinus meiguensis</i>), which has just 12 pairs of chromosomes.<br />
<br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg9gJBkpoxFpjF_H5MvPiysh_2yjW4B1S02hRpc00sGuv1dGqDk3-baQfa5x4MHS3reRWr02wXPsP6qE1TBh7RODrVN7Y-la2iAn_ibziU90WrCjDpc998OJaK_FZg1Lq-pPqCjeotTTUEa/s1600/Achalinus+werneri+Takehiko+Sato+Amami-takachiho-snake-amongst-leaf-litter.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="222" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg9gJBkpoxFpjF_H5MvPiysh_2yjW4B1S02hRpc00sGuv1dGqDk3-baQfa5x4MHS3reRWr02wXPsP6qE1TBh7RODrVN7Y-la2iAn_ibziU90WrCjDpc998OJaK_FZg1Lq-pPqCjeotTTUEa/s320/Achalinus+werneri+Takehiko+Sato+Amami-takachiho-snake-amongst-leaf-litter.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Amami Odd-scaled Snake (<i>Achalinus werneri</i>)</td></tr>
</tbody></table>
From the 1960s to the 1980s, before DNA sequencing became cheap and easy, scientists invested heavily in collecting karyotypes from a diversity of species for comparative purposes, so we can say with pretty good certainty that the ancestral state for all snakes is 36 (18 pairs). That's the number in filesnakes, pareids, most vipers, homalopsids, and many colubrids, lamprophiids, and elapids, although there are <a href="https://www.researchgate.net/profile/Nancy_Oguiura/publication/41848785_Cytogenetics_and_molecular_data_in_snakes_a_phylogenetic_approach/links/0fcfd5062ef29db787000000.pdf">lots of exceptions in the latter three groups</a>. The fusions in xenodermids emphasize their uniqueness, but unfortunately don't shed any new light on their phylogenetic placement.<br />
<br />
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgAr7Pr8gm5h9QKqJulPQ8a7Ve1oJ4oD9gSEevzoYjP18PsWR8TdyTdwxyHHhvc3FZmlA9fjd6ipZvjly2K5ygO6o5xStCXaWLbbynQry9G-CPZphsaI70xHkjbEHaxEkkzw84VqqSeLHpn/s1600/Stoliczkia+borneensis+Sam+Howard+2.jpeg" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="213" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgAr7Pr8gm5h9QKqJulPQ8a7Ve1oJ4oD9gSEevzoYjP18PsWR8TdyTdwxyHHhvc3FZmlA9fjd6ipZvjly2K5ygO6o5xStCXaWLbbynQry9G-CPZphsaI70xHkjbEHaxEkkzw84VqqSeLHpn/s320/Stoliczkia+borneensis+Sam+Howard+2.jpeg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;"><i>Stoliczkia borneensis</i></td></tr>
</tbody></table>
<a href="http://rspb.royalsocietypublishing.org/content/282/1821/20151992.abstract">The other study</a> focused on the sex chromosomes. In humans, sex is determined by which combination of sex chromosomes a baby receives from its parents: two X chromosomes make a female, whereas an X and a Y chromosome make a male. It's pretty similar in snakes, with a twist: the sex chromosomes are called Z and W instead of X any Y, and <a href="https://www.researchgate.net/profile/Lukas_Kratochvil/publication/229951554_Phylogeny_of_sex-determining_mechanisms_in_squamate_reptiles_Are_sex_chromosomes_an_evolutionary_trap/links/0deec51f76b9d5a372000000.pdf">females are the heterogametic sex</a> (meaning that a Z and a W chromosome make a female, and two Z chromosomes make a male). Birds and many other reptiles also have ZW sex determination. In many colubroid snakes, the W chromosome is about twice the size of the Z, and it is often unusual in other ways as well, such as having sections of highly condensed
chromatin or a different centromere position. In contrast, filesnakes, boids, and other more basal snakes have morphologically indistinguishable Z and W chromosomes, although they still contain different genes and perform different functions.<br />
<br />
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgdfIOMXT5oyyd8uCqJb4wmHZ0p_zkL_VdpyRUoLcEBJTPIzwLyk5FE8_kCEXV9CHlQxNz_koabwuCANxQZ6uWkpN7KtcokQbvxIGSE3gC3IuOm7jn8rL-9qSAQxu5tlGsjn-0rE0IirpZd/s1600/Xylophis+perroteti+Zeeshan+Mirza.jpeg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="192" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgdfIOMXT5oyyd8uCqJb4wmHZ0p_zkL_VdpyRUoLcEBJTPIzwLyk5FE8_kCEXV9CHlQxNz_koabwuCANxQZ6uWkpN7KtcokQbvxIGSE3gC3IuOm7jn8rL-9qSAQxu5tlGsjn-0rE0IirpZd/s320/Xylophis+perroteti+Zeeshan+Mirza.jpeg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Perrotet's Narrow-headed Snake (<i>Xylophis perroteti</i>)<br />
Are members of this genus really xenodermids? Or, like the<br />
former xenodermids <i>Oxyrhabdium</i> and <i>Nothopsis</i>, will they<br />
prove to be more closely related to something else?</td></tr>
</tbody></table>
One reason the W chromosome looks so different from the Z in colubroids is that it contains repetitive elements called Bkm ('banded krait minorsatellite') repeats, which consist of the sequence "GATA" (sometimes "GACA") repeated thousands of times. Mammalian X chromosomes <a href="https://www.researchgate.net/profile/Stephen_Sarre/publication/45822187_Non-homologous_sex_chromosomes_of_birds_and_snakes_share_repetitive_sequences/links/00b7d51c7f39ff3a3e000000.pdf">and avian W chromosomes</a> also have these repeats. Cell biologists think that these repeats function to inactivate all the genes on the W chromosome except for those that determine sex<a href="#3" name="top3"><sup>3</sup></a>. Both mammalian X chromosomes and snake W chromosomes become very dense in body cells, so that none of the genes on them can be expressed. They only decondense and plays their brief, female-determining roles, in maturing eggs that are destined to become females. Unlike in mammals, the sex chromosomes of snakes span the gamut from completely identical to markedly differentiated, allowing biologists to study the evolution of chromosomal sex determination. <a href="http://rspb.royalsocietypublishing.org/content/282/1821/20151992.abstract">The new study</a> showed that female dragonsnakes have two different-looking sex chromosomes, with many Bkm repeats in the W, whereas the two Z sex chromosomes of male dragonsnakes look similar and lacked Bkm repeats, bolstering the relationship between xenodermids and other colubroids and diminishing the relationship between xenodermids and filesnakes.<br />
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The other major finding of the new study is the documentation that at least part of the sex chromosomes are homologous across all families of caenophidian snakes, suggesting that snake sex chromosomes emerged in the common ancestor of Caenophidia some 60-80 million years ago. One gene that is only on the Z chromosome in all caenophidians, including dragonsnakes, is also found on the W chromosome in filesnakes. The Z-chromosome-specific genes in caenophidians were on both the Z and W chromosomes in boas, pythons, and sunbeam snakes (Xenopeltidae), as well as in bearded dragons and anoles. Other toxicoferan lizards with ZW sex chromosomes, including chameleons and monitor lizards, seem to have evolved them independently.<br />
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<hr width="80%"><p><span class="Apple-style-span" style="font-size: x-small;"><br />
<a name="1"><b>1 </b></a><a href="http://www.zenscientist.com/index.php/pdflibrary2/Open-Access-Files/ssar_public/Herpetological-Review-1967-2015/2015-Herpetological-Review-46(4)-December/Savage-2015--HR46(4)664-665/">A recent article in the journal </a><i><a href="http://www.zenscientist.com/index.php/pdflibrary2/Open-Access-Files/ssar_public/Herpetological-Review-1967-2015/2015-Herpetological-Review-46(4)-December/Savage-2015--HR46(4)664-665/">Herpetological Review</a> </i>pointed out that the grammatical rules for structuring family and subfamily names from genus names have recently been incorrectly applied in two cases involving snakes which concern this article: 1) Xenodermatidae/inae for the family/subfamily containing <i>Xenodermus</i>, the root of which is "dermus", a masculine noun with which the masculine specific epithet <i>javanicus</i> is correctly coupled (not the neuter <i>javanicum</i>; in contrast think of the neuter <i>Heloderma horridum</i> in family Helodermatidae). The correct family or subfamily name is thus Xenodermidae/inae. 2) Pareatidae or Pareatinae for the family containing <i>Pareas</i>, which is also masculine, making the correct family/subfamily name Pareidae/inae.<a href="#top1"><sup>↩</sup></a><br />
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<a name="2"><b>2 </b></a>Don't confuse this snake genus (<i>Stoliczkia</i>) with a genus of extinct ammonite (<i>Stoliczkaia</i>), both named for <a href="https://en.wikipedia.org/wiki/Ferdinand_Stoliczka">Czech biologist Ferdinand Stoliczka</a>. The extra "a" was added to <a href="http://www.biodiversitylibrary.org/item/130586#page/127/mode/1up">the original spelling</a> of the snake genus by <a href="http://biodiversitylibrary.org/item/81048#page/468/mode/1up">Boulenger</a> in 1899, probably by accident, and this genus is still widely misspelled today (<i>e.g., </i><a href="http://www.ncbi.nlm.nih.gov/nuccore/?term=Stoliczkaia">on GenBank</a> and <a href="https://en.wikipedia.org/w/index.php?title=Talk:Stoliczkaia_khasiensis&redirect=no">on Wikipedia before I fixed it</a> while writing this article).<a href="#top2"><sup>↩</sup></a><br />
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<a name="3"><b>3 </b></a>It's also thought that "GATA" is a particularly potent regulatory sequence, with the power to turn nearby genes on and off. In a way, the sex genes have essentially 'hijacked' the W chromosome, turning off all its other genes, and simultaneously creating a concentrated source of mutation-causing elements. Chromosomal sex determination <a href="http://www.plosbiology.org/article/fetchObject.action?uri=info:doi/10.1371/journal.pbio.1001643&representation=PDF">may therefore constitute a unique and potentially very powerful genotypic mechanism for abruptly enhancing evolutionary rates</a>, which might have contributed to the explosive radiations of species in clades with chromosomal sex determination, such as mammals, birds, squamates, and <a href="http://digital.csic.es/bitstream/10261/81858/4/IJDB%2052%20(837-856)%202008.pdf">certain groups of insects</a>.<a href="#top3"><sup>↩</sup></a><br />
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ACKNOWLEDGMENTS</div>
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Thanks to <a href="https://www.flickr.com/photos/thomascalame/">Thomas Calame</a>, <a href="http://calphotos.berkeley.edu/cgi/img_query?enlarge=0000+0000+0914+0422">Sam Howard</a>, <a href="https://www.researchgate.net/profile/Konrad_Mebert">Konrad Mebert</a>, <a href="http://calphotos.berkeley.edu/cgi/img_query?enlarge=0000+0000+0311+1785">Zeeshan Mirza</a>, <a href="http://www.arkive.org/amami-takachiho-snake/achalinus-werneri/">Takehito Sato</a>, and <a href="https://www.flickr.com/photos/euprepiosaur/">Stephen Zozaya</a> for the use of their photos.<br />
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REFERENCES</div>
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<span style="font-size: x-small;">Pyron, R. A., C. R. Hendry, V. M. Chou, E. M. Lemmon, A. R. Lemmon, and F. T. Burbrink. 2014. Effectiveness of phylogenomic data and coalescent species-tree methods for resolving difficult nodes in the phylogeny of advanced snakes (Serpentes: Caenophidia). Molecular Phylogenetics and Evolution 81:221-231 <<a href="http://med.fsu.edu/userFiles/file/TSL/Genomics__Pyron_et_al,_Molecular_Phylogenetics_and_Evolution_(2014)_81_p221_231.pdf">link</a>></span><br />
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<span style="font-size: x-small;">Zheng, Y., and J. J. Wiens. 2016. Combining phylogenomic and supermatrix approaches, and a time-calibrated phylogeny for squamate reptiles (lizards and snakes) based on 52 genes and 4162 species. Molecular Phylogenetics and Evolution 94:537-547 <<a href="http://www.wienslab.com/Publications_files/Zheng_Wiens_2015b_MPE.pdf">link</a>></span></div>
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="http://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com0Logan, UT, USA41.7369803 -111.833835941.6422018 -111.9951974 41.831758799999996 -111.6724744tag:blogger.com,1999:blog-7443075087825368900.post-77957230065402658682016-01-26T07:19:00.000-07:002017-02-16T09:04:04.148-07:00Book Review: Bushmaster by Dan Eatherley<div style="text-align: center;">
<span style="font-size: x-small;"><a href="https://drive.google.com/open?id=0B2jxDXUJh_X7eG1yaEhndDJfYms">Click here to read this post in Spanish</a></span></div>
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<span style="font-size: x-small;"><a href="https://drive.google.com/open?id=0B2jxDXUJh_X7eG1yaEhndDJfYms">Haga clic aquí para leer este artículo en español</a></span></div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgQp9PceclD9po1yySztD0QkIQlCHvW8fxNAHHv9nCodzR6uN7vBdRgL-HaeRUzysdO4yv52zsIUjGAGUzeOnMFoEmLTLMWX7ONoh9asvu489lswaOUCWvaD1hncquKYzNeLvGR1Lm5R32W/s1600/Lachesis+muta_Marisa+Ishimatsu.jpg" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="250" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgQp9PceclD9po1yySztD0QkIQlCHvW8fxNAHHv9nCodzR6uN7vBdRgL-HaeRUzysdO4yv52zsIUjGAGUzeOnMFoEmLTLMWX7ONoh9asvu489lswaOUCWvaD1hncquKYzNeLvGR1Lm5R32W/s400/Lachesis+muta_Marisa+Ishimatsu.jpg" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Bushmaster (<i>Lachesis muta</i>) from Peru</td></tr>
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Lachesis is the name of <a href="http://www.greekmythology.com/Other_Gods/The_Fates/the_fates.html">one of the three Greek Moirai or Fates</a>, sister-deities who determined the destiny of every human life by spinning each life as a thread on a loom. Her role in the process was to determine the length of a mortal's life, and so she is appropriately immortalized<a href="https://www.blogger.com/blogger.g?blogID=7443075087825368900#1" name="top1"><sup>1</sup></a> in modern biology in the genus name of Bushmasters, huge Latin American pitvipers that occasionally play the same role and are herpetologically mythical in their own right. Her sisters, Clotho (who spun the threads of life) and Atropos (who did the actual thread-cutting), are similarly honored in the Latin name of vipers of the genus <i><a href="http://reptile-database.reptarium.cz/advanced_search?genus=atropoides&submit=Search">Atropoides</a></i><i> </i>and in <i><a href="http://reptile-database.reptarium.cz/advanced_search?common_name=clotho&submit=Search">Clotho</a></i>, an old synonym for some members of the African viper genus <i>Bitis</i>.</div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgvKVX2RTv4_8wtoPlDdXS5gyVqrIYEM1wiy-m0IjC17nqgkL4u1ZGYEh9tDOK_aJYcbprFsymzpokkyfFMdJ5cE3GuyabziGjgCFkPYlgFamHvSVYv7Gdcwjrb5JuyiXiStbpqPHKBC7tB/s1600/Ditmars+filming+Lecky+Eatherly+p255.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="277" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgvKVX2RTv4_8wtoPlDdXS5gyVqrIYEM1wiy-m0IjC17nqgkL4u1ZGYEh9tDOK_aJYcbprFsymzpokkyfFMdJ5cE3GuyabziGjgCFkPYlgFamHvSVYv7Gdcwjrb5JuyiXiStbpqPHKBC7tB/s400/Ditmars+filming+Lecky+Eatherly+p255.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Ditmars filming the Bushmaster "Lecky" at the Bronx Zoo in 1934<br />
©WCS. Courtesy of the <a href="http://ielc.libguides.com/wcs">WCS Archives</a></td></tr>
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If you're interested in Bushmasters and herpetological history, check out <a href="http://www.daneatherley.com/bushmaster.html">Dan Eatherley's new book, <i>"Bushmaster: Raymond Ditmars and the Hunt for the World's Largest Viper"</i></a>, which chronicles the life and times of the Bronx Zoo's first reptile curator and one of America's first and most successful popular herpetological writers. Ditmars authored 24 books, >200 articles, and pioneered nature films in an era when video technology was still in its infancy. Eighty years ago, he was a household name in New York, enjoying a celebrity attained by few herpetologists. President Theodore Roosevelt praised Ditmars's <i>The Reptile Book</i> and invited him to the White House. One of the reasons for his popularity was his "obsession" with keeping large, exotic, sexy, venomous snakes—such as Bushmasters—in captivity, an endeavor on which the press regularly reported. Ditmars was reporter for <i>New York Times</i> when he was young, and the paper published 12-15 stories a year on his exploits between 1899 and 1942. Such was the popularity drummed up for snakes that, when a short-lived Bushmaster named "Lecky" was exhibited at the Bronx Zoo in 1934, <a href="http://www.bioone.org.dist.lib.usu.edu/doi/pdf/10.1643/OT-15-325">it was credited with attracting</a> an estimated 100,000 additional guests at the zoo and a 60% increase in visitors at the nearby American Museum of Natural History’s reptile hall.<br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi1RU-uAlPwQXtOuIbTDMwMCieYoVsTxupyqt9niOTeugbEKQpijqsX4KgQRT_E2tHdlOJJWoPMwI9vFhvaJneirq4ltaKjC_vPedN-07gNLijTD8AK6FPmdC0qzgIqHgyXw-QhfyfV07LD/s1600/Lachesis+muta+with+eggs+Eatherly+p48.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEi1RU-uAlPwQXtOuIbTDMwMCieYoVsTxupyqt9niOTeugbEKQpijqsX4KgQRT_E2tHdlOJJWoPMwI9vFhvaJneirq4ltaKjC_vPedN-07gNLijTD8AK6FPmdC0qzgIqHgyXw-QhfyfV07LD/s400/Lachesis+muta+with+eggs+Eatherly+p48.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">The first photograph of a female Bushmaster guarding her eggs,<br />
taken by C.S. Rogers in Trinidad, was published in <a href="http://www.biodiversitylibrary.org/item/24597#page/465/mode/1up">Ditmars (1910)</a>,<br />
and subsequently as a postcard sold at the Bronx Zoo.<br />
The snake was a captive in the possession of R.R. Mole.</td></tr>
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Bushmasters are unique among New World vipers, with the possible exception of the rare <i>Bothrops colombianus</i>, in laying eggs rather than giving birth to live young. Because they guard their eggs, a phenomenon that Ditmars and his correspondent R.R. Mole first described, they may offer insight into the <a href="http://snakesarelong.blogspot.com/2015/11/snakes-that-are-good-parents.html">complex evolution of parental care in pitvipers</a>. In Ditmars's time, there was a single, widespread species of bushmaster, with four subspecies separated by tropical mountain ranges; we now recognize those four subspecies as species <a href="http://www.eeb.cornell.edu/zamudio/KZ_home/KZ_Publications_files/1997Lachesis.pdf">on the basis of morphological, behavioral, and molecular differences</a>. Bushmasters are also the only pitvipers where <a href="http://www.sciencedirect.com/science/article/pii/004101019090080Q">the venom of juveniles appears to lack the chemical potency of adults</a>, at least towards mammals. Many vipers feed on amphibians or other reptiles when they are young and switch to mammals as they grow up<a href="https://www.blogger.com/blogger.g?blogID=7443075087825368900#2" name="top2"><sup>2</sup></a>, which might explain this observation. Bushmasters are the world's longest vipers<a href="https://www.blogger.com/blogger.g?blogID=7443075087825368900#3" name="top3"><sup>3</sup></a> (<a href="http://snakesarelong.blogspot.com/2013/01/africas-giant-gaboon-vipers.html">Gaboon Vipers </a>exceed them in weight) and the longest venomous snakes in the Americas (<a href="http://snakesarelong.blogspot.com/2015/11/snakes-that-are-good-parents.html">King Cobras</a> exceed them in length).<br />
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<tr><td class="tr-caption" style="text-align: center;">Ditmars wears a snake fang tie pin<br />
on the book's cover</td></tr>
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Eatherley's book is well-researched and accurate. I found it to be an exciting read with an excellent historical perspective. My biggest criticism was that it was a little sensational at times, as are most popular accounts dealing with venomous snakes. I particularly enjoyed the author's description of his experiences meeting up with some New York City herpers to seek Gartersnakes (<i>Thamnophis sirtalis</i>), Brownsnakes (<i>Storeria dekayi</i>), and other snakes that could still be found in the northern part of Central Park in the 1880s, when Ditmars was cutting his herpetological teeth. I was also interested to learn that Ditmars supplied snake venom to early antivenom producers and set a precedent, still in place today, of zoos stocking exotic snake antivenoms for the dual purpose of protecting their keepers and providing them to the medical community when bites from exotic species occur elsewhere.<br />
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In his writing, Ditmars often portrayed Bushmasters as aggressive, in contrast to many other herpetologists who have described their manner as relatively gentle, even timid. In reality, they are, like most venomous snakes, cowards first, then bluffers, and lastly warriors, and their large size has earned them a reputation as formidable warriors as well as a prominent position in folklore throughout Latin America<a href="https://www.blogger.com/blogger.g?blogID=7443075087825368900#4" name="top4"><sup>4</sup></a>. Their mystique and biology effectively drive Eatherley's book, only the second biography of Ditmars ever written (the first, by <a href="http://socialarchive.iath.virginia.edu/ark:/99166/w67q6w5h">Laura Newbold Wood</a>, was written for children and published in 1944, just two years after Ditmars's death). Throughout the book, Eatherley goes from stating that negative responses towards reptiles are “of course, the norm for most of us” (p. 11) to tracing a rapid path from ecstasy to palpable disappointment, familiar to any snake enthusiast, when informed during his search for a wild Bushmaster in Trinidad that a nearby farmer has found one, but killed it (p. 255). I think that Ditmars would be pleased with his abiding influence, nearly 75 years after his death, in inspiring passion for and love of snakes.<br />
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You can read two other reviews of Eatherley's book, published last month in <i><a href="http://www.bioone.org.dist.lib.usu.edu/doi/pdf/10.1643/OT-15-325">Copeia</a></i> and <a href="http://www.zenscientist.com/index.php/pdflibrary2/Open-Access-Files/ssar_public/Herpetological-Review-1967-2015/2015-Herpetological-Review-46(4)-December/Book-Reviews-2015-HR46(3)655-662/" style="font-style: italic;">Herpetological Review</a>.</div>
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<a href="https://www.blogger.com/null" name="1"><b>1 </b></a>I suppose she was already immortal, since she's a Greek Goddess.<a href="https://www.blogger.com/blogger.g?blogID=7443075087825368900#top1"><sup>↩</sup></a><br />
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<a href="https://www.blogger.com/null" name="2"><b>2 </b></a>Strangely, bushmasters seem to be one of the only vipers where this shift is not well-documented. Collecting data on young snakes is hard, and <a href="http://www.sciencedirect.com/science/article/pii/004101019090080Q">the venom study</a> found that venom chemistry became more adult-like after just one year, so perhaps we've just missed the shift. Another hypothesis is that bushmasters tend to hold onto their prey after striking it, unlike other vipers which strike, release, and relocate, so perhaps the rapid immobilizing venom components have been replaced by a mechanical means of immobilization.<a href="https://www.blogger.com/blogger.g?blogID=7443075087825368900#top2"><sup>↩</sup></a><br />
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<a href="https://www.blogger.com/null" name="3"><b>3 </b></a>Regarding their maximum length, <a href="http://www.amazon.com/Venomous-Reptiles-Hemisphere-Comstock-Herpetology/dp/0801441412">Campbell & Lamar's authoritative reference</a> on venomous reptiles of the western hemisphere says: "Documented reports of measured specimens are scarce, however, and the maximum length has been the subject of some hyperbole. Hoge and Lancini (1962) claimed 4.5 m, Abalos (1977) claimed 3.5 m, Ditmars (1937) mentioned specimens of 11 feet (3.35 m) but apparently never saw one exceeding 3m, Bellairs (1969) gave the maximum length as between 3.05 and 3.36m, Dunn (1951) gave the maximum length as 14 feet (4.27 m), and Mertens (1960) listed 13 feet (3.96 m) as the maximum size. Sandner-Montilla (1994) claimed a record of 5.28 m for a Venezuelan specimen of <i>L. muta </i>(with 6-cm fangs!), but such records must be placed in the same realm as 20-m anacondas and other legendary monsters.", and concludes "The great </span><span style="font-size: x-small;">majority of adult specimens of all species of <i>Lachesis </i>measure </span><span style="font-size: x-small;">less than 2.5 m, and 3.5 m is likely near the maximum size."<a href="https://www.blogger.com/blogger.g?blogID=7443075087825368900#top3"><sup>↩</sup></a><br />
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<a href="https://www.blogger.com/null" name="4"><b>4 </b></a>Bushmasters play other roles in human culture as well—as food. </span><span style="font-size: x-small;">Bora and </span><span style="font-size: x-small;">Yagua Indians in eastern Peru consider them a delicacy. They are certainly one of the few snakes large enough to make a filling meal for a family.<a href="https://www.blogger.com/blogger.g?blogID=7443075087825368900#top4"><sup>↩</sup></a><br />
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ACKNOWLEDGMENTS</span></div>
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Thanks to <a href="http://www.daneatherley.com/bushmaster.html">Dan Eatherley and Arcade Publishing</a> for producing such a wonderful book, of which they kindly provided me a copy, to Drew Foster for sharing an advance copy of his review of this book for <i>Copeia</i>, to <a href="http://flickr.com/photos/marisa_ishimatsu">Marisa Ishimatsu</a> and the <a href="http://ielc.libguides.com/wcs">Wildlife Conservation Society</a> for the use of their photographs, and to Harry Greene for shedding a little more light on the diets of juvenile bushmasters.</span></div>
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REFERENCES</span>
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<span style="font-size: x-small;">Adler, K. 1989. Contributions to the History of Herpetology. Volume 1. Society for the Study of Amphibians and Reptiles, Oxford, Ohio <<a href="http://www.ssarbooks.com/?page=shop/flypage&product_id=8707">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Campbell, J. A., and W. W. Lamar. 2004. The Venomous Reptiles of the Western Hemisphere (2 Vol.). Cornell University Press, Ithaca, New York <<a href="http://www.amazon.com/Venomous-Reptiles-Hemisphere-Comstock-Herpetology/dp/0801441412">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Ditmars, R. L. 1910. Reptiles of the World : Tortoises and Turtles, Crocodilians, Lizards, and Snakes of the Eastern and Western Hemispheres. Macmillan Co., New York <<a href="http://www.biodiversitylibrary.org/item/24597#page/9/mode/1up">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Gutiérrez, J., C. Avila, Z. Camacho, and B. Lomonte. 1990. Ontogenetic changes in the venom of the snake <i>Lachesis muta stenophrys</i> (bushmaster) from Costa Rica. Toxicon 28:419-426 <<a href="http://www.sciencedirect.com/science/article/pii/004101019090080Q">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Eatherley, D. 2015. Bushmaster: Raymond Ditmars and the Hunt for the World's Largest Viper. Arcade Publishing, New York, New York <<a href="http://www.daneatherley.com/bushmaster.html">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Foster, C. D. 2015. Bushmaster: Raymond Ditmars and the Hunt for the World’s Largest Viper [book review]. Copeia 103:1107-1109 <<a href="http://www.bioone.org.dist.lib.usu.edu/doi/pdf/10.1643/OT-15-325">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Novotny, R. J. 2015. Bushmaster: Raymond Ditmars and the Hunt for the World's Largest Viper [book review]. Herpetological Review 46:657-659 <<a href="http://www.zenscientist.com/index.php/pdflibrary2/Open-Access-Files/ssar_public/Herpetological-Review-1967-2015/2015-Herpetological-Review-46(4)-December/Book-Reviews-2015-HR46(3)655-662/">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Wood, L. N. 1944. Raymond L. Ditmars: His Exciting Career With Reptiles, Animals and Insects. The Junior Literary Guild and Julian Messner, Inc., New York <<a href="http://www.worldcat.org/title/raymond-l-ditmars-his-exciting-career-with-reptiles-animals-and-insects/oclc/4084207">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Zamudio, K. R., and H. W. Greene. 1997. Phylogeography of the bushmaster (<i>Lachesis muta</i>: Viperidae): implications for neotropical biogeography, systematics, and conservation. Biological Journal of the Linnean Society 62:421-442 <<a href="http://www.eeb.cornell.edu/zamudio/KZ_home/KZ_Publications_files/1997Lachesis.pdf">link</a>></span></div>
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="http://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
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Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com2Logan, UT, USA41.7369803 -111.833835941.6422018 -111.9951974 41.831758799999996 -111.6724744tag:blogger.com,1999:blog-7443075087825368900.post-32683653640876090232015-12-29T07:00:00.000-07:002017-02-16T10:04:00.455-07:00Life is Short but Snakes are Long 2015 Milestones<div style="text-align: justify;">
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<span style="font-size: x-small;"><a href="https://drive.google.com/open?id=0B2jxDXUJh_X7dENiZDBsTDR6b2M">Click here to read this post in Spanish</a></span></div>
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<span style="font-size: x-small;"><a href="https://drive.google.com/open?id=0B2jxDXUJh_X7dENiZDBsTDR6b2M">Haga clic aquí para leer este blog en español</a></span></div>
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Dear reader,<br />
<br />
Instead of debuting my planned new content for this month, I wanted to take a moment to thank you for your readership, to review the several milestones reached by Life is Short but Snakes are Long in 2015, and to outline where it's headed in the future.<br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhFJES3kj4JqcXDlOl3fZ-otJz7pQQypmzyBj1brmoM5LEh3fXP2FwlsPObSCXy0yCOJpohNsza8sNcflswWGFbIwg2_cyDAd1H0dAYHB9cf7uQzCYN9anjpdJB0qY_tTFp0ZNHTbeGBMtM/s1600/Snakes_back_cover.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="205" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhFJES3kj4JqcXDlOl3fZ-otJz7pQQypmzyBj1brmoM5LEh3fXP2FwlsPObSCXy0yCOJpohNsza8sNcflswWGFbIwg2_cyDAd1H0dAYHB9cf7uQzCYN9anjpdJB0qY_tTFp0ZNHTbeGBMtM/s400/Snakes_back_cover.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">The back cover of the paperback edition of Harry Greene's opus <i>Snakes:<br />The Evolution of Mystery in Nature</i>, bearing the review by eminent<br />
nature writer David Quammen from which this blog takes its name</td></tr>
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Life is Short but Snakes are Long reached half a million unique views this year (by over 320,000 unique readers from nearly every country) on September 28th, 2015. When <a href="http://snakesarelong.blogspot.com/2012/04/identifying-snake-sheds.html">I began writing it</a> on April 4, 2012, I would never have imagined that so many people would want to read about snakes. Since that time, the pace and intensity of my dissertation research has increased steadily, and my goal for this year was to publish one article a month, which I am proud to say I have achieved (unless you think that this one is cheating, which I kind of do). Even though I wrote fewer articles this year, many of them were more ambitious than my past articles, in that they synthesized large bodies of literature that I personally knew little about before I started writing.<br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhfxu8CTzWQwnpWvRN0xN9iz9a2no8-SPBM9Has0XQMU4sVw6sCJbUeQ6GSIWoBL4hrZbceF1SiwRk3IhyEizOEEN1xV045mRv-DidVpI_h0l9y2aYXuen8H8mahO61x7BFyTMfeE1USUKm/s1600/map+of+visits.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="205" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhfxu8CTzWQwnpWvRN0xN9iz9a2no8-SPBM9Has0XQMU4sVw6sCJbUeQ6GSIWoBL4hrZbceF1SiwRk3IhyEizOEEN1xV045mRv-DidVpI_h0l9y2aYXuen8H8mahO61x7BFyTMfeE1USUKm/s320/map+of+visits.png" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Countries and regions from which readers have accessed <i>Life is Short</i><br />
From what I can see on this tiny map, we're missing only Svalbard,<br />
Western Sahara, Turkmenistan, and North Korea</td></tr>
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My other goals for this blog include: 1) to provide referenced, reputable information that is not available elsewhere, 2) to synthesize & translate information from the peer-reviewed literature, and 3) to indulge my own broad interests. I know I've been successful with #3, which was important to me because I was afraid of becoming too specialized in the process of getting my PhD. Whether or not I've succeeded at numbers 1 & 2 you'll have to tell me. Apparently at least a few people think so, because articles from Life is Short but Snakes are Long have been syndicated by <a href="http://herpnation.wpengine.com/category/hn-blog/andrew-durso/">HerpNation Media</a> and linked to, covered, or republished (with permission) by:</div>
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<ul>
<li><a href="http://0.0.7.221/06/01/ive-got-your-missing-links-right-here-01-june-2013/">National Geographic</a> (<a href="http://phenomena.nationalgeographic.com/2014/03/29/ive-got-your-missing-links-right-here-29-march-2014/">twice!</a>)</li>
<li><a href="http://www.scientificamerican.com/author/andrew-durso/">Scientific American</a></li>
<li><a href="http://www.bbc.com/earth/story/20150709-the-snake-that-decapitates-its-prey">BBC Earth</a></li>
<li><a href="http://blogs.discovermagazine.com/but-not-simpler/2014/03/30/nerds-words-week-13/">Discover Magazine</a></li>
<li><a href="https://theconversation.com/explainer-why-do-snakes-flick-their-tongues-29935">The Conversation</a>/<a href="http://www.huffingtonpost.com/2014/08/02/why-snakes-flick-their-tongues_n_5638421.html">Huffington Post</a></li>
<li><a href="http://blog.biodiversitylibrary.org/2015/06/life-is-short-but-snakes-are-long.html">Biodiversity Heritage Library</a></li>
<li><a href="http://smallpondscience.com/2014/02/28/friday-recommended-reads-19/">Small Pond Science</a></li>
<li><a href="http://upr.org/post/snakes-and-their-bad-reputation-fridays-access-utah">Access Utah on Utah Public Radio</a></li>
<li><a href="http://news.hjnews.com/allaccess/article_649fd626-5c9f-11e3-aabe-001a4bcf887a.html">Logan Herald-Journal</a></li>
<li><a href="http://archive.sltrib.com/story.php?ref=/sltrib/news/57229913-78/snakes-snake-durso-shed.html.csp">Salt Lake City Tribune</a></li>
<li><a href="http://www.usu.edu/today/index.cfm?id=53451">Utah State Today</a></li>
<li><a href="http://sierraclub.org/sierra/2015-2-march-april/speed/two-months-environmental-news-one-page">The Sierra Club</a></li>
<li><a href="http://www.blogtalkradio.com/theotherherpcast-/2013/12/10/20-the-other-herpcast-tippin-the-scales-1292013">The Other Herpcast</a></li>
<li><a href="https://snaketalk.wordpress.com/2012/12/12/the-snakes-that-eat-caviar-andrew-durso/">Center for Snake Conservation</a></li>
<li><a href="http://www.parcplace.org/news-a-events/2013-year-of-the-snake.html?id=271">Partners in Amphibian and Reptile Conservation</a></li>
<li><a href="http://ncherps.org/">North Carolina Herpetological Society</a></li>
<li><a href="http://herpdigest.org/">HerpDigest</a></li>
</ul>
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I also found out that one of my posts was nominated for a ‘Best Science Writing Online 2013’ contest (although it did not win). Because of the blog, I've also been asked to provide review services on snake biology to Bones on Fox TV, The Blacklist on NBC, and the children's book publisher <a href="http://cherrylakepublishing.com/shop/show/50182">Cherry Lake Publishing</a>. Finally, I was invited to travel to San Antonio, Texas, in May for the <a href="http://www.internationalherpetologicalsymposium.com/">International Herpetological Symposium</a> to speak at their Science Café and also in their general program about Life is Short but Snakes are Long, which I really enjoyed. I want to thank the many editors, writers, scientists, publicists, and reporters who thought my writing was good enough to republish or pay attention to in some form.</div>
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjgB0Fsn9Cv64hrkVvb2oB8HebjljdOp_NEArtpQxb8OFoWoHLXagNpyUwW_S-1jHG1WB-ofmtkw2gMCIkYTX2nOQDGVY0NSOaovl29VXX61ZUsjyk4PAECisp2dVim8WdXozN5wIRsYlo0/s1600/IMG_1155.JPG" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjgB0Fsn9Cv64hrkVvb2oB8HebjljdOp_NEArtpQxb8OFoWoHLXagNpyUwW_S-1jHG1WB-ofmtkw2gMCIkYTX2nOQDGVY0NSOaovl29VXX61ZUsjyk4PAECisp2dVim8WdXozN5wIRsYlo0/s320/IMG_1155.JPG" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">My Sonoran Coralsnake (<i>Micruroides euryxanthus</i>)</td></tr>
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I also reached a personal herping milestone this year: 100 snake species seen alive and in the wild, on July 30th, 2015, with a Sonoran Coralsnake (<i>Micruroides euryxanthus</i>) that I found on Portal Road in Cochise County, Arizona. This was an especially exciting snake for me because it was my first wild elapid and because I was there with <a href="http://www.amnh.org/our-research/southwestern-research-station/education/field-herpetology-of-the-southwest">the American Museum of Natural History Southwest Research Station's Field Herpetology of the Southwest class</a>, where the enthusiasm was nothing short of infectious. I also published several peer-reviewed journal articles and short notes this year, including <a href="http://www.zenscientist.com/index.php/pdflibrary2/Open-Access-Files/ssar_public/Herpetological-Review-1967-2015/2015-Herpetological-Review-46(1)-March/Natural-History-Notes-2015-HR46(1)69-109/">one that I came across as a direct result of writing Life is Short but Snakes are Long</a>. Because I'll be writing and (hopefully) completing my dissertation in 2016, I'll likely be relying more heavily on updating and re-posting existing material, since I'll have less time to research and write new material. But, I have some new articles planned that I've already begun working on, so there should be a mix of old and new in 2016.</div>
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Life is Short but Snakes are Long would not be possible without support from volunteer translators <a href="http://malariaworld.org/blogs/alvaro-pemartin">Alvaro Pemartin</a> & Estefania Carrillo, from Utah State University, particularly my advisor <a href="https://sites.google.com/site/sfrenchlab/home">Susannah French</a> and <a href="http://www.usu.edu/ecology/">the Ecology Center</a>, and from my loving girlfriend and editor Kendal Morris.<br />
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Thank you, and happy 2016!</div>
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="http://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
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Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com2tag:blogger.com,1999:blog-7443075087825368900.post-5640581476905417752015-11-30T17:44:00.004-07:002018-02-27T15:25:33.223-07:00Snakes that are Good Parents<div style="text-align: center;">
<span style="font-size: x-small;"><a href="https://drive.google.com/open?id=0B2jxDXUJh_X7UzBrN2pnUjhwUnc">Click here to read this article in Spanish</a></span><br />
<span style="font-size: x-small;"><a href="https://drive.google.com/open?id=0B2jxDXUJh_X7UzBrN2pnUjhwUnc">Haga clic aquí para leer este artículo en español</a></span><br />
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Almost all mammals and birds care for their young to some extent, but most amphibians and reptiles do not. We tend to think of snakes as particularly asocial, and in many cases this is probably true. But, a growing body of evidence contradicts the generalization, <a href="http://www.jstor.org/stable/2460163">made as recently as 1978</a>, that "all reptiles produce precocial offspring without postnatal parental care", and shows that some snakes, in particular, are more caring parents than we typically think.<br />
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<b>Vipers</b><br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEholMOCujwj1aUo19iLXpURSoIGhMpEY8O1xh6AqKypczU_Jpw4-Q54N0RuUL3nt9nDaMH89Jfzk74baPnjQi1KCot73jQvEtFMOgaG_I3vLzQkWtaYueo5ISWz4dKzmnw58ulJrIfSDLKJ/s1600/Crotalus+horridus+with+newborns+Jim+Williams.jpg" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="232" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEholMOCujwj1aUo19iLXpURSoIGhMpEY8O1xh6AqKypczU_Jpw4-Q54N0RuUL3nt9nDaMH89Jfzk74baPnjQi1KCot73jQvEtFMOgaG_I3vLzQkWtaYueo5ISWz4dKzmnw58ulJrIfSDLKJ/s320/Crotalus+horridus+with+newborns+Jim+Williams.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">A female Timber Rattlesnake (<i>Crotalus horridus</i>)<br />
with her newly-born young
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</tbody></table>
Probably the group of snakes most well-known for parental care are now the vipers, which is somewhat ironic considering the fierce but undeserved reputation of these venomous snakes. Although it was documented as early as 1850, parental care by vipers was not widely known or accepted by the scientific community until the 1990s; <a href="http://beheco.oxfordjournals.org/content/15/3/520.full">like crocodilians</a>, it was assumed that these animals were too vicious to exhibit such caring behavior. When <a href="http://www.sdnhm.org/about-us/our-museum/history/klauber/">Laurence Klauber</a>, at the time the world's foremost authority on rattlesnakes, wrote in 1956 that <i><a href="https://books.google.com/books?id=ijblNZQrk_wC&pg=PA736&dq=does+not+result+from+any+maternal+solicitude;+rather+it+is+only+because+the+refuge+sought+by&hl=en&sa=X&ved=0CB0Q6AEwAGoVChMIzsq_r5qRyQIVE-BjCh3QDAPW#v=onepage&q=does%20not%20result%20from%20any%20maternal%20solicitude%3B%20rather%20it%20is%20only%20because%20the%20refuge%20sought%20by&f=false">"Their propinquity [to aggregate]...does not result from any maternal solicitude; rather it is only because the refuge sought by the mother is also used as a hiding place by the young."</a></i>, he was uncharacteristically incorrect; in hindsight, his words now seem almost willfully ignorant. In the 1990s, <a href="http://www.zenscientist.com/index.php/pdflibrary2/Open-Access-Files/ssar_public/Herpetological-Review-1967-2015/1996-Herpetological-Review-27/Herpetological-Review-27(1)-1996-pages-0-27/">credible reports</a> of <a href="http://www.zenscientist.com/index.php/pdflibrary2/Open-Access-Files/ssar_public/Herpetological-Review-1967-2015/1996-Herpetological-Review-27/Herpetological-Review-27(4)-1996-pages-194-224/">parental care</a> in <a href="http://www.zenscientist.com/index.php/pdflibrary2/Open-Access-Files/ssar_public/Herpetological-Review-1967-2015/1999-Herpetological-Review-30/Herpetological-Review-30(1)-1999-pages-35-64/">wild pitvipers</a> began to accumulate, corroborating the many older stories listed by Klauber, and in 2002, <a href="http://eaglemountainpublishing.s3.amazonaws.com/PDF/Biology%20of%20the%20Vipers/CH%2012_greene_.pdf">a seminal review paper based around two studies using radio-telemetry and DNA proved once and for all that mother rattlesnakes do stay with and care for their young</a>. Today, you can read <a href="http://blog.socialsnakes.org/category/behavior/parental-care/">a whole blog about parental care in rattlesnakes</a>, and we think that parental care is widespread (but not ubiquitous) among the <a href="http://reptile-database.reptarium.cz/advanced_search?taxon=Crotalinae&submit=Search">~230 species</a> of pitvipers (aka <a href="http://snakesarelong.blogspot.com/2013/05/basics-of-snake-taxonomy.html">crotalines</a> or New World vipers). This is particularly remarkable because many of them give birth to live young, which they guard until the young's first shed, even though they may not have eaten for 9-10 months beforehand. It appears that the completion of the first shed cycle is the cue for them to separate, an event which is <a href="http://www.sciencedirect.com/science/article/pii/S0016648014003220">mediated by the same hormone in snakes</a> as it is in birds and mammals. Because snakes swallow their food whole, the mother can't really feed her offspring, and they forage for themselves after they disperse. Pitvipers are the only snakes known to care for their living young; other snakes with parental care limit themselves to care of their eggs.<br />
<br />
<b>Pythons</b><br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhAPHLvTUzwPEcbH01-octVwQGS27I7lUg_6xQLvsbHNxkH1cOk0o9lDRPtRcGAf9Z_ozQnsL2TvYtCS_cC74f-ggXIHVUgdKdQ_-9zDcuBhu3hVdSx58PNFmsKYR-QgNIR9vyK-_FG7HLF/s1600/Python_sebae_brooding_eggs.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhAPHLvTUzwPEcbH01-octVwQGS27I7lUg_6xQLvsbHNxkH1cOk0o9lDRPtRcGAf9Z_ozQnsL2TvYtCS_cC74f-ggXIHVUgdKdQ_-9zDcuBhu3hVdSx58PNFmsKYR-QgNIR9vyK-_FG7HLF/s320/Python_sebae_brooding_eggs.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">A mother African Rock Python (<i>Python sebae</i>)<br />
brooding her eggs</td></tr>
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The next most well-known example of parental care in snakes is egg-brooding behavior in pythons, <a href="http://www.jstor.org.dist.lib.usu.edu/stable/pdf/1717551.pdf?acceptTC=true">first documented in 1835</a>. All <a href="http://reptile-database.reptarium.cz/advanced_search?taxon=Pythonidae&submit=Search">40 species</a> of pythons lay eggs, and most of them coil tightly around them throughout incubation, forsaking food. As with vipers, early reports of this behavior were dismissed, but <a href="http://www.pnas.org/content/18/2/209.full.pdf">by the 1930s</a> observations of pythons in zoos showed that they did indeed brood their eggs. Some species that live in cold climates, such as Indian Pythons (<i>Python molurus</i>) and Carpet Pythons (<i>Morelia spilota</i>), also generate heat using muscle contractions ("shivering"). Measurements taken of brooding Indian Pythons have shown that they can increase the temperature of their clutch by 7-10°F. Even though mother pythons may brood for up to 2 months, <a href="http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.497.8063&rep=rep1&type=pdf">studies have found that</a>, at normal temperatures, they rarely shiver and lose only about 6% of their body mass, suggesting that the costs of brooding are relatively small compared to the benefits, which also include reduced water loss by the eggs and <a href="http://www.rcreptiles.com/articles/why-do-female-ball-pythons-python-regius-coil-so-tightly-around-their-eggs.pdf">hatchlings that develop faster and are larger and more active</a>. The brooding instinct in mother pythons is very strong—<a href="http://onlinelibrary.wiley.com/doi/10.1111/j.1439-0310.2012.02070.x/abstract">lab experiments have shown</a> that they will brood the eggs of other pythons just as readily as they will brood their own, and they will even brood rocks that are the same size as their eggs (a behavior reminiscent of the well-known fixed-action pattern of egg-retrieval behavior in graylag geese). Today, <a href="https://www.researchgate.net/profile/Olivier_Lourdais/publication/236944241_Are_females_maternal_manipulators_selfish_mothers_or_both_Insight_from_pythons/links/0f3175333fdc81b7fa000000.pdf">pythons are frequently used as models</a> to study female reproductive behavior and life-history trade-offs.<br />
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<b>King Cobras</b><br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgpe1FQJ_2Ipf8tVID0fRal9uanBDD9TOr65kXGPAHTTtU1HHBk-yjanxcJ7D8APMGQZwtRobLiYuUvC4R06gBsriaIvvlAr-yzFDWII9RHjQBjAod9UBuRg7sck0PkYyVoHpFRxPc66r3d/s1600/Ophiophagus+hannah+figs+combined+Hrima+et+al+2014.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="400" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgpe1FQJ_2Ipf8tVID0fRal9uanBDD9TOr65kXGPAHTTtU1HHBk-yjanxcJ7D8APMGQZwtRobLiYuUvC4R06gBsriaIvvlAr-yzFDWII9RHjQBjAod9UBuRg7sck0PkYyVoHpFRxPc66r3d/s400/Ophiophagus+hannah+figs+combined+Hrima+et+al+2014.png" width="353" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Top: A female King Cobra guarding her nest<br />
Bottom left: A diagram of a typical King Cobra nest<br />
Bottom right: King Cobra eggs in an excavated nest chamber<br />
From <a href="https://www.researchgate.net/profile/Samuel_Lalronunga2/publication/263008786_Nesting_ecology_of_the_King_Cobra_Ophiophagus_hannah_%28Reptilia_Squamata_Elapidae%29_in_Aizawl_District_Mizoram_India/links/0f3175398708745259000000.pdf">Hrima et al. 2014</a></td></tr>
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That female King Cobras (<i>Ophiophagus hannah</i>) <a href="http://www.arkive.org/king-cobra/ophiophagus-hannah/video-09b.html">use their coils to build a nest</a> of sticks and bamboo leaves and guard their eggs for two to three months <a href="https://books.google.com/books?id=GpDQYoNvPrwC&pg=PA257&lpg=PA257&dq=A+nest+of+king+cobra%27s+eggs+wasey&source=bl&ots=l7KBl-nKnv&sig=OAN4-WEEJK11-e2Iu3bDfPcxoI4&hl=en&sa=X&ved=0ahUKEwiyhbHwh6XJAhXPN4gKHZTQBHMQ6AEINDAD#v=onepage&q&f=false">has been known at least since 1892</a><a href="#1" name="top1"><sup>1</sup></a>. Detailed observation of nest-building and attendance were made in captivity at the Bronx Zoo from 1953-1956, and wild King Cobra nests were surveyed and detailed observations made in 1969<a href="#2" name="top2"><sup>2</sup></a>. King Cobra nests are the largest and most complex of any snake's, measuring up to four feet in diameter and rising to a similar height, with an internal chamber for the 20-50 eggs and sometimes a second one above for the snake, which abandons the nest just before the eggs hatch. The female must select her nesting material and bring it to the nest site, because the species of bamboo that are most commonly used in building the nest <a href="https://www.researchgate.net/profile/Samuel_Lalronunga2/publication/263008786_Nesting_ecology_of_the_King_Cobra_Ophiophagus_hannah_%28Reptilia_Squamata_Elapidae%29_in_Aizawl_District_Mizoram_India/links/0f3175398708745259000000.pdf">are not the most abundant species in the surrounding area</a>. There are also some anecdotal reports that male King Cobras will guard the nest and/or the female. Some sources suggest that female King Cobras are more aggressive towards humans when they are guarding their nests, but most suggest that their behavior is no different than at any other time.</div>
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<b>Other snakes</b><br />
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<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEijRW2375OF5h_i-HIdpXiaulr4JAJVj55mwnzQx3Me7pqhhxTdAQJXgJdLLtsmTrC1JGpK3EjHi0o2BZcqYjKbif60UrL09pfptYqPIw0BidkD9YcTDzMQn6eKZh5VaSJXkLQEjzh7r90Y/s1600/Farancia_abacura_mother_on_nest_Matt_Nordgren.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEijRW2375OF5h_i-HIdpXiaulr4JAJVj55mwnzQx3Me7pqhhxTdAQJXgJdLLtsmTrC1JGpK3EjHi0o2BZcqYjKbif60UrL09pfptYqPIw0BidkD9YcTDzMQn6eKZh5VaSJXkLQEjzh7r90Y/s320/Farancia_abacura_mother_on_nest_Matt_Nordgren.jpg" width="213" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">A female Mudsnake (<i>Farancia abacura</i>)<br />
coils around her eggs in a subterranean nest</td></tr>
</tbody></table>
Maternal attendance or guarding of clutches of eggs is widespread in snakes, but observations in the wild are still fairly uncommon, mostly due to the difficulty of locating nesting sites. There are several excellent reviews of this topic, including those written by <a href="http://carlgans.org/bor-view/?borv=16&borpage=275">Rick Shine (1988)</a>, <a href="https://books.google.com/books?hl=en&lr=&id=IdpVchaptYkC&oi=fnd&pg=PA145&ots=bADMJiD7-D&sig=q3SNIEy_QspSX251elQHPG0jOL4#v=onepage&q&f=false">Carl Gans (1996)</a>, <a href="http://www.linnean.org/Resources/LinneanSociety/Documents/Publications/The-Linnaen/Lin%20Vol%2019_%20no%203_%20July%202003.pdf#page=44">Louis Somma (2003)</a>, and <a href="http://www.crcnetbase.com/doi/abs/10.1201/b10879-19">Zach Stahlschmidt and Dale DeNardo (2011)</a>.</div>
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Other snakes that have been observed guarding their eggs in the wild include:</div>
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<div style="text-align: start;">
<ul>
<li>Other species of true cobras <a href="http://pages.bangor.ac.uk/~bss166/Publications/2007_Cobras_MPE.pdf">(King Cobras are more closely related to mambas)</a>, such as <a href="https://www.researchgate.net/publication/233259217_Captive_biology_of_the_shield-nosed_snake_%28Aspidelaps_scutatus_intermedius%29">Shield-nosed Cobras</a> (<i>Aspidelaps scutatus</i>) and some Asian cobras (genus <i>Naja</i>). Unlike King Cobras, they do not build nests but some may dig holes or enlarge existing holes.</li>
<li><a href="http://carlgans.org/bor-view/?borv=16&borpage=287">Other elapids</a>, including kraits (genus <i>Bungarus</i>), coralsnakes (genus <i>Micrurus</i>), and possibly some sea snakes (genus <i>Laticauda</i>) and terrestrial Australian elapids (genera <i>Demansia</i>, <i>Pseudechis</i>, and <i>Pseudonaja</i>).</li>
<li>Mud and <a href="http://snakesarelong.blogspot.com/2012/04/eel-mocassins.html">Rainbow Snakes</a> (genus <i>Farancia</i>), which spend most of their lives in water but lay and brood their eggs on land.</li>
<li>South American Pondsnakes (<i>Pseudoeryx plicatilis</i>), which are also interesting because they <a href="http://www.ecoevo.com.br/publicacoes/pesquisadores/selma_santos/2016_Reproductive%20modes%20of%20the%20South%20American%20water%20snakes.pdf">are part of a lineage that is close to an evolutionary transition between egg-laying and live-bearing</a></li>
<li><a href="http://repository.kulib.kyoto-u.ac.jp/dspace/bitstream/2433/197269/1/018.029.0204.pdf">Malagasy Hog-nosed Snakes (<i>Leioheterodon madagascarensis</i>)</a>, which may leave their nest and return several times over a period of a few months.</li>
<li>Rhombic Skaapstekers (<i>Psammophylax rhombeatus</i>), which guard their eggs under rocks in southern African grasslands, and <a href="http://www.tandfonline.com/doi/abs/10.1080/04416651.1970.9650762">possibly other species of </a><i><a href="http://www.tandfonline.com/doi/abs/10.1080/04416651.1970.9650762">Psammophylax</a> </i>as well.</li>
<li><a href="http://www.jstor.org.dist.lib.usu.edu/stable/pdf/1440862.pdf?acceptTC=true">New Mexico Blindsnakes (<i>Rena dissecta</i>)</a>, which have been found with their eggs in small colonies in sandstone in Kansas. There are hints of parental care in other blindsnakes as well.</li>
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<div style="text-align: start;">
<div style="text-align: justify;">
It's worth noting that, unlike the case with pythons, survival or physiological benefits to the eggs have not been documented in any of these cases. In addition, there are <a href="http://carlgans.org/bor-view/?borv=16&borpage=287&borp=296">numerous anecdotal reports of egg attendance in other snakes</a>, many of which are based on hearsay and are not backed up by data, photographs, or even descriptions. So, expect this list to grow, but keep in mind that parental care in snakes is still, and will probably always be, the exception rather than the rule.</div>
</div>
<br />
<b>Costs and benefits</b><br />
<b><br /></b>
Except for pythons and pitvipers, the costs and benefits of parental care in snakes have not been examined, and I've mentioned some of the evidence for both in pythons already. Why do rattlesnakes and other pitvipers care for their eggs or young? There are several non-mutually-exclusive theories, including:<br />
<b></b><br />
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhcrrN6S4p5gqBtMXrpUTo3PuTA68PNLYEfXVD34V9PCB_Tdvl9sakv6rlKzZxtN_N0VvYvfupJtZRXDVrxpjSsf6tTZmUWXnzEW0dsh0dlsRkYg5n7iDU2pb2NyeJABDMYBL3fObVFzSkO/s1600/Sistrurus_miliarius_061311_8_mother+%252B+litter_Peter+May.jpg" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="240" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhcrrN6S4p5gqBtMXrpUTo3PuTA68PNLYEfXVD34V9PCB_Tdvl9sakv6rlKzZxtN_N0VvYvfupJtZRXDVrxpjSsf6tTZmUWXnzEW0dsh0dlsRkYg5n7iDU2pb2NyeJABDMYBL3fObVFzSkO/s320/Sistrurus_miliarius_061311_8_mother+%252B+litter_Peter+May.jpg" width="320" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">A mother Pigmy Rattlesnake (<i>Sistrurus miliarius</i>) with<br />
her brood. Because rattlesnake rattles are made of segments<br />
that form each time the snake sheds its skin, newborn snakes<br />
have only one segment and cannot yet make sound.</td></tr>
</tbody></table>
1. To protect them from predators. This might involve any or all of the following:<br />
<br />
<ul>
<li>Physical concealment, especially of the eggs, which are less well-camouflaged than the adults.</li>
<li>Deterrence of predators, which may recognize an adult viper as a threat but not an egg or a juvenile.</li>
<li>Active defense from predators, using venom or the threat thereof. This may be especially important prior to the first shed of the young, since they would probably suffer their heaviest mortality during this stage because of their small size, inexperience, hampered <a href="http://snakesarelong.blogspot.com/2013/11/how-snakes-see-through-closed-eyes.html">eyesight</a> and <a href="http://snakesarelong.blogspot.com/2012/09/snakes-that-can-see-without-eyes.html">pit organ</a> sensitivity, and, in rattlesnakes, their inability to use their rattle.</li>
<li>Socially-facilitated retreat from predators, in which the parent helps the young escape an attack by physically moving them, showing them what to do, or distracting the predator. These may seem like surprisingly sophisticated behaviors for snakes, but <a href="http://eaglemountainpublishing.s3.amazonaws.com/PDF/Biology%20of%20the%20Vipers/CH%2012_greene_.pdf">several observations of mother snakes and their young</a> support this idea, and we are learning that <span style="text-align: justify;">many snakes have subtle but complex social lives and communication abilities </span><a href="http://blog.socialsnakes.org/removing-the-blinders/" style="text-align: justify;">that have long been underappreciated</a><span style="text-align: justify;">.</span></li>
</ul>
<br />
<span style="text-align: justify;">Antipredator benefits of parental care in snakes may vary geographically or in other ways, because </span>some species of pitvipers do not seem to change their defensive behavior when they are guarding their young, but others <a href="http://www.jstor.org/stable/1445520">are more defensive</a>, and still others are <a href="http://www.bio.sdsu.edu/pub/clark/Site/Publications_files/cottonmouth_maternal_behavior.pdf">less defensive but more distracting</a>.<br />
<ol><ol>
</ol>
</ol>
<table align="center" cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEio03Nkb20nKyz4DSqPjs5QAnMRkJZWiZJ9WI96UTcmaI9PvDdi6uh3tNzuDykiUpq8xOupwf29sLMa5GgyiX_4WqXYCvdMewipBp1DqP6-uvilZDLall5PGdXU2CnZk7Nsm4CPgGvBCGkC/s1600/Notechis+scutatus+litter+snuggling+with+data+Aubret+%2526+Shine+2009.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEio03Nkb20nKyz4DSqPjs5QAnMRkJZWiZJ9WI96UTcmaI9PvDdi6uh3tNzuDykiUpq8xOupwf29sLMa5GgyiX_4WqXYCvdMewipBp1DqP6-uvilZDLall5PGdXU2CnZk7Nsm4CPgGvBCGkC/s320/Notechis+scutatus+litter+snuggling+with+data+Aubret+%2526+Shine+2009.png" style="cursor: move;" width="260" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Young Tiger Snakes (<i>Notechis scutatus</i>) snuggling<br />
and data showing that the more litter-mates<br />
they snuggle with, the more slowly they cool off<br />
From <a href="https://www.researchgate.net/profile/Richard_Shine/publication/229792308_Causes_and_consequences_of_aggregation_by_neonatal_tiger_snakes_%28Notechis_scutatus_Elapidae%29/links/0912f503da9f76ec5c000000.pdf">Aubret & Shine 2009</a></td></tr>
</tbody></table>
2. <a href="https://www.researchgate.net/publication/229792308_Causes_and_consequences_of_aggregation_by_neonatal_tiger_snakes_%28Notechis_scutatus_Elapidae%29">Litters or clutches of several species of young snakes</a>, including <a href="http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.384.3448&rep=rep1&type=pdf">some rattlesnakes</a>, aggregate together, without their mothers, in order to conserve water or heat—which, if they were mammals, we would call snuggling. Experiments have shown that they prefer to snuggle—sorry, I mean aggregate—inside shelters that contain their own scent cues, and that snuggling kept them warm, which helped them slither to shelter faster. No one has tested whether young pitvipers that snuggle with their mothers have higher body temperatures or lower rates of evaporative water loss than those snuggling with one another, but physics suggests that they would, since larger animals have a lower surface-area-to-volume ratio and thus lose heat and water more slowly. The presence of the mother may also offset the increased visibility or olfactory conspicuousness to predators of a bunch of aggregated young snakes. If this is the primary benefit, it is easy to see how maternal attendance of eggs could evolve into maternal attendance of the young, because <a href="http://home.gwu.edu/~rpyron/publications/Pyron_Burbrink_2014a.pdf">we think that live birth has evolved many times in snakes</a>, and parental care may have evolved and been lost as many as six and ten times, respectively, in vipers. It's probable that we will continue to fill in the gaps in our knowledge. For example, perhaps we're overlooking the behavior in some poorly-studied vipers, as we did in North American pitvipers for over a century.</div>
<div style="text-align: justify;">
<br /></div>
<div style="text-align: justify;">
<div style="text-align: right;">
</div>
<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiPdrPqEuX9fGadUlehmjkIShVaU7tN4sAU0cBnLZa3wUZyjJWBGoNo0oJTOxf_3aFEsopzAcKJT_5x9VwPSISTx4ai-tkW0_9pPqR1d6KFjtYcN8ib2-6855blJ6fwLfhU6TrL7lFY_wVl/s1600/Greene+et+al+2002+Fig4+phylogeny+of+PC+in+vipers.png" imageanchor="1" style="clear: right; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="400" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEiPdrPqEuX9fGadUlehmjkIShVaU7tN4sAU0cBnLZa3wUZyjJWBGoNo0oJTOxf_3aFEsopzAcKJT_5x9VwPSISTx4ai-tkW0_9pPqR1d6KFjtYcN8ib2-6855blJ6fwLfhU6TrL7lFY_wVl/s400/Greene+et+al+2002+Fig4+phylogeny+of+PC+in+vipers.png" width="293" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Viper family tree showing the evolution of parental care.<br />
A few details have changed but the basic shape of the tree<br />
is the same. Abbreviations: O=oviparous, V=viviparous;<br />
Tr=tropical, Te=temperate. From <a href="http://eaglemountainpublishing.s3.amazonaws.com/PDF/Biology%20of%20the%20Vipers/CH%2012_greene_.pdf">Greene et al. 2002</a></td></tr>
</tbody></table>
3. The week or so of parental care may represent an imprinting period for the young snakes to learn the scent of their mother and of one another, similar to the time a young sea turtle spends imprinting on its natal beach or a young salmon on its natal stream. This would be especially important for snakes in cold climates because <a href="http://www.jstor.org/stable/3892539">they use each others' scent trails to locate hibernation sites</a>. Although there is no direct evidence for the third hypothesis, it is suggestive that, at least in the Americas, temperate pitvipers stay with their young, but live-bearing tropical pitvipers, which do not need to hibernate, do not<a href="#3" name="top3"><sup>3</sup></a>. Other explanations include that memories of their siblings' scents help young snakes avoid inbreeding later in life, or that they promote other social behaviors, such as <a href="http://rsbl.royalsocietypublishing.org/content/early/2012/02/13/rsbl.2011.1217.short">communal basking</a>. Some <a href="https://www.researchgate.net/profile/Rulon_Clark/publication/280917086_Post-birth_separation_affects_the_affiliative_behaviour_of_kin_in_a_pitviper_with_maternal_attendance/links/55dc9cad08aec156b9b184b4.pdf">new data suggest that the adult behavior of pitvipers differs</a> when they are deprived of a maternal attendance period. Tall tales about snakes abound, and initially social behavior ranked among them (there are still false tales about parental behavior in snakes, such as the idea that they swallow their young). Parental care in vipers may <a href="http://165.193.122.153/science/science_news/articles/social_lives_snakes.html">just be the tip of their social iceberg</a>. Research over the last decade has shown that <a href="http://beheco.oxfordjournals.org/content/18/2/487.full">vipers make use of chemical information left behind by other vipers</a> when they choose their foraging sites, like a dog sniffing a fire hydrant. This kind of cryptic sociality in snakes can lead to things like inheritance of <a href="http://rsbl.royalsocietypublishing.org/content/early/2012/02/13/rsbl.2011.1217.short">birthing rookeries</a>, <a href="http://rsbl.royalsocietypublishing.org/content/3/2/131.short">nesting sites</a>, and <a href="http://www.jstor.org/stable/3892539">hibernation sites</a> over many generations. Some research even suggests that <a href="http://media.wix.com/ugd/ecb3e9_96a2f3a5bf234bdea457431499ba4869.pdf">pair-bonding might happen</a> between male and female copperheads. <a href="http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0019041">Some lizards build multi-generational homes</a>; might we one day discover snakes doing the same? If we do, my money is on vipers.</div>
<br />
<div style="text-align: justify;">
<div style="text-align: justify;">
<hr width="80%"><p><span class="Apple-style-span" style="font-size: x-small;"><br />
<a name="1"><b>1 </b></a>Rudyard Kipling's <a href="http://www.gutenberg.org/ebooks/236"><i>The Jungle Book</i></a>, containing the story <i>Rikki Tikki Tavi</i>, which describes a King Cobra pair, nest, and parental behavior, was originally published in 1893-4, just a year later.<a href="#top1"><sup>↩</sup></a><br />
</span>
<br />
<hr width="80%" />
<span class="Apple-style-span" style="font-size: x-small;"><br />
<a href="https://www.blogger.com/null" name="2"><b>2 </b></a>The Kenyan herpetologist J.H.E. Leakey collected eggs from these nests and <a href="http://www.thaiscience.info/Journals/Article/NRCT/10629872.pdf">acknowledged in his paper</a> the support of "the management of the International Hotel, who never once raised any objections to our housing live King Cobras in our rooms."<a href="#top2"><sup>↩</sup></a><br />
</span>
<br />
<hr width="80%" />
<span class="Apple-style-span" style="font-size: x-small;"><br />
<a href="https://www.blogger.com/null" name="3"><b>3 </b></a>Intriguingly, the only two Neotropical pitvipers known to have parental care are also the only two that lay eggs. One is <a href="http://www.nsrl.ttu.edu/publications/opapers/ops/OP153.pdf">the Colombian toad-headed pitviper (<i>Bothrops colombianus</i>)</a>, about which very little is known. The other, the Bushmaster (<i>Lachesis muta</i>), well-known by comparison, is nevertheless a secretive denizen of primary rain forests. In 1910, <a href="http://www.daneatherley.com/bushmaster.html">Inaugural Bronx Zoo herp curator Raymond Ditmars</a> and his Trinidad correspondent, R. R. Mole, were the first to publish a photograph of a female Bushmaster guarding her eggs. They wrote of Bushmasters guarding their eggs in the wild, and numerous subsequent captive snakes have borne these observations out. Although Eyelash Pitvipers (<i>Bothriechis schlegelii</i>) have not been observed to guard their young, they may do so because their young shed several days after birth, like those of temperate pitvipers, rather than within 24 hours of birth, like most tropical live-bearing pitvipers. The pattern of parental care in Old World vipers, about which we have far less information, appears to be more complicated still.<a href="#top3"><sup>↩</sup></a><br />
</span>
</div>
<div style="text-align: center;">
ACKNOWLEDGMENTS</div>
<div style="text-align: center;">
<br /></div>
<div style="text-align: justify;">
Thanks to my parents, for indulging my interest in snakes and encouraging me to pursue a career studying them, and to <a href="https://www.flickr.com/photos/williajb1979/">Jim Williams</a>, <a href="http://volusianaturalist.com/">Peter May</a>, <a href="https://commons.wikimedia.org/wiki/File:Female_Python_sebae_brooding_eggs_Tropicario,_FIN.jpg">J. Lanki</a>, and <a href="https://www.flickr.com/photos/happyherper/">Matt Nordgren</a> for the use of their photos.</div>
<div style="text-align: center;">
<br /></div>
<div style="text-align: center;">
REFERENCES</div>
<br />
<div style="text-align: justify;">
<span style="font-size: x-small;">Aubret, F., X. Bonnet, R. Shine, and S. Maumelat. 2005. Energy expenditure for parental care may be trivial for brooding pythons, <i>Python regius</i>. Animal Behaviour 69:1043-1053 <<a href="http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.497.8063&rep=rep1&type=pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Aubret, F., X. Bonnet, R. Shine, and S. Maumelat. 2005. Why do female ball pythons (<i>Python regius</i>) coil so tightly around their eggs? Evolutionary Ecology Research 7:743-758 <<a href="http://www.rcreptiles.com/articles/why-do-female-ball-pythons-python-regius-coil-so-tightly-around-their-eggs.pdf">link</a>></span><br />
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<span style="font-size: x-small;">Aubret, F., and R. Shine. 2009. Causes and consequences of aggregation by neonatal tiger snakes (<i>Notechis scutatus</i>, Elapidae). Austral Ecology 34:210-217 <<a href="https://www.researchgate.net/profile/Richard_Shine/publication/229792308_Causes_and_consequences_of_aggregation_by_neonatal_tiger_snakes_%28Notechis_scutatus_Elapidae%29/links/0912f503da9f76ec5c000000.pdf">link</a>></span><br />
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<span style="font-size: x-small;">Bates, M. F. 1985. Notes on egg clutches in <i>Lamprophis inornatus</i> and <i>Psammophylax rhombeatus rhombeatus</i>. The Journal of the Herpetological Association of Africa 31:21-22.</span><br />
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<span style="font-size: x-small;">Benedict, F. G., E. L. Fox, and V. Coropatchinsky. 1932. The incubating python: a temperature study. Proceedings of the National Academy of Sciences 18:209-212 <<a href="http://www.pnas.org/content/18/2/209.full.pdf">link</a>></span><br />
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<span style="font-size: x-small;">Brashears, J., and D. F. DeNardo. 2012. Do brooding pythons recognize their clutches? Investigating external cues for offspring recognition in the Children's Python, <i>Antaresia childreni</i>. Ethology 118:793-798 <<a href="http://onlinelibrary.wiley.com/doi/10.1111/j.1439-0310.2012.02070.x/abstract">link</a>></span><br />
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<span style="font-size: x-small;">Brown, G. P., and R. Shine. 2007. Like mother, like daughter: inheritance of nest-site location in snakes. Biology Letters 3:131-133 <<a href="http://rsbl.royalsocietypublishing.org/content/3/2/131.short">link</a>></span><br />
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<span style="font-size: x-small;">Brown, W. S., and F. M. MacLean. 1983. Conspecific scent-trailing by newborn timber rattlesnakes, <i>Crotalus horridus</i>. Herpetologica 39:430-436 <<a href="http://www.jstor.org/stable/3892539">link</a>></span><br />
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<span style="font-size: x-small;">Butler, J.A., T.W. Hull, and R. Franz. 1995. Neonate aggregations and maternal attendance of young in the Eastern Diamondback Rattlesnake, <i>Crotalus adamanteus</i>. Copeia 1995:196–198 <<a href="http://www.jstor.org/stable/1446814">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Campbell, J. A., and W. W. Lamar. 1992. The taxonomic status of miscellaneous Neotropical viperids, with the description of a new genus. Occasional Papers of the Museum, Texas Tech University 153:1-31 <<a href="http://www.nsrl.ttu.edu/publications/opapers/ops/OP153.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Case, T. J. 1978. Endothermy and parental care in the terrestrial vertebrates. American Naturalist 112:861-874 <<a href="http://www.jstor.org/stable/2460163">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Clark, R. W. 2007. Public information for solitary foragers: timber rattlesnakes use conspecific chemical cues to select ambush sites. Behavioral Ecology 18:487-490 <<a href="http://beheco.oxfordjournals.org/content/18/2/487.full">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Clark, R. W., W. S. Brown, R. Stechert, and H. W. Greene. 2012. Cryptic sociality in rattlesnakes (<i>Crotalus horridus</i>) detected by kinship analysis. Biology Letters 8:523-525 <<a href="http://rsbl.royalsocietypublishing.org/content/early/2012/02/13/rsbl.2011.1217.short">link</a>></span><br />
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<span style="font-size: x-small;">Cunningham, G. R., S. M. Hickey, and C. M. Gowen. 1996. <i>Crotalus viridis viridis</i> (Prairie Rattlesnake). Behavior. Herpetological Review 27:24 <<a href="http://www.zenscientist.com/index.php/pdflibrary2/Open-Access-Files/ssar_public/Herpetological-Review-1967-2015/1996-Herpetological-Review-27/Herpetological-Review-27(1)-1996-pages-0-27/">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">DeNardo, D. F., O. Lourdais, and Z. R. Stahlschmidt. 2012. Are females maternal manipulators, selfish mothers, or both? Insight from pythons. Herpetologica 68:299-307 <<a href="https://www.researchgate.net/profile/Olivier_Lourdais/publication/236944241_Are_females_maternal_manipulators_selfish_mothers_or_both_Insight_from_pythons/links/0f3175333fdc81b7fa000000.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Gans, C. 1996. An overview of parental care among the Reptilia. Pages 145-157 in J. S. Rosenblatt and C. T. Snowdon, editors. Parental Care: Evolution, Mechanisms, and Adaptive Significance. Academic Press, San Diego, California, USA <<a href="https://books.google.com/books?hl=en&lr=&id=IdpVchaptYkC&oi=fnd&pg=PA145&ots=bADMJiD7-D&sig=q3SNIEy_QspSX251elQHPG0jOL4#v=onepage&q&f=false">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Graves, B. M. 1989. Defensive behavior of female prairie rattlesnakes (<i>Crotalus viridis</i>) changes after parturition. Copeia 1989:791-794 <<a href="http://www.jstor.org/stable/1445520">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">G</span><span style="font-size: x-small;">reene, H.W., P.G. May, D.L. Hardy, J.M. Sciturro, and T.M. Farrell. 2002. Parental behavior by vipers. Pp. 179-206 In Biology of the Vipers. Schuett, G.W., M. Höggren, M.E. Douglas, and H.W. Greene (Eds.). Eagle Mountain Publishers, Eagle Mountain, UT <<a href="http://eaglemountainpublishing.s3.amazonaws.com/PDF/Biology%20of%20the%20Vipers/CH%2012_greene_.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Hibbard, C. W. 1964. A brooding colony of the blind snake, <i>Leptotyphlops dulcis dissecta</i>. Copeia 1964:222 <<a href="http://www.jstor.org.dist.lib.usu.edu/stable/pdf/1440862.pdf?acceptTC=true">link</a>></span></div>
<div style="text-align: justify;">
<span style="font-size: x-small;"><br /></span></div>
<div style="text-align: justify;">
<span style="font-size: x-small;">Hoss, S.K. and R.W. Clark. 2014. Mother Cottonmouths (<i>Agkistrodon piscivorus</i>) alter their antipredator behavior in the presence of neonates. Ethology 120:933-941 <<a href="http://www.bio.sdsu.edu/pub/clark/Site/Publications_files/cottonmouth_maternal_behavior.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Hoss, S. K., D. H. Deutschman, W. Booth, and R. W. Clark. 2015. Post-birth separation affects the affiliative behaviour of kin in a pitviper with maternal attendance. Biological Journal of the Linnean Society 116:637-648 <<a href="https://www.researchgate.net/profile/Rulon_Clark/publication/280917086_Post-birth_separation_affects_the_affiliative_behaviour_of_kin_in_a_pitviper_with_maternal_attendance/links/55dc9cad08aec156b9b184b4.pdf">link</a>></span></div>
<div style="text-align: justify;">
<span style="font-size: x-small;"><br /></span></div>
<div style="text-align: justify;">
<span style="font-size: x-small;">Hoss, S.K., M.J. Garcia, R.L. Earley, and R.W. Clark. 2014. Fine-scale hormonal patterns associated with birth and maternal care in the cottonmouth (<i>Agkistrodon piscivorus</i>), a North American pitviper snake. General and Comparative Endocrinology 208:85-93 <<a href="http://www.sciencedirect.com/science/article/pii/S0016648014003220">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Leakey, J. 1969. Observations made on king cobras in Thailand during May 1966. Journal of the National Research Council of Thailand 5:1-10 <<a href="http://www.thaiscience.info/Journals/Article/NRCT/10629872.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Mori, A., and T. M. Randriamboavonjy. 2010. Field observation of maternal attendance of eggs in a Madagascan snake, <i>Leioheterodon madagascariensis</i>. Current Herpetology 29:91-95 <<a href="http://repository.kulib.kyoto-u.ac.jp/dspace/bitstream/2433/197269/1/018.029.0204.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Oliver, J. A. 1956. Reproduction in the king cobra, <i>Ophiophagus hannah</i> Cantor. Zoologica 41:145-152.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Reiserer, R., G. Schuett, and R. Earley. 2008. Dynamic aggregations of newborn sibling rattlesnakes exhibit stable thermoregulatory properties. Journal of Zoology 274:277-283 <<a href="http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.384.3448&rep=rep1&type=pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Savary, W. 1999. <i>Crotalus molossus molossus</i> (northern blacktail rattlesnake). Brood defense. Herpetological Review 30:45 <<a href="http://www.zenscientist.com/index.php/pdflibrary2/Open-Access-Files/ssar_public/Herpetological-Review-1967-2015/1999-Herpetological-Review-30/Herpetological-Review-30(1)-1999-pages-35-64/">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Schuett, G., R. Repp, M. Amarello, and C. Smith. 2013. Unlike most vipers, female rattlesnakes (<i>Crotalus atrox</i>) continue to hunt and feed throughout pregnancy. Journal of Zoology 289:101-110 <<a href="http://rattlesnakesofarizona.org/files/PDFs/Crotalus%20atrox%20Pregnancy%20Feeding%20JZ%202013.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Shine, R. 1988. Parental care in reptiles. Pp. 275-330 In Biology of the Reptilia. Gans, C. and R.B. Huey (Eds.). Alan Liss, New York <<a href="http://carlgans.org/bor-view/?borv=16&borpage=275">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Smith, C. F., and G. W. Schuett. 2015. Putative pair-bonding in <i>Agkistrodon contortrix</i> (Copperhead). Northeastern Naturalist 22:N1-N5 <<a href="http://media.wix.com/ugd/ecb3e9_96a2f3a5bf234bdea457431499ba4869.pdf">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Somma, L. A. 2003a. Parental Behavior in Lepidosaurian and Testudinian Reptiles: A Literature Survey. Krieger Publishing Company, Malabar, Florida, USA.</span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Somma, L. A. 2003b. Reptilian parental behaviour. The Linnean 19:42-44 <<a href="http://www.linnean.org/Resources/LinneanSociety/Documents/Publications/The-Linnaen/Lin%20Vol%2019_%20no%203_%20July%202003.pdf#page=44">link</a>></span></div>
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<span style="font-size: x-small;"><br /></span></div>
<div style="text-align: justify;">
<span style="font-size: x-small;">Stahlschmidt, Z.R. and D.F. DeNardo. 2011. Parental care in snakes. Pp. 673-702 In Reproductive Biology and Phylogeny of Snakes. Aldridge, R.D. and D.M. Sever (Eds.). Science Publishers, Enfield, New Hampshire <<a href="http://www.crcnetbase.com/doi/abs/10.1201/b10879-19">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">van Mierop, L. H. S., and E. L. Bessette. 1981. Reproduction of the ball python, <i>Python regius</i> in captivity. Herpetological Review 12:20-22 <<a href="http://www.zenscientist.com/index.php/pdflibrary2/Open-Access-Files/ssar_public/Herpetological-Review-1967-2015/1981-Herpetological-Review-12/Herpetological-Review-12(1)-1981/">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Wall, F. 1924. The Hamadryad or King Cobra, <i>Naja hannah</i> (Cantor). Journal of the Bombay Natural History Society 30:189-195 <<a href="http://www.biodiversitylibrary.org/page/47561556#page/273/mode/1up">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Walters, A. C., and W. Card. 1996. <i>Agkistrodon piscivorus conanti</i> (Florida Cottonmouth). Brood defense. Herpetological Review 27:203 <<a href="http://www.zenscientist.com/index.php/pdflibrary2/Open-Access-Files/ssar_public/Herpetological-Review-1967-2015/1996-Herpetological-Review-27/Herpetological-Review-27(4)-1996-pages-194-224/">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Wasey, G. K. 1892. A nest of King Cobra's eggs. Journal of the Bombay Natural History Society 7:257 <<a href="https://books.google.com/books?id=GpDQYoNvPrwC&pg=PA257&lpg=PA257&dq=A+nest+of+king+cobra%27s+eggs+wasey&source=bl&ots=l7KBl-nKnv&sig=OAN4-WEEJK11-e2Iu3bDfPcxoI4&hl=en&sa=X&ved=0ahUKEwiyhbHwh6XJAhXPN4gKHZTQBHMQ6AEINDAD#v=onepage&q&f=false">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Whitaker, N., P. G. Shankar, and R. Whitaker. 2013. Nesting ecology of the King Cobra (<i>Ophiophagus hannah</i>) in India. Hamadryad 36:101-107.</span></div>
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="http://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
</div>Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com3tag:blogger.com,1999:blog-7443075087825368900.post-59959731211615081082015-10-30T07:44:00.000-06:002017-02-16T10:06:35.490-07:00Are there any countries without snakes?<div style="text-align: justify;">
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: left; margin-right: 1em; text-align: left;"><tbody>
<tr><td style="text-align: center;"><a href="https://commons.wikimedia.org/wiki/File:World_distribution_of_snakes.svg#cite_note-Spawls1995-5" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="216" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhKwOMmew9RyPqNtWcukASFlGfpi2whn09kIEuJvVSrynNkekW_yMhyphenhyphenLcwBP0ZhZxEXeLw8xilAjEmePoHv3FERwLVL7wS8rhpr4IrxcFM9gAptKXZpl-1WxhdU5PDmHRhP6-yxZn_SMJas/s400/World_distribution_of_snakes.png" width="400" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Global distribution of all snake species combined<br />
<a href="https://en.wikipedia.org/wiki/Snake#/media/File:World_distribution_of_snakes.svg">Public domain from Wikipedia</a><br />
Terrestrial data from <a href="https://commons.wikimedia.org/wiki/File:World_distribution_of_snakes.svg#cite_note-Ernst2011-1">Ernst & Ernst (2011)</a> and <a href="https://commons.wikimedia.org/wiki/File:World_distribution_of_snakes.svg#cite_note-Cogger1998-2">Cogger et al. (1998)</a><br />
Sea snake data based on <a href="https://commons.wikimedia.org/wiki/File:World_distribution_of_snakes.svg#cite_note-Campbell2004-3">Campbell & Lamar (2004)</a>, <a href="https://commons.wikimedia.org/wiki/File:World_distribution_of_snakes.svg#cite_note-Phillips2002-4">Phillips (2002)</a>,<br />
<a href="https://commons.wikimedia.org/wiki/File:World_distribution_of_snakes.svg#cite_note-Ernst2011-1">Ernst & Ernst (2011)</a>, and <a href="https://commons.wikimedia.org/wiki/File:World_distribution_of_snakes.svg#cite_note-Spawls1995-5">Spawls & Branch (1995)</a></td></tr>
</tbody></table>
Snakes are found in almost every country in the world, but there are a few places without wild<a href="#1" name="top1"><sup>1</sup></a> snakes. Snake-free land generally falls into two categories: remote islands, mostly <a href="https://en.wikipedia.org/wiki/Category:Volcanic_islands">formed by volcanism</a> or <a href="https://en.wikipedia.org/wiki/Atoll">as atolls</a>, that have never been part of a continental land mass and/or have been isolated from continents for a long time, and continental areas that are or were covered by ice <a href="https://en.wikipedia.org/wiki/Last_Glacial_Maximum">within the last 26,000 years</a> and haven't been recolonized since (for example, there are <a href="http://fossilworks.org/bridge.pl?a=collectionSearch&collection_no=15583">snake fossils</a> from northern Canada, where no snakes live now, <a href="http://www.scotese.com/newpage9.htm">from a time when it was much warmer</a>). There are also snake-free parts of the oceans, and probably there are some urban areas that are so disturbed that no snakes live there any more (<i>e.g., </i>downtown Manhattan), although they once did.<br />
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<b>Iceland</b><br />
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<a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhqUAXGAmDOJDxGLrNUq0D8q7VhwGDrOCzLB6FVLiOwgeL5KdLC5uve6XNH7o8DTJRn2TiZc0UIqLGipKwueh44taOvKQngxAj5GAG170d-PPhEoYdv9zxLPw8O9BON8012fxmV4HEpElPX/s1600/iceland.jpg" imageanchor="1" style="clear: right; float: right; margin-bottom: 1em; margin-left: 1em;"><img border="0" height="161" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEhqUAXGAmDOJDxGLrNUq0D8q7VhwGDrOCzLB6FVLiOwgeL5KdLC5uve6XNH7o8DTJRn2TiZc0UIqLGipKwueh44taOvKQngxAj5GAG170d-PPhEoYdv9zxLPw8O9BON8012fxmV4HEpElPX/s200/iceland.jpg" width="200" /></a></div>
Iceland is a volcanic archipelago just outside the Arctic Circle. Despite its high latitude, Iceland is warmed by the Gulf Stream and has a temperate climate, so snakes might actually do fairly well there, especially if they could take advantage of its plentiful geothermal features, as the high-altitude <a href="http://snakesarelong.blogspot.com/2013/05/hot-spring-snakes.html">hot-spring snakes of Tibet (genus <i>Thermophis</i>)</a> have done. However, Iceland has never been connected to any continent—instead, it was formed about 20 million years ago by a series of volcanic eruptions in the Mid-Atlantic Ridge, which separates the Eurasian and North American plates. It's been at about its current latitude the entire time, and, as far as anyone knows, has never been colonized by snakes. Today, the closest snakes are adders (<i>Vipera berus</i>) in both Scotland (470 mi away) and Norway (600 mi away), both of which are separated by a great deal of very cold ocean.</div>
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<b>Ireland</b><br />
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<a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg0E1P3ungMFG2BHZa93CYWv5Ewn42vmnbp_7cbaNrbmuX0xja1kAN3kbarveYsD1Iatjxt2_l6cTryjNeGQtSXYqTwnhbrDD50ID6f7r-QKcKo2OYJR4z8aAl5C5RczqfoaxbsZPlYE2Vr/s1600/ireland.jpg" imageanchor="1" style="clear: left; float: left; margin-bottom: 1em; margin-right: 1em;"><img border="0" height="200" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEg0E1P3ungMFG2BHZa93CYWv5Ewn42vmnbp_7cbaNrbmuX0xja1kAN3kbarveYsD1Iatjxt2_l6cTryjNeGQtSXYqTwnhbrDD50ID6f7r-QKcKo2OYJR4z8aAl5C5RczqfoaxbsZPlYE2Vr/s200/ireland.jpg" width="158" /></a></div>
Unlike Iceland, Ireland was once connected to other land masses. Parts of it are at least 1.7 billion years old. At the end of the Precambrian, two pieces of rock that would become Ireland could be found beneath the sea, one piece connected to the continent of Laurentia and the other piece to the smaller continent of Avalonia, both around 80° South. Over the next 50 million years, these two parts drifted northward, eventually uniting and breaking sea level near the equator about 440 million years ago, in the Silurian Period. Throughout the late Paleozoic Era, Ireland sank back under the sea and gained 65% of its modern mass as limestone deposits from huge coral reefs. At the beginning of the Mesozoic, Ireland was at the latitude of present-day Egypt and had a desert climate, and <a href="https://en.wikipedia.org/wiki/Snake#Origins">by the time snakes evolved</a> (150 million years ago, in the late Jurassic-early Cretaceous) <a href="http://www.scotese.com/jurassic.htm">Ireland had separated from any other land mass</a>, and has been connected on and off to this day. <a href="https://www.researchgate.net/publication/259141117_The_Island_of_Ireland_Drowning_the_Myth_of_an_Irish_Land-bridge">There is some debate</a> over how recently a land bridge connected Ireland with Great Britain and, by extension, mainland Europe, with the consensus resting on the idea that Ireland was isolated by ocean by 16,000 years ago, at which time the climate was still quite cold and there was a lot more ice in Ireland than there is now. Although it's not insane to think that snakes might have colonized Ireland from Europe sometime during the 90 million years that preceded the Pleistocene Ice Ages, as they have since re-colonized Great Britain, so far no one has found any snake fossils in Ireland. But, viviparous lizards, natterjack toads, and common frogs have managed to make it to Ireland, and the slowworm has been introduced there, so it could happen one day. Likely successful colonists include adders (<i>Vipera berus</i>), grass snakes (<i>Natrix natrix</i>), or smooth snakes (<i>Coronella austriaca</i>) from Great Britain, France, or Scandinavia. The Irish climate is highly moderated by the gulf stream, with much milder winters than expected for such a northerly area, so snakes could do quite well there.<br />
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<b>Cape Verde</b><br />
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<a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjfkqBgwWzyt4hB8bCPEePfCN27knbSjRjS3YX4etos0qm2Q87kEz8Nkp-Z0N3DIv-KB1MJYvx5XxgV_8wcL9BlH3lRS8-ZvJTfbOuDHbyDsMWjy1Yy9iOsCzpTabN7wwOap5Zb8WCGY3KS/s1600/Cape-Verde.gif" imageanchor="1" style="clear: right; float: right; margin-bottom: 1em; margin-left: 1em;"><img border="0" height="138" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEjfkqBgwWzyt4hB8bCPEePfCN27knbSjRjS3YX4etos0qm2Q87kEz8Nkp-Z0N3DIv-KB1MJYvx5XxgV_8wcL9BlH3lRS8-ZvJTfbOuDHbyDsMWjy1Yy9iOsCzpTabN7wwOap5Zb8WCGY3KS/s200/Cape-Verde.gif" width="200" /></a></div>
Cape Verde is an island country consisting of 10 volcanic islands in the central Atlantic Ocean, 350 miles off the coast of the western African countries of Mauritania and Senegal. The Cape Verde Islands were all formed by the same volcanic hot spot, the oldest 26 million years ago and the youngest just 100,000 years ago. They have never been colonized by snakes from mainland Africa. There is a single reference to the Striped Sand Snake (<i>Psammophis sibilans</i>) on the island of Sal <a href="http://molevol.cmima.csic.es/carranza/pdf/Vasconcelos%20etal_2013_Oryx_Atlas%20CV_suppMat.pdf">in a 1951 paper</a> that, according to the authors, was an accidental introduction from Guinea-Bissau. Neither this snake nor any other has ever been recorded again from Cape Verde, although the archipelago is <a href="http://molevol.cmima.csic.es/carranza/pdf/Vasconcelos%20etal_2013_Oryx_Atlas%20CV_suppMat.pdf">home to 31 endemic lizard species</a>, more than any other island chain in the <a href="https://en.wikipedia.org/wiki/Macaronesia">Macaronesian region</a>.<br />
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<b>New Zealand</b><br />
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<a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgN0uOP_9TeIWIgl3yAYUWS3QwqdJmrwSFVIcrVwbFAw-qtMlnANx1mckHuOiuOcHQ64XzKxp1m7cDa-63NhXOURrjLS4F1PiR4PWqoz0s7jtCoTY-Y1w6i60JssDNBz9RQTOWdS8F5Nc51/s1600/new-zealand-map-big.jpg" imageanchor="1" style="clear: left; float: left; margin-bottom: 1em; margin-right: 1em;"><img border="0" height="200" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgN0uOP_9TeIWIgl3yAYUWS3QwqdJmrwSFVIcrVwbFAw-qtMlnANx1mckHuOiuOcHQ64XzKxp1m7cDa-63NhXOURrjLS4F1PiR4PWqoz0s7jtCoTY-Y1w6i60JssDNBz9RQTOWdS8F5Nc51/s200/new-zealand-map-big.jpg" width="161" /></a></div>
New Zealand was part of Gondwana (aka Gondwanaland), the more southerly of the two supercontinents formed by the breakup of Pangaea 200-180 million years ago. Gondwana comprised the present-day continents of South America, Africa, Australia, India, and Antarctica as well as New Zealand. Today, New Zealand is the highest part of a mostly-submerged continent called Zealandia that broke away from Gondwana between 100 and 80 million years ago. Since that time, New Zealand has developed a unique flora and fauna that does not include any terrestrial snakes, which makes sense since it has been isolated since around the dawn of their evolution (and has been mostly submerged several times since). However, a steady trickle of reports of sea snakes, borne by oceanic currents beyond their normal range to New Zealand waters and beaches, was <a href="http://www.tandfonline.com/doi/pdf/10.1080/00288330.1997.9516781">summarized in 1997</a>, at which time an amazing 69 records of 2 species were known, dating back to 1837 (more records and a third species have been added since). About 90% are of <a href="http://snakesarelong.blogspot.com/2013/08/pelagic-sea-snakes-and-animals-that.html">pelagic sea snakes</a> (<i>Hydrophis platurus</i>;<i> </i>formerly <i>Pelamis platurus</i>, also known as yellow-bellied sea snakes), a very widespread species that is infamous for vagrancy and recently <a href="http://www.npr.org/2015/10/19/450030340/rare-yellow-bellied-sea-snake-washes-ashore-in-southern-california">made headlines</a> when one washed ashore in Ventura County, California. The remaining 10% of records are of banded sea snakes (<i>Laticauda colubrina</i>), a species that normally sticks more closely to shores, and judging by their morphology most of these have likely come to New Zealand from Fiji or Tonga. In 1995, one specimen in the British Museum collected in New Zealand in 1925 and formerly classified as <i>L. colubrina</i> was <a href="http://www.jstor.org.dist.lib.usu.edu/stable/pdf/4093026.pdf?acceptTC=true">re-identified as a new species from New Caledonia, <i>L. saintgironsi</i>,</a> by herpetologists revising the widespread <i>Laticauda colubrina</i> complex.<br />
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<table cellpadding="0" cellspacing="0" class="tr-caption-container" style="float: right; margin-left: 1em; text-align: right;"><tbody>
<tr><td style="text-align: center;"><a href="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgYzwHig38U0ysO19k6-jzEWrlHmaZQhUV_xP_5a_nyscZE1WcGDdxA34QqhZs7mIAV_6LBWZKlycIGG__GfMdT8RGvTKLRnOmI6JFR2iHifID3d5Tiw1vuy9Tmyo8gyUkqkPEn6O1EsXhZ/s1600/Gill+1997+Pelamis+map+NZ.png" imageanchor="1" style="clear: left; margin-bottom: 1em; margin-left: auto; margin-right: auto;"><img border="0" height="320" src="https://blogger.googleusercontent.com/img/b/R29vZ2xl/AVvXsEgYzwHig38U0ysO19k6-jzEWrlHmaZQhUV_xP_5a_nyscZE1WcGDdxA34QqhZs7mIAV_6LBWZKlycIGG__GfMdT8RGvTKLRnOmI6JFR2iHifID3d5Tiw1vuy9Tmyo8gyUkqkPEn6O1EsXhZ/s320/Gill+1997+Pelamis+map+NZ.png" width="240" /></a></td></tr>
<tr><td class="tr-caption" style="text-align: center;">Map of pelagic sea snake records from New Zealand<br />
(1837-1997)<br />
From <a href="http://www.tandfonline.com/doi/pdf/10.1080/00288330.1997.9516781">Gill 1997</a></td></tr>
</tbody></table>
High sea surface temperatures in 1969-1975 and again in 1988-1990 coincided with major influxes of tropical and subtropical fishes, sea turtles, and sea snakes (up to 16 a year) carried to New Zealand waters by the East Australian Current. Most records are of single animals, but in March 1985 four <i>H. platurus</i> were found on Tokerau Beach in Northland. About three-quarters of sea snake records are from Austral autumn (March-May), and many are from the north coast of the north island, but <i>H. platurus</i> has been found all around the North Island, including in the Cook Strait, and once even on the north coast of the South Island (at Pakawau, Golden Bay, in March 1974)! All <i>L. colubrina</i> records are from the north-east coast of the North Island, except for one at Castlepoint, Wairarapa, in August 1977. All records are of adult snakes, and most (79%) were alive when found, usually washed ashore, but occasionally swimming freely. One even swam up a stream near the sea! Even more amazingly, several sea snakes have been found alive inland from the coast, including a May 1938 record of <i>H. platurus</i> "some distance" from the sea at Table Cape on the Mahia Peninsula, a January 1990 record of <i>L. colubrina</i> "well above" the high-tide line at Whangaruru Harbour, an April 1938 record of <i>H. platurus</i> 200 feet from the sea on a lawn at New Plymouth, and, most incredible, a September 1945 record of <i>L. colubrina</i> alive at Te Aroha, near Hamilton, which is over 12 miles from an estuary over a range of hills or over 27 miles from the ocean along the Waihou River. Unlike <i>H. platurus</i>, which is almost incapable of moving on land, <i>L. colubrina</i> is reasonably good at terrestrial locomotion, which could explain the inland presence of these snakes. Alternatively, <a href="http://www.tandfonline.com/doi/pdf/10.1080/00288330.1997.9516781">the author of the review paper</a> suggested that the snakes could have been carried inland by birds.<a href="#2" name="top2"><sup>2</sup></a><br />
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New Zealand also owns the Chatham Islands 560 miles to the east, the Kermadec Islands 620 miles to the north, and Tokelau 2000 miles to the northeast<a href="#3" name="top3"><sup>3</sup></a>, but no sea snakes have been reported from these islands, probably because so few people live there. Like vagrant birds, even the records from mainland New Zealand surely represent just a small percentage of the total number of marine reptiles that have reached New Zealand over the years. However, New Zealand is still widely considered to have no native snakes, since <i>H. platurus</i> stop feeding at sea temperatures below 18°C and die at temperatures between 14.5 and 17°C (<a href="http://www.seatemperature.org/australia-pacific/new-zealand/taipa-september.htm">the average sea temperature in the coldest month in northern New Zealand is 16°C</a>).<br />
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<b>Kiribati</b><br />
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Kiribati is a Pacific Island nation that straddles the region of the central Pacific Ocean where the Equator and the International Date Line cross, making it the only country that is in all four hemispheres. It consists of four island groups totaling 32 atolls and one coral island. Of these, approximately the eastern half (the Phoenix and Line Islands) are apparently devoid of snakes; at least, they are listed as having no snakes in <a href="https://books.google.com/books/about/Reptiles_and_Amphibians_of_the_Pacific_I.html?id=3H65ngEACAAJ">the most up-to-date and authoritative guide to the reptiles of the Pacific Islands</a>. This guide takes a conservative approach in listing only species that are confirmed by a museum specimen or literature record, so it's possible that at least <a href="http://snakesarelong.blogspot.com/2013/08/pelagic-sea-snakes-and-animals-that.html">pelagic sea snakes</a> are found in the waters of eastern Kiribati. What is certain is that the western half of Kiribati (Banaba and the Gilbert Islands) is home to breeding populations of banded sea snakes (<i>Laticauda colubrina</i>), and possibly pelagic sea snakes as well. Additionally, there is a single record of an ornate reef seasnake (<i>Hydrophis ornatus</i>), a species that is normally found much farther west, from the Gilbert Islands. This might represent a vagrant, but more likely it is a misidentified or mislabeled specimen. So, Kiribati has no terrestrial snakes, unless you count banded sea snakes, which mate, lay eggs, and sometimes digest food on land, but hunt, catch prey, and spend much of their time in the ocean.<br />
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<b>Tuvalu</b><br />
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Tuvalu is a Pacific Island nation south of Kiribati comprising three reef islands and six atolls and totaling 10 square miles, making it the fourth smallest country in the world. Like Kiribati, Tuvalu has no terrestrial snakes unless you count <i>L. colubrina</i>, but unlike Kiribati it has literature records of pelagic sea snakes off its shores. Happily, Tuvalu has decided to honor this species by putting it <a href="http://talismancoins.com/products/64957">on one of its coins</a>! It's a commemorative coin rather than a coin that's actually part of normal circulation, but still, it's pretty cool to have a snake on your money. Tuvalu is also home to at least 9 species of lizards and the introduced cane toad, so it's possible that snakes could show up there one day. In fact, it's even possible that a native, endemic blindsnake could have escaped detection on Tuvalu (or any other Pacific island) to this day. The only reason the Federated States of Micronesia aren't on this list is because of <a href="http://www.fs.fed.us/psw/publications/falanruw/psw_2012_falanruw001_wynn.pdf?">two unexpected species of endemic blindsnakes</a>, <i>Ramphotyphlops adocetus</i> and <i>R. hatmaliyeb</i>, described in 2012 from two small islands, one in the eastern part of FSM and the other in the western part.<br />
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<b>Nauru</b><br />
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Nauru is a relatively isolated Pacific Island nation and is one of the only countries smaller than Tuvalu (at 8.1 square miles, only Monaco and Vatican City, both in Europe, are smaller). Unlike many Pacific Island nations, Nauru is a single island. Nauru has no native terrestrial snakes, but it does have <i>H. platurus</i> off its shores, and it also has what is likely an introduced species, the ubiquitous <i>Indotyphlops braminus</i> or <a href="http://snakesarelong.blogspot.com/2014/04/the-most-widespread-snake-in-world.html">Brahminy Blindsnake</a>, the only unisexual species of snake. It's actually amazing to me that we're on the seventh entry and haven't encountered this species yet, considering <a href="http://snakesarelong.blogspot.com/2014/04/the-most-widespread-snake-in-world.html">how widespread it is globally</a>. The original native range of <i>I. braminus</i> is unknown, but it probably evolved in continental Asia. Because a single individual constitutes a reproductively-competent population, it has since spread all over the world, and it's unclear how long it has been established on Nauru or elsewhere in the Pacific. Many similarly-widespread species in the Pacific owe their distribution to human-assisted transport, the precise timeline of which is difficult to determine. Given the harm done to Nauru's environment by phosphate mining during the 20th century, it's unlikely that any native terrestrial snake would have survived.</div>
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<b>Marshall Islands</b><br />
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The Marshall Islands (see above map) have close political ties with the USA, but they are self-governing. They are located north of Kiribati, west of the FSM, and south of Wake Island. The <a href="https://books.google.com/books/about/Reptiles_and_Amphibians_of_the_Pacific_I.html?id=3H65ngEACAAJ">authoritative guide to the reptiles of the Pacific Islands</a> lists only <i>I. braminus</i> from the Marshall Islands, but <a href="http://www.columbia.edu/itc/cerc/danoff-burg/invasion_bio/inv_spp_summ/boiga_irregularis.html">other sources </a>suggest that at least a few brown treesnakes (<i>Boiga irregularis</i>), infamously introduced to Guam, have been found there as well, and it's possible that <i>H. platurus</i> and possibly other sea snakes are found off its shores. Both the Gilbert Islands in Kiribati to the south and Pohnpei and Kosrae in FSM to the west have <i>L. colubrina</i>, although <a href="http://biormi.org/index_nav_left.shtml?en/non_dangerous_env.html">an official page</a> states that the Marshall Islands have no sea snakes. So, as far as we know the Marshall Islands have no snakes that are native and terrestrial (unless you count <i>I. braminus</i> as native, considering that we don't know how long it's been there).<br />
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<b>Vatican City</b><br />
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The Vatican is a walled enclave within the city of Rome, Italy, with an area of 110 acres and a population of 842, making it the smallest internationally-recognized independent state in the world, both by area and population. I couldn't find any references confirming or denying the presence of wild snakes in the Vatican, but <a href="https://books.google.com/books?id=PJ9bCAAAQBAJ&pg=PT38&lpg=PT38&dq=wildlife+in+vatican+city&source=bl&ots=XZsoR-ArOg&sig=4zW4E244odTSQ1dl5HVXFiQEvoA&hl=en&sa=X&ved=0CDUQ6AEwBGoVChMIlI-Ci8XoyAIVDc5jCh2xXQZ2#v=onepage&q=wildlife%20in%20vatican%20city&f=false">other wildlife seem to be pretty minimal</a>, which makes sense considering that Rome has been a large city for thousands of years. But, snakes and other wildlife can hang on in some amazingly urbanized places, so I wouldn't completely rule out the presence of a few of the eight species of snakes that can surely be found in the surrounding Italian countryside. <b>Monaco</b>, another European microstate with a very dense population and a high degree of urbanization, is another possibility for a snake-less nation, although, given Monaco's reputation as a playground for the rich and famous (30% percent of its population are millionaires), there are certainly some who meet <a href="http://www.urbandictionary.com/define.php?term=snake&defid=3940106">an alternate definition of the word "snake"</a> within its walls.<br />
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<tr><td class="tr-caption" style="text-align: center;">Cover of a joke book that's blank inside</td></tr>
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So there you have it: a maximum of ten countries out of 196 "without snakes", depending on where you want to draw the line. If we start expanding into territories or disjunct sections of larger countries, the list grows considerably, including places like Greenland, the Falkland Islands, Bermuda, Hawaii<a href="#4" name="top4"><sup>4</sup></a>, Wake Island, Johnston Atoll, Howland & Baker Islands, the Marquesas Islands, the Pitcairn Islands, Sala y Gomez, Isla Malpelo, St. Helena, the Faroe Islands, the <a href="http://www.iomtoday.co.im/news/isle-of-man-news/snake-s-alive-walker-s-slippery-surprise-1-7141066">Isle of Man</a>, many Arctic and Antarctic islands, and Antarctica itself, which is owned by no country. And of course, as you can see from the map at the top, there are also large mainland areas of northern Europe, Asia, and North America, as well as the southern tip of Patagonia, that are too cold for snakes (although <i>Vipera</i> <i>berus</i> gets above the Arctic Circle in Scandinavia), not to mention the Atlantic, Arctic, and Antarctic Oceans<a href="#5" name="top5"><sup>5</sup></a>.<br />
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In the course of the research I did for this post, I found many <a href="http://www.npr.org/2015/10/13/448182595/island-travel-tips-miss-the-hiss-pick-the-purr">travel articles promoting the snakelessness of some of these places</a> as overwhelmingly positive, as I'm sure it is for many ophidiophobic travelers. But, <a href="http://snakesarelong.blogspot.com/2013/11/the-truth-about-snakebite.html">the risk that snakes pose is way, way smaller than the fear we have of them</a>, and in my mind the real danger is that many people see eradication of snakes as a positive thing, despite the fact that many of them are in real danger of extinction. Mauritius barely made it off this list, with <a href="http://snakesarelong.blogspot.com/2012/04/round-island-splitjaw-snakes.html">one of two native species extinct and the other hanging on</a> thanks only to captive breeding and reintroduction efforts. St. Kitts & Nevis could lose <a href="http://www.iucnredlist.org/details/942/0">its only native snake, the Saba or orange-bellied Racer (<i>Alsophis rufiventris</i>)</a>, and <a href="http://snakesarelong.blogspot.com/2015/01/dwarf-boas.html">native snakes have gone extinct or become critically endangered on many other islands</a> throughout the Pacific and Caribbean due to centuries of forest clearance, overgrazing, development, and the introduction of invasive species, not to mention the many <a href="http://snakesarelong.blogspot.com/2012/09/recent-conservation-successes-with.html">continental snake species threatened by sprawling development and habitat fragmentation</a>. So, please, let's keep this list from growing.</div>
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<a href="https://www.blogger.com/null" name="1"><b>1 </b></a>Given the growing popularity of herpetoculture, I'd be willing to bet that there are captive snakes in every country, although <a href="http://knowledgenuts.com/2013/10/09/snakes-are-illegal-in-new-zealand/">a few countries have stringent laws banning any captive snakes, including as pets as well as in zoos and research facilities</a>.<a href="#top1"><sup>↩</sup></a><br />
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<span class="Apple-style-span" style="font-size: x-small;"><br />
<a href="https://www.blogger.com/null" name="2"><b>2 </b></a><a href="http://adsabs.harvard.edu/abs/1970Natur.228.1288R">Studies have shown that</a>, although many Pacific birds avoid pelagic sea snakes, naive Atlantic birds will try eat them (only to throw them up, since they are apparently poisonous as well as venomous). New Zealand's birds might be sufficiently naive to try to eat one.<a href="#top2"><sup>↩</sup></a><br />
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<span class="Apple-style-span" style="font-size: x-small;"><br />
<a href="https://www.blogger.com/null" name="3"><b>3 </b></a>Zug's <a href="https://books.google.com/books/about/Reptiles_and_Amphibians_of_the_Pacific_I.html?id=3H65ngEACAAJ">Reptiles and Amphibians of the Pacific Islands</a> lists Tokelau as having no snakes, not even sea snakes, but does not cover the Chatham or Kermadec Islands.<a href="#top3"><sup>↩</sup></a><br />
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<span class="Apple-style-span" style="font-size: x-small;"><br />
<a href="https://www.blogger.com/null" name="4"><b>4 </b></a>Hawaii has introduced Brahminy Blindsnakes and, unlike many Pacific Islands, <a href="http://eol.org/pages/1055435/details">it is known</a> that these colonized the island chain more recently, in 1930, when they were imported from the Philippines in potted palm trees. Hawaii also has pelagic sea snakes and there are a few records of introduced brown treesnakes and boa constrictors, but neither species has established a breeding population (yet).<a href="#top4"><sup>↩</sup></a><br />
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<span class="Apple-style-span" style="font-size: x-small;"><br />
<a href="https://www.blogger.com/null" name="5"><b>5 </b></a><a href="http://www.nature.com/nature/journal/v228/n5278/abs/2281288a0.html">A study</a> evaluating the probability that pelagic sea snakes could enter the Caribbean and Atlantic through the Panama canal, as lionfish have, concluded that there were no real barriers to their colonization of the eastern side of the Americas, but so far this has not happened.<a href="#top5"><sup>↩</sup></a><br />
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ACKNOWLEDGMENTS</div>
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Thanks to Kerry Nelson for doing some of the background research for this post as part of a discussion in the <a href="https://www.facebook.com/groups/snakeED/">Wild Snakes: Education & Discussion</a> Facebook group.<br />
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REFERENCES</div>
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<span style="font-size: x-small;">Edwards, R. J., and A. J. Brooks. 2008. The Island of Ireland: Drowning the Myth of an Irish Land-bridge? Pages 19-34 in J. J. Davenport, D. P. Sleeman, and P. C. Woodman, editors. Mind the Gap: Postglacial Colonisation of Ireland. Special Supplement to The Irish Naturalists’ Journal <<a href="http://www.tara.tcd.ie/bitstream/handle/2262/40560/Edwards&Brooks_INJ08_TARA.pdf;jsessionid=086FD4079DD2778AFEE38FCC788E835E?sequence=1">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Gill, B. J. 1997. Records of turtles and sea snakes in New Zealand, 1837-1996. New Zealand Journal of Marine and Freshwater Research 31:477-486 <<a href="http://www.tandfonline.com/doi/pdf/10.1080/00288330.1997.9516781">link</a>></span><br />
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<span style="font-size: x-small;">Heatwole, H., S. Busack, and H. Cogger. 2005. Geographic variation in sea kraits of the <i>Laticauda colubrina</i> complex (Serpentes: Elapidae: Hydrophiinae: Laticaudini). Herpetological Monographs 19:1-136 <<a href="http://www.bioone.org/doi/full/10.1655/0733-1347(2005)019%5B0001%3AGVISKO%5D2.0.CO%3B2">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Hecht, M. K., C. Kropach, and B. M. Hecht. 1974. Distribution of the yellow-bellied sea snake, <i>Pelamis platurus</i>, and its significance in relation to the fossil record. Herpetologica 30:387-396 <<a href="http://www.jstor.org/stable/3891437">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">McKeown, S. 1996. A Field Guide to Reptiles and Amphibians in the Hawaiian Islands. Diamond Head Publishing.</span><br />
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<span style="font-size: x-small;">Vasconcelos, R., J. C. Brito, S. Carranza, and D. J. Harris. 2013. Review of the distribution and conservation status of the terrestrial reptiles of the Cape Verde Islands. Oryx 47:77-87 <<a href="http://molevol.cmima.csic.es/carranza/pdf/Vasconcelos%20etal_2013_Oryx_Atlas%20CV_suppMat.pdf">link</a>></span><br />
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<span style="font-size: x-small;">Wynn, A. H., R. P. Reynolds, D. W. Buden, M. Falanruw, and B. Lynch. 2012. The unexpected discovery of blind snakes (Serpentes: Typhlopidae) in Micronesia: two new species of <i>Ramphotyphlops </i>from the Caroline Islands. Zootaxa 3172:39–54 <<a href="http://www.fs.fed.us/psw/publications/falanruw/psw_2012_falanruw001_wynn.pdf?">link</a>></span><br />
<span style="font-size: x-small;"><br /></span>
<span style="font-size: x-small;">Zug, G. R. 2013. Reptiles and Amphibians of the Pacific Islands: A Comprehensive Guide. University of California Press, Berkeley, California, USA <<a href="https://books.google.com/books/about/Reptiles_and_Amphibians_of_the_Pacific_I.html?id=3H65ngEACAAJ">link</a>></span><br />
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<span style="font-size: x-small;"><span property="dct:title" xmlns:dct="http://purl.org/dc/terms/">Life is Short, but Snakes are Long</span> by <a href="http://www.snakesarelong.blogspot.com/" property="cc:attributionName" rel="cc:attributionURL" xmlns:cc="http://creativecommons.org/ns#">Andrew M. Durso</a> is licensed under a <a href="http://creativecommons.org/licenses/by-nc-nd/3.0/deed.en_US" rel="license">Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License</a>.</span></div>
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</div>Andrew Dursohttp://www.blogger.com/profile/04720822623046554913noreply@blogger.com8