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Showing posts with label Thermophis. Show all posts
Showing posts with label Thermophis. Show all posts

Friday, October 30, 2015

Are there any countries without snakes?


Global distribution of all snake species combined
Public domain from Wikipedia
Terrestrial data from Ernst & Ernst (2011) and Cogger et al. (1998)
Sea snake data based on Campbell & Lamar (2004), Phillips (2002),
Ernst & Ernst (2011), and Spawls & Branch (1995)
Snakes are found in almost every country in the world, but there are a few places without wild1 snakes. Snake-free land generally falls into two categories: remote islands, mostly formed by volcanism or as atolls, 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 within the last 26,000 years and haven't been recolonized since (for example, there are snake fossils from northern Canada, where no snakes live now, from a time when it was much warmer). 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 (e.g., downtown Manhattan), although they once did.

Iceland

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 hot-spring snakes of Tibet (genus Thermophis) 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 (Vipera berus) in both Scotland (470 mi away) and Norway (600 mi away), both of which are separated by a great deal of very cold ocean.

Ireland

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 by the time snakes evolved (150 million years ago, in the late Jurassic-early Cretaceous) Ireland had separated from any other land mass, and has been connected on and off to this day. There is some debate 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 (Vipera berus), grass snakes (Natrix natrix), or smooth snakes (Coronella austriaca) 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.

Cape Verde

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 (Psammophis sibilans) on the island of Sal in a 1951 paper 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 home to 31 endemic lizard species, more than any other island chain in the Macaronesian region.

New Zealand

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 summarized in 1997, 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 pelagic sea snakes (Hydrophis platurus; formerly Pelamis platurus, also known as yellow-bellied sea snakes), a very widespread species that is infamous for vagrancy and recently made headlines when one washed ashore in Ventura County, California. The remaining 10% of records are of banded sea snakes (Laticauda colubrina), 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 L. colubrina was re-identified as a new species from New Caledonia, L. saintgironsi, by herpetologists revising the widespread Laticauda colubrina complex.

Map of pelagic sea snake records from New Zealand
(1837-1997)
From Gill 1997
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 H. platurus 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 H. platurus 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 L. colubrina 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 H. platurus "some distance" from the sea at Table Cape on the Mahia Peninsula, a January 1990 record of L. colubrina "well above" the high-tide line at Whangaruru Harbour, an April 1938 record of H. platurus 200 feet from the sea on a lawn at New Plymouth, and, most incredible, a September 1945 record of L. colubrina 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 H. platurus, which is almost incapable of moving on land, L. colubrina is reasonably good at terrestrial locomotion, which could explain the inland presence of these snakes. Alternatively, the author of the review paper suggested that the snakes could have been carried inland by birds.2

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 northeast3, 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 H. platurus  stop feeding at sea temperatures below 18°C and die at temperatures between 14.5 and 17°C (the average sea temperature in the coldest month in northern New Zealand is 16°C).

Kiribati

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 the most up-to-date and authoritative guide to the reptiles of the Pacific Islands. 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 pelagic sea snakes 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 (Laticauda colubrina), and possibly pelagic sea snakes as well. Additionally, there is a single record of an ornate reef seasnake (Hydrophis ornatus), 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.

Tuvalu

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 L. colubrina, 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 on one of its coins! 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 two unexpected species of endemic blindsnakes, Ramphotyphlops adocetus and R. hatmaliyeb, described in 2012 from two small islands, one in the eastern part of FSM and the other in the western part.

Nauru

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 H. platurus off its shores, and it also has what is likely an introduced species, the ubiquitous Indotyphlops braminus or Brahminy Blindsnake, 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 how widespread it is globally. The original native range of I. braminus 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.

Marshall Islands

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 authoritative guide to the reptiles of the Pacific Islands lists only I. braminus from the Marshall Islands, but other sources suggest that at least a few brown treesnakes (Boiga irregularis), infamously introduced to Guam, have been found there as well, and it's possible that H. platurus 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 L. colubrina, although an official page 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. braminus as native, considering that we don't know how long it's been there).

Vatican City

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 other wildlife seem to be pretty minimal, 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. Monaco, 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 an alternate definition of the word "snake" within its walls.

Cover of a joke book that's blank inside
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, Hawaii4, Wake Island, Johnston Atoll, Howland & Baker Islands, the Marquesas Islands, the Pitcairn Islands, Sala y Gomez, Isla Malpelo, St. Helena, the Faroe Islands, the Isle of Man, 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 Vipera berus gets above the Arctic Circle in Scandinavia), not to mention the Atlantic, Arctic, and Antarctic Oceans5.

In the course of the research I did for this post, I found many travel articles promoting the snakelessness of some of these places as overwhelmingly positive, as I'm sure it is for many ophidiophobic travelers. But, the risk that snakes pose is way, way smaller than the fear we have of them, 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 one of two native species extinct and the other hanging on thanks only to captive breeding and reintroduction efforts. St. Kitts & Nevis could lose its only native snake, the Saba or orange-bellied Racer (Alsophis rufiventris), and native snakes have gone extinct or become critically endangered on many other islands throughout the Pacific and Caribbean due to centuries of forest clearance, overgrazing, development, and the introduction of invasive species, not to mention the many continental snake species threatened by sprawling development and habitat fragmentation. So, please, let's keep this list from growing.



1 Given the growing popularity of herpetoculture, I'd be willing to bet that there are captive snakes in every country, although a few countries have stringent laws banning any captive snakes, including as pets as well as in zoos and research facilities.



2 Studies have shown that, 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.



3 Zug's Reptiles and Amphibians of the Pacific Islands lists Tokelau as having no snakes, not even sea snakes, but does not cover the Chatham or Kermadec Islands.



4 Hawaii has introduced Brahminy Blindsnakes and, unlike many Pacific Islands, it is known 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).



5 A study 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.


ACKNOWLEDGMENTS

Thanks to Kerry Nelson for doing some of the background research for this post as part of a discussion in the Wild Snakes: Education & Discussion Facebook group.

REFERENCES

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 <link>

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 <link>

Heatwole, H., S. Busack, and H. Cogger. 2005. Geographic variation in sea kraits of the Laticauda colubrina complex (Serpentes: Elapidae: Hydrophiinae: Laticaudini). Herpetological Monographs 19:1-136 <link>

Hecht, M. K., C. Kropach, and B. M. Hecht. 1974. Distribution of the yellow-bellied sea snake, Pelamis platurus, and its significance in relation to the fossil record. Herpetologica 30:387-396 <link>

McKeown, S. 1996. A Field Guide to Reptiles and Amphibians in the Hawaiian Islands. Diamond Head Publishing.

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 <link>

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 Ramphotyphlops from the Caroline Islands. Zootaxa 3172:39–54 <link>

Zug, G. R. 2013. Reptiles and Amphibians of the Pacific Islands: A Comprehensive Guide. University of California Press, Berkeley, California, USA <link>

Creative Commons License

Life is Short, but Snakes are Long by Andrew M. Durso is licensed under a Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License.




Wednesday, May 15, 2013

Tibetan Hot-spring Snakes


Everyone likes a good soak in a hot spring now and again, but imagine spending your whole life in one! Now imagine being the size of a pencil and unable to regulate your own body temperature, and you're doing a pretty good approximation of a Tibetan Hot-spring Snake (Thermophis). These tiny snakes reach only 2.5 feet in length and are found at fewer than ten sites on the Tibetan plateau in the Himalayan Mountains of south-central China, all above 14,000 feet elevation. For comparison, that's at least as tall as Mt. Rainier in Washington, Pike's Peak in Colorado, or Mont Blanc in the Alps. To cope with the cold, hot-spring snakes inhabit marshes, rivers, and rocky areas around sulfur-free hot springs, where they eat amphibians and fishes, including the dicroglossid frog Nanorana parkeri, the minnow Schizothorax oconnori, and elongate stone loaches in the genus Triplophysa. As you can see from this video, these are highly charismatic snakes.

Frank Wall
Hot-spring Snakes were first described in 1907 by a physician and herpetologist living in India named Frank Wall. Wall received specimens of this snake sent  from Tibet by Lieutenant F. M. Bailey, who reported that local people familiar with the snake told him that it could be found within half a mile of certain hot springs at any time of the year (although he stated that they did not enter the spring water, which has since been shown to be false). Wall was impressed by the altitude at which the snakes were found, which to date is still higher than any other snake known! Wall named the snake Natrix baileyi after Bailey, and in 1953 herpetologist Edmond Malnate moved it into a newly erected genus, Thermophis, meaning "heat snake" in Greek, giving it the name is has today: Thermophis baileyi. In 2008 a second species of Thermophis was discovered which differs slightly in scale characters and body proportions. Peng Guo of Yibin University named it Thermophis zhaoermii for preeminent Chinese herpetologist Zhao Ermi.

Just how remarkable these snakes are was not fully realized until recently. In the past, analyses of evolutionary relationships were limited to comparisons of morphological characteristics (for snakes, early taxonomists primarily relied on features of the scales and of the male reproductive organs, called hemipenes, to inform their hypotheses on how snakes were related to one another). Modern advances in molecular biology have enabled taxonomists to compare genetic sequences of related organisms and discover the intricate branching pattern of the evolutionary tree of life, essentially the family tree of all life on Earth. Although molecular phylogenetics, as this branch of science is called, is not flawless, it can provide incredible insight into the ancestry of species that have no close living relatives and therefore are very unique morphologically, making them difficult to compare with other organisms. Hot-spring Snakes are in this very situation, and although to a non-specialist they look pretty much like any other snake, their evolutionary history remained a mystery until 2009, when a group of biologists led by Zhao Ermi published two papers on the evolutionary origins of Thermophis.

Thermophis baileyi
As it turns out, Hot-spring Snakes are most closely related to South American snakes called xenodontines. Xenodontinae is one of the largest subfamilies of colubrid snakes, with about 90 genera and more than 500 species known. They are primarily tropical snakes previously thought to be restricted to the Americas, and they include several well-known (and many poorly-known) species, among them the South American Hog-nosed Snakes (genus Xenodon). Similarities of hemipenal morphology had hinted at a relationship between these taxa, but who would have guessed that the closest relatives of Hot-spring Snakes lived nearly 10,000 miles away on the tropical other side of the world? Not I, for one.

Thermophis baileyi
Hot-spring Snakes probably diverged from their "nearest" relatives about 28 million years ago. Despite the strengths of molecular phylogenetics, there is still some uncertainty about the position of Thermophis relative to other colubrid snakes because their branch of the tree arises near the base of a major clade (Xenodontinae), meaning that, as suspected, they have no close living relatives. In some phylogenies, Hot-spring Snakes are clustered with the "relict snakes of North America": CarphophisContia, Diadophis, Farancia, and Heterodon. Some of my favorite snakes, these are thought to have dispersed from Asia into North America during the Miocene, about 16 million years ago. (Diligent readers will recall that I've told this story before in my post on Rainbow Snakes, although I didn't know then about the involvement of Thermophis.)

Reproduced with permission from
Story in the Stone: The Formation of a Tropical Land Bridge
by Tom Gidwitz, illustration by David Stevenson & Greg Wenzel
Probably the common ancestor of all modern colubrids (Thermophis and NA relicts included) lived in Asia more than 30 million years ago. When the Bering Land Bridge connected North America and Asia, some of these snakes dispersed eastward across it, just like the ancestors of sabre-toothed tigers, woolly mammoths, and even Tyrannosaurus rex1. These evolved into a North American snake fauna, now largely extinct except for the few aforementioned relicts, and a hugely successful South American snake fauna, which was isolated from North America for a 5 million year period during the late Miocene-early Pliocene when the Isthmus of Panama was submerged by the ocean. One reason for this disparity is that two other groups of colubrid snakes, which are today the dominant colubrids of North America, the colubrines and the natricines, dispersed from Asia to North America around the same time as the xenodontines. Apparently ancestral colubrines and natricines dispersed more slowly than xenodontines, because they didn't reach South America before it separated. Instead, they only moved into South America following the most recent closing of the Isthmus of Panama in the late Pliocene, in an event known as the Great American Biotic Interchange. The GABI was responsible for allowing toads, treefrogs, opossums, armadillos, hummingbirds, and vampire bats to colonize North America, and salamanders, pit vipers, rabbits, squirrels, raccoons, deer, and jaguars (and colubrine and natricine snakes) to colonize South America. Assuming that Thermophis are all that's left of the original Asian proto-xenodontine snake stock, this pattern explains the evolutionary and biogeographic relationships of the Hot-spring Snakes and their relatives. However, given other recent discoveries in Asia, I wouldn't rule out the future discovery of another Asian proto-xenodontine more closely related to Thermophis than to any other known snake.

One reason we know only a little about Thermophis is its high mountain habitat. Most of the mountain ranges in China run east-west, but the Hengduan Mountains, where Hot-spring Snakes are found, stretch north-south (the name "Hengduan" means "to transect" and "cut downward" in Chinese). Parallel north-south sub-ranges of the Hengduans are separated by deep river valleys through which flow the famous Three Parallel Rivers: the Nujiang (Salween), Lantsang (Mekong), and Jinshajiang (Upper Changjiang or Yangtze). Thermophis baileyi is distributed west of the Salween, whereas T. zhaoermii is distributed east of the Changjiang. Geologic uplift of the intervening region of southern Tibet has lasted for about the last 20 million years, about the same age as the divergence between the two extant species of Thermophis. It is hypothesized that refuges in the Kyi Chu/Lhasa and Yarlung Zhangbo valleys during the last glacial maximum probably allowed T. baileyi to persist in the west, alongside such glacial relicts as neo-endemic ground beetles, juniper trees, and even humans. Following the end of the last Ice Age, they dispersed to other hot spring sites, and today connectivity among these sites is maintained when male snakes make rare movements among them, probably facilitated by the rivers and streams that connect the sites. Female snakes are less likely to disperse, because the plateau's short summers necessitate highly seasonal reproduction. Whether Thermophis are oviparous or viviparous is still unknown.

Sylvia Hofmann's photo of T. baileyi
made the cover of  the
Herpetological Bulletin
in 2007
Although the advantages of living around hot springs at high altitudes, where the temperature is relatively cold, are pretty obvious, recent surveys by Ding-qi Rao found that Hot-spring Snakes also live in fields and other areas far from hot springs, suggesting that the species' ecological niche may be wider than previously thought. This is fortunate, both because the growing exploitation of geothermal energy has led to destruction and degradation of hot spring habitats, and because global climate change will likely continue to cause mountaintop habitats around the world to shrink, necessitating a shift upward in elevation by high-altitude species in order to follow their habitat. This problem has been documented for pikas and for birds and will likely affect Hot-spring Snakes too. Because the ability of mountaintop species to disperse across intervening areas to higher mountain ranges is limited, many may go extinct. Will we one day see the top of Mount Everest as the last foothold for Hot-spring Snakes? Let's hope not.



1 Not all of these dispersal events happened at the same time. Evidence suggests that the Bering Land Bridge has connected North America with Asia several times over the last seventy million years: at least once during the time of the dinosaurs, again about 55 million years ago, another 20-16 mya, and more recently both 35,000 and 22-7,000 years ago. The ancestors of the New World xenodontines probably came across 20-16 million years ago.


ACKNOWLEDGMENTS

Thanks to photographers Kai Wang, Daniel Winkler, Sylvia Hofmann, Brian McDiarmant, and Gavin Maxwell for use of their photographs, and to Tom Gidwitz, David Stevenson, and Greg Wenzel for allowing me to reproduce their artwork.
REFERENCES

Guo, P, Liu S, Feng J, He M (2008) The description of a new species of Thermophis (Serpentes: Colubridae). Sichuan Journal of Zoology 27:321 <link>

Guo, P., S. Y. Liu, S. Huang, M. He, Z. Y. Sun, J. C. Feng, and E. M. Zhao. 2009. Morphological variation in Thermophis Malnate (Serpentes: Colubridae), with an expanded description of T. zhaoermii. Zootaxa 1973:51-60 <link>

He M, Feng J, Zhao E (2010) The complete mitochondrial genome of the Sichuan hot-spring keel-back (Thermophis zhaoermii; Serpentes: Colubridae) and a mitogenomic phylogeny of the snakes. Mitochondrial DNA 21:8-18 <link>

Hofmann S (2012) Population genetic structure and geographic differentiation in the hot spring snake Thermophis baileyi (Serpentes, Colubridae): indications for glacial refuges in southern-central Tibet. Molecular Phylogenetics and Evolution 63:396-406 <link>

Hofmann S, Fritzsche P, Solhøy T, Dorge T, Miehe G (2012) Evidence of sex-biased dispersal in Thermophis baileyi inferred from microsatellite markers. Herpetologica 68:514-522 <link>

Huang S, Liu S, Guo P, Zhang Y, Zhao E (2009) What are the closest relatives of the hot-spring snakes (Colubridae, Thermophis), the relict species endemic to the Tibetan Plateau? Molecular Phylogenetics and Evolution 51:438-446 <link>

Pinou, T., S. Vicario, M. Marschner, and A. Caccone. 2004. Relict snakes of North America and their relationships within Caenophidia, using likelihood-based Bayesian methods on mitochondrial sequences. Molecular Phylogenetics and Evolution 32:563-574 <link>

Sekercioglu, C. H., S. H. Schneider, J. P. Fay, and S. R. Loarie. 2008. Climate change, elevational range shifts, and bird extinctions. Conservation Biology 22:140-150 <link>

Wall, F. 1907. Some new Asian snakes. The Journal of the Bombay Natural History Society 17:612-618 <link>

Creative Commons License

Life is Short, but Snakes are Long by Andrew M. Durso is licensed under a Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License.