The 9,999th Reptile

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Number of new snake species by decade, with highlights
Data from The Reptile Database

Linnaeus's 1758 Systema Naturae, the starting point of zoological nomenclature, described 118 species of reptiles, including 74 snakes (not counting the limbless lizards and amphibians he included in the same group). It took over 100 years for the number of described species of snakes to reach 1000, an event that probably passed without much notice amid the American Civil War. Since that time, new snake species descriptions have been added at the rate of about 15 a year, although molecular taxonomy has increased this pace over the last few decades. The trends for snakes and for reptiles as a whole have been similar, and on July 9th, 2014, a team of American, German, Lao, and Vietnamese scientists described a new species of gecko from Laos, which the journal Herpetological Review reported as the 10,000th reptile species. Needless to say, I was excited, but I was also extremely disappointed because I had been so hoping that it would be a snake! Rather than admit defeat and scrap this planned post, I emailed Peter Uetz at The Reptile Database, an incredible resource that I've praised before, to confirm that this gecko was indeed #10,000. As usual for taxonomy and as I should have suspected, the reality was a bit more complicated.

Although Cyrtodactylus vilaphongi was the 10,000th reptile species for a while, the order and position of entries in The Reptile Database is constantly changing. Although new species get added to the end of the list, it's common for two or more existing species to get synonymized or merged, which moves the position of all subsequent species up. Furthermore, sometimes species that were described long ago and subsequently synonymized are revalidated, leading to 'new' species that aren't really new in the sense that they have existed before. Finally, often existing species get split up, leading to additions that aren't as dramatic as legitimate new discoveries. This last complication is on the rise now that molecular systematics has enabled us to describe the cryptic diversity of some lineages, which are not all that morphologically distinct but may contain considerable genetic diversity.

At the time of my email to Peter last month, C. vilaphongi was the 9988th species, and (happily), a new snake, Siphlophis ayauma, was #10,000. Although this has probably changed again by now, I'm going to operate under the assumption that, since we can't really say with certainty that any particular species was #10,000, if it was a snake, it was probably one of the 11 brand new snake species that have been described so far this year. You can read about many of these on the blog 'Species New to Science', but I'm going to highlight them in a little more detail here.

Rhabdophis guangdongensis
From Zhu et al. 2014

The first new snake described this year, Rhabdophis guangdongensis, was collected by a team of Chinese herpetologists in Guangdong Province in 2008. The reason it wasn't described until the February 20th issue of the journal Zootaxa is because, as is often the case, it takes a couple of years to compare both the anatomy and the DNA of a suspected new species to reference specimens of known similar species and establish that the species really is new. In the past, particularly prior to the internet, the difficulty of doing this was a huge problem, resulting in close to half of all 'new' species later being invalidated as duplicates. The genus Rhabdophis  is distributed in southern and eastern Asia, and this is the 21st species. It's an extremely interesting genus from a chemical ecology perspective, because at least one species sequesters defensive chemicals from its prey and provisions them to its young (which I wrote about for Scientific American shortly after I started this blog). A recent paper by Yosuke Kojima and Akira Mori on the Japanese species R. tigrinus showed that females periodically leave wetlands for forest streams where they forage on toads, likely to obtain the necessary toxins for provisioning their offspring. The new species also has specialized structures, known as nuchal glands, on the back of its neck, so presumably it stores bufotoxins there as well, although this has yet to be verified.

Opisthotropis durandi
From Teynié et al. 2014

On March 3rd, a team of French, German, and Vietnamese scientists published a description of Opisthotropis durandi, a highly aquatic snake collected from the base of a waterfall in northern Laos. This is the seventh species of Opisthotropis described in the past 20 years, and the first from Laos (although other species are likely to occur there based on their occurrence in surrounding countries). Like the new Rhabdophis, it is also the 21st species in its genus. It is important to realize that, like most species new to science, this snake was already known by local people. It is called Ngou Koung or Ngou Kung, meaning “shrimp snake”, suggesting that it may eat shrimp. The pools at the base of the waterfall where the first specimen was found contained many small shrimp.

Eutrachelophis bassleri and its weird penis
From Myers & McDowell 2014
A color photo of E. bassleri was published in Echevarría & Venegas 2015

Harvey Bassler, a petroleum geologist, explored many of the Amazon's upper tributaries for his work during the 1920s and 30s, during which time he collected over 4,200 snakes on the side. Bassler deposited his magnificent collection in the American Museum of Natural History in 1934, and on March 6th this year Charles Myers and Samuel McDowell published a monograph in the Bulletin of the American Museum of Natural History describing a species of snake collected by Bassler in 1927, for which they erected a new genus, Eutrachelophis (‘beautiful-necked snakes’’). They also placed in this genus a species originally described by Boulenger in 1905, Rhadinaea steinbachi, which they renamed Eutrachelophis steinbachi. Although the two species (and a third, yet undescribed) have very similar skeletons, muscles, glands, viscera, and markings, they probably would have been placed in separate genera had they been described in the 19th or early 20th century because their hemipenes are so different. E. steinbachi has long but relatively normal-looking hemipenes, whereas E. bassleri  has extremely unusual heimpenes tipped with a dome-like structure so strange (at least within the world of snake hemipenes) that the authors wrote "we have seen nothing quite like [it]." Hemipenes were traditionally considered one of the most taxonomically-important structures in snakes¹ because they were considered to be evolutionarily neutral (that is, unlikely to change in response to selection), but a growing awareness that evolution by both natural and especially sexual selection can influence the morphology of male genitalia led these authors to recognize that these two snakes were in fact close relatives. Although we await molecular confirmation, the authors propose a mechanism by which differential expression of Hox genes² could cause such a rapid divergence in hemipenal morphology between two sister species.

Siphlophis ayauma
From Sheehy et al. 2014

On January 12th, 2008, a group of American and Ecuadorian herpetologists stopped for lunch at a grilled-chicken restaurant in Paute, Azuay province, Ecuador. They noticed a peculiar sun-faded snake on display in a jar of alcohol that they couldn't quite put a name to. Following negotiation with the restaurant owner, the specimen was acquired and determined to belong to the genus Siphlophis, but could not be identified to any known species. A few months later, another specimen was found alive about 100 miles to the north, and two more were discovered in 2011 about the same distance to the south. A fifth individual is now recognized to have been hiding out unnoticed in the collection of the Museo de Zoología, Pontificia Universidad Católica del Ecuador. Because of its red-banded head and its occurrence in the mountains near cold (achachay) streams, the new species was named Siphlophis ayauma after the Kichwa spirit Aya Uma, a good spirit devil who derives strength from nature, particularly from cold mountain pacchas (cascades) and is represented in Kichwa folklore as having a colorful red-banded head. This is the seventh species in the genus, the third species known from Ecuador, and the first new species of Siphlophis since 1940. The results are published in the April 1st issue of the South American Journal of Herpetology.

Philodryas amaru
From Zaher et al. 2014

In a montane grassland high in the Andes Mountains of southern Ecuador, another genus gained its 21st species this year: Philodryas amaru. Known to the Ecuadorian and Brazilian authors since 2005, a small population of these striped racers was formally described in Zootaxa on April 4th this year. The new species resembles Philodryas simonsii in color pattern, but differs noticeably in its hemipenis morphology. "Amaru" means "snake" in Kichwa, and is also the name of a snake deity who influences water and the economy. This diurnal snake lays clutches of 9-13 eggs underground in galleries and under decaying logs, and probably eats frogs and lizards. It is a close relative of the Galapagos racers that I've written about before.

Causus rasmusseni
From Broadley 2014; photo by Paul L. Lloyd

Night adders (genus Causus) are a small and unusual group of vipers found in sub-Saharan Africa. They were once thought to be the most primitive vipers and were placed in their own subfamily, but they are now grouped with the viperines even though they have a plethora of unusual features: platelike head scales, round pupils, a different hinge mechanism for their erectile fangs, incomplete fang canal closure, and elongate venom glands in most species. On April 25th of this year, Don Broadley³ described the first new species of Causus since 1905. He named it Causus rasmusseni after the late Jens Rasmussen, a Dutch expert on African snakes who died in 2005. This species is found only in the watershed between the Congo and Zambezi basins, where it co-occurs with three other species of Causus. Broadley first became aware that there might be a new species of night adder in this region in 1991, when he noticed pale gray C. rhombetaus from northwestern Zambia with black markings and low ventral scale counts. In 2013, someone sent him a picture of one eating a toad (another unusual adaptation that night adders share with several other snakes), which prompted him to look again at the unusual specimens and describe them as a new species. Few molecular data are available for Causus, so this diagnosis is based on morphology alone.

Micrurus potyguaraFrom Pires et al. 2014

Brazil is graced with nearly 400 species of snakes, including 30 of the world's ~80 species of coralsnakes. The morphology of coralsnakes is highly variable, and there are many misidentified specimens in museum collections, so it is often difficult to recognize new species. A group of Brazilian herpetologists working on the tri-colored coralsnakes from the endangered northeastern coastal forests discovered a new species, which they described in the June 5th issue of Zootaxa (if any of these dates are your birthday, then you share a birthday with that of a new species of snake!).

Top: Jaw of Lycodon aulicus
From Jackson & Fritts 2004
Middle: Lycodon zoosvictoriae
From Neang et al. 2014
Bottom: Lycodon cavernicolus
From Grismer et al. 2014

Wolfsnakes (genus Lycodon) are named for their fearsome-looking fang-like anterior maxillary teeth. Unlike the true fangs of vipers, elapids, and atractaspidids, wolfsnake teeth are not grooved or hollow and they have no venom. Instead, their strongly arched upper jaw helps them feed on skinks, whose hard, cylindrical bodies fit snugly into their diastema, or the gap between their anterior and posterior teeth. The wolf-like anterior teeth keep the skink from being squeezed out of the mouth, while the posterior teeth slice through the skink's cycloid scales. At least 16 of the nearly 60 species of Lycodon have been described since the 1990s, including two this June: Lycodon zoosvictoriae from the Cardamom Mountains of southwestern Cambodia, and L. cavernicolus from a limestone cave in peninsular Malaysia. The latter is a cave-adapted species, both specimens of which were found climbing several feet above the cave floor, in total darkness. It's likely that they eat a cave-adapted gecko. Many of the caves in this region are in immediate danger of being quarried for cement before their endemic fauna and flora can be fully documented. Both of these species were also described in Zootaxa, which is a relatively new journal dedicated almost exclusively to rapid publication of new species descriptions, with the stated goal of aiding conservation efforts by circumventing the lengthy delays normally associated with publication of new science. Since its inception in 2001, Zootaxa has become a daily journal that has published nearly one quarter of all new animal taxa and nomenclatural acts in the last five years, including over 400 new species of reptiles and the 7000th species of amphibian.

"Cloudogram" of Crotalus triseriatus species group
showing the new nine-species arrangement
From Bryson et al. 2014

Just three days before the new gecko, a team of scientists from Mexico, the USA, and Canada published a genetic analysis of the Crotalus triseriatus species group, which contains small montane rattlesnakes found in Mexico and the southwestern USA. Although five species were historically recognized within the group, an analysis of seven nuclear genes revealed that there are at least nine species, including two that were previously recognized as subspecies and two more that have not heretofore been formally recognized. The paper described the two new species: Crotalus tlaloci, named for Tláloc, the Aztec god of rain, and Crotalus campbelli, named for herpetologist Jonathan Campbell. The authors of this paper suggest that these rattlesnakes speciated rapidly from a single common ancestor during the uplifting of the Trans-Mexican Volcanic Belt near the end of the Neogene period 2.6 million years ago, which makes sense because they are not very mobile and populations of their common ancestor likely would have become isolated from one another  on various "Sky Islands" of suitable habitat during the genesis of this new mountain range. Many species are endemic to the high-altitude pine-oak forests and grasslands of this region, which has become famous as the overwintering grounds of the Monarch Butterfly.

Chironius diamantina
From Fernandes & Hamdan 2014

Surprise! Just when you thought we were through, at press time the description of four more new species of snake had just been published, all from relatively recent issues of Zootaxa. One is a Brazilian species of Chironius, one of my favorite genera. Chironius diamantina is the 16th species in the genus, which is unusual is having a very low, even number of dorsal scale rows (10 or 12), the central pair of which are strongly keeled, giving the snake a distinctly flat-backed appearance. This species is found in riparian forests along rocky streams in coastal Brazil, not too far south of the new coralsnake (above). Chironius are diurnal and generally eat birds and mammals. Another is a new Asian keelback, Herpetoreas burbrinki, from near the border of China, India, and Burma. which is relatively closely related to the Rhabdophis above. Finally, two new species from the large ground-dwelling Latin American genus Atractus, both small and described from single specimens collected decades ago in Colombia (perhaps they will one day be rediscovered). More new species from both of these groups will likely follow, given the taxonomic untidiness of their genera. [Update: shortly after publication David Salazar-Valenzuela alerted me to the fact that I had missed his description, with colleagues, of a third new Atractus from the cloud forests of northern Ecuador earlier this year, in the journal Herpetologica. They mention that some of the specimens were collected from under logs alongside an undescribed species of slender blindsnake of the genus Trilepida, so it seems we are at 3,500 this year without a doubt!] [[Update II: It seems I missed more than I thought - a new species of Trimeresurus from Sumatra was described in September from specimens collected in 1899, and a new Ninia from Trinidad was described in August from a 1988 specimen.]]

In addition to these 15 species, there are a couple of species of snake which were described long ago but that were revalidated recently, including several scolecophidians (Typhlops silus, first described in 1959; Afrotyphlops angeli, first described in 1952; and Letheobia acutirostrata, first described in 1916) and a rattlesnake (Crotalus armstrongi, originally described as a subspecies in 1979 and elevated by the same group that described C. tlaloci and C. campbelli). These are typically announced with less fanfare than the truly new descriptions that I've highlighted above.

Although it's actually been the slowest year for new snakes since 1997⁴, we have 15 new snakes this year, bringing snakes to a total of 3,499 (and 2014 isn't over yet!). We could make it to 3,500 snakes in the same year that we hit 10,000 reptiles. I think these milestones in taxonomy emphasize the importance of reptiles and how much we have left to learn about them. I doubt that the pace of new species descriptions will slow down anytime soon, as experts estimate that less than 15% of the species on Earth have yet been described. Increasingly, reptiles, and snakes in particular, are becoming poster-children for biodiversity and conservation, a welcome change from their history of being overlooked and maligned. Soon, we will have high-quality global range maps for all species of reptiles, an achievement reached some time ago by amphibians, mammals, and birds, which will enable their incorporation into global assessments of vertebrate diversity and conservation planning. It's an exciting time.

For a complete list of all 24 snake species eventually described in 2014, click here.

ACKNOWLEDGMENTS

Thanks to Peter Uetz at The Reptile Database for sharing with me some inside information, and to the authors of these papers for their photos.

REFERENCES

Newspaper clipping from 10 January 1960
showing Broadley with his amputated finger.
You can see more at the finger's Facebook page
or listen to Broadley describe the experience here.

Angarita-Sierra, T. 2014. Hemipenial Morphology in the Semifossorial Snakes of the Genus Ninia and a New Species from Trinidad, West Indies (Serpentes: Dipsadidae). South American Journal of Herpetology 9:114-130 <link>

Broadley, D. G. 2014. A new species of Causus Lichtenstein from the Congo/Zambezi watershed in north-western Zambia (Reptilia: Squamata: Viperidae). Arnoldia Zimbabwe 10:341-350 <link>

Bryson, R. J., C. W. Linkem, M. E. Dorcas, A. Lathrop, J. M. Jones, J. Alvarado-Diaz, C. I. Grunwald, and R. W. Murphy. 2014. Multilocus species delimitation in the Crotalus triseriatus species group (Serpentes: Viperidae: Crotalinae), with the description of two new species. Zootaxa 3826:475-496 <link>

Cope, E. D. 1895. The classification of the Ophidia. Transactions of the American Philosophical Society 18:186-219 <link>

Dowling, H. G. 1967. Hemipenes and other characters in colubrid classification. Herpetologica 23:138–142 <link>

Grismer, L. L., E. S. H. Quah, S. Anuar, M. A. Muin, P. L. Wood Jr, and S. A. M. Nor. 2014. A diminutive new species of cave-dwelling Wolf Snake (Colubridae: Lycodon Boie, 1826) from Peninsular Malaysia. Zootaxa 3815:51-67 <link>

Guo, P., Q. Liu, L. Zhang, J. X. Li, Y. Huang, and R. A. Pyron. 2014. A taxonomic revision of the Asian keelback snakes, genus Amphiesma (Serpentes: Colubridae: Natricinae), with description of a new species. Zootaxa 3873:425-440 <link>

Fernandes, D. and B. Hamdan. 2014. A new species of Chironius Fitzinger, 1826 from the state of Bahia, Northeastern Brazil (Serpentes: Colubridae). Zootaxa 3881:563-575 <link>

Trimeresurus gunaleni
From Vogel et al 2014

Jackson, K. and T. H. Fritts. 2004. Dentitional specialisations for durophagy in the Common Wolf snake, Lycodon aulicus capucinus. Amphibia-Reptilia 25:247-254 <link>

Köhler, G. and M. Kieckbusch. 2014. Two new species of Atractus from Colombia (Reptilia, Squamata, Dipsadidae). Zootaxa 3872:291-300 <link>

Linnaeus, C. 1758. Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. [10th ed.]. Laurentii Salvii, Holmiae, Stockholm, Sweden <link>

Myers, C. W. and S. B. McDowell. 2014. New Taxa and Cryptic Species of Neotropical Snakes (Xenodontinae), with Commentary on Hemipenes as Generic and Specific Characters. Bulletin of the American Museum of Natural History 385:1-112 <link>

Neang, T., T. Hartmann, S. Hun, N. J. Souter, and N. M. Furey. 2014. A new species of wolf snake (Colubridae: Lycodon Fitzinger, 1826) from Phnom Samkos Wildlife Sanctuary, Cardamom Mountains, southwest Cambodia. Zootaxa 3814:68-80 <link>

Pires, M. G., N. J. da Silva Jr., D. T. Feitosa, A. L. d. C. Prudente, G. A. P. Filho, and H. Zaher. 2014. A new species of triadal coral snake of the genus Micrurus Wagler, 1824 (Serpentes: Elapidae) from northeastern Brazil. Zootaxa 3811:569-585 <link>

Atractus savagei
From Salazar-Valenzuela et al. 2014

Salazar-Valenzuela, D., O. Torres-Carvajal, and P. Passos. 2014. A New Species of Atractus (Serpentes: Dipsadidae) from the Andes of Ecuador. Herpetologica 70:350-363 <link>

Schneider, N., T. Q. Nguyen, M. D. Le, L. Nophaseud, M. Bonkowski, and T. Ziegler. 2014. A new species of Cyrtodactylus (Squamata: Gekkonidae) from the karst forest of northern Laos. Zootaxa 3835:80-97 <link>

Sheehy, C. M., M. H. Yánez-Muñoz, J. H. Valencia, and E. N. Smith. 2014. A new species of Siphlophis (Serpentes: Dipsadidae: Xenodontinae) from the eastern Andean slopes of Ecuador. South American Journal of Herpetology 9:30-45 <link>

Teynié, A., A. Lottier, P. David, T. Q. Nguyen, and G. Vogel. 2014. A new species of the genus Opisthotropis Günther, 1872 from northern Laos (Squamata: Natricidae). Zootaxa 3774:165-183 <link>

Uetz, P. 2010. The original descriptions of reptiles. Zootaxa 2334:59-68 <link>

Vogel, G., P. David, and I. Sidik. 2014. On Trimeresurus sumatranus (Raffles, 1822), with the designation of a neotype and the description of a new species of pitviper from Sumatra (Squamata: Viperidae: Crotalinae). Amphibian and Reptile Conservation 8:1–29 <link>

Zaher, H., J. C. Arredondo, J. H. Valencia, E. Arbeláez, M. T. Rodrigues, and M. Altamirano-Benavides. 2014. A new Andean species of Philodryas (Dipsadidae, Xenodontinae) from Ecuador. Zootaxa 3785:469–480 <link>

Zhu, G.-X., Y.-Y. Wang, H. Takeuchi, and E.-M. Zhao. 2014. A new species of the genus Rhabdophis Fitzinger, 1843 (Squamata: Colubridae) from Guangdong Province, southern China. Zootaxa 3765:469-481 <link>

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

What the State Snakes Should Be: Part I

Click here to read this post in Spanish!
Haga clic aquí para leer este blog en español!

The 50 United States have various state symbols, including state trees, flowers, songs, drinks, and even fossils. Just over half the states have a state reptile, and none have a state snake (although a few of the state reptiles are snakes). This post was inspired by the witty and spot-on post 'The State Birds: What They SHOULD Be' from thebirdist.com, which struck a chord with me because I've always been underwhelmed by the state birds, many of which are uncreative and absurd. The selection of state reptiles has been characterized by the same lack-of-creativity, bordering on willful ignorance of a state’s unique and endemic biodiversity, as the selection of state birds. In fact, it's probably worse, since most people can at least name several different kinds of birds whereas even distinguishing reptiles from amphibians or fishes (or worms) is apparently beyond many. Perhaps having a state snake would help remedy this problem as well as improve upon the massive and undeserved PR problem faced by snakes in general.

At the risk of catalyzing a scandal on par with my home state of North Carolina's State Fruit Fiasco of 2001, which in an effort not to alienate any of several fruit growing lobbies resulted in an official state red berry (the strawberry), state blue berry (wait for it…the blueberry), and state fruit (the scuppernong grape), I've taken the liberty of choosing a state snake for each of the 50 U. S. states, 44 of which I've been to and almost half of which I've caught snakes in. This was surprisingly hard to do, because some snakes have too many good choices and others have too few. I used the same rules as thebirdist: I didn't repeat any species. Feel free to chime in with your opinion about what your state's snake should be, if it differs from my choice.

1. Alabama. Eastern Diamond-backed Rattlesnake (Crotalus adamanteus)

Crotalus adamanteus

The Eastern Diamondback is iconic in the southeast, appearing on America's first flag, the Gadsden flag (better known as the 'Don't Tread on Me' flag) and in the first political cartoon appearing in an American newspaper (Ben Franklin's 'Join, or Die'). State symbolism of the world's largest rattlesnake might help abate pressure from rattlesnake roundups in Alabama and other states, which have caused declines so serious that this species has been considered for listing under the Endangered Species Act.

2. Alaska. Northwestern Gartersnake (Thamnophis ordinoides)

Thamnophis ordinoides

Alaska's only got one snake, and I do mean one: a road-killed juvenile gartersnake was found outside of Haines, Alaska, in August of 2005. The poor condition of the specimen prevented a positive identification based on morphology alone, so some of my colleagues including my fellow USU graduate student Lori Neuman-Lee sequenced genes from the snake and identified it as a T. ordinoides. This highly variable species eats mostly slugs. Because the closest known population of Northwestern Gartersnakes is over 600 miles south of Haines, the snake might have been transported there accidentally. However, reports of gartersnakes on the banks of the Taku and Stikine rivers in Alaska, including one ostensibly vouchered (now lost) specimen, keep the door open for future discovery of a naturally occurring population of snakes in The Last Frontier. Don't lose hope, Alaska! If nothing else, gartersnakes from British Columbia will probably disperse there eventually if climate change keeps up the way it's been going.

3. Arizona. Arizona Ridge-nosed Rattlesnake (Crotalus willardi)

Crotalus willardi

This actually is the state reptile of Arizona! Way to go, Arizona! This snake is a phenomenal choice for its beauty and uniqueness. Principally a Mexican species, it is found in scattered, isolated "Sky Island" mountain ranges separated by uncrossable desert. Former Staten Island Zoo reptile curator and herping godfather Carl Kauffeld, in his classic 1957 memoir Snakes and Snake Hunting, called this species "sublime" and said of finding one that "no greater triumph is possible". It was the last rattlesnake species discovered in the United States, in  1905 (just as the "Baby State," as it was once known, was one of the last states to enter the union, seven years after C. willardi was discovered). These small rattlesnakes grow to be only one to two feet long and as a result are relatively unthreatening. Few bites have been recorded, none serious, although I wouldn't go picking one up. Runner up: Glossy Snake (Arizona elegans), which shares its genus name with the state.

4. Arkansas. Cottonmouth (Agkistrodon piscivorus)

Agkistrodon piscivorus

When I asked my colleague Geoff Smith, a native Arkansan, about snakes iconic to Arkansas, he first informed me that, as someone born above the Arkansas River, he was an Arkansawyer rather than an Arkansan. He went on to suggest that the Cottonmouth was a good choice because every snake someone sees in Arkansas is identified as a Cottonmouth (a joke that is all too true, poor Nerodia). He also suggested some similarities between Arkansans and Cottonmouths: they stink¹, they're fat, they like to eat fish. I decided not to contradict him because 1. I haven't been to Arkansas, and 2. he's from there so he can say that. In the end, we decided that a generalist state needed a generalist snake. Because Arkansas' diverse landscapes encompass the range edges of many species that are more characteristic of other places (for example, Queensnakes, Coralsnakes, and Western Diamond-backed Rattlesnakes can all be found in the state), choosing a widespread generalist is appropriate. I know of few better examples than the Cottonmouth, a denizen of river swamps that eats a variety of other vertebrates. Furthermore, Cottonmouths might have the worst PR problem of any snake: in addition to being misidentified, they suffer from exaggerated accounts of their aggression, size, breeding habits and even basking behavior ("A Water Moccasin tried to get in my boat!"). State snakehood might help with that.

5. California. Giant Gartersnake (Thamnophis gigas)

Thamnophis gigas

This largest of gartersnakes is endemic to California's Central Valley, where it historically occupied tule marshes but today persists mostly in rice fields and their associated canals and drains, where it mostly eats introduced fishes and Bullfrogs. Because of the massive agricultural use of this area, this species has lostmuch of its habitat and is on the edge of extinction, so it could use some positive PR. It is highly aquatic, with habits more similar to eastern Nerodia watersnakes than to its  western congeners - the same could be said of many Golden State residents in terms of their political opinions and attitudes.

6. Colorado. Great Plains Ratsnake (Pantherophis emoryi)

Pantherophis emoryi

Once considered a western subspecies of the Cornsnake, Great Plains Ratsnakes were re-elevated to full species status in 2002 by College of Staten Island snake taxonomist Frank Burbrink. Although Colorado's majestic landscapes are home to many beautiful snakes, not one but two geographically-separated forms of Great Plains Ratsnakes are found in the state: the larger, more brightly colored nominate subspecies in the high plains and tablelands of southeastern Colorado, and the smaller, drabber Intermountain Ratsnake (P. e. intermontanus) on the Colorado Plateau in the west-central part of the state. Originally named by Baird & Girard for surveyor and U.S. Army officer William H. Emory, these large constrictors are more subtly colored than other ratsnakes but just as harmless and beautiful.

7. Connecticut. Northern Watersnake (Nerodia sipedon)

Nerodia sipedon

Northern Watersnakes are some of the most common snakes in eastern North America. Found in almost every aquatic habitat, they are frequently seen basking on fallen logs, from which they will drop into the water at the slightest provocation. Leo Finneran reported this species from the Branford area in the 1940s, saying that it was "a very common snake occurring along water bodies in all parts of the town" and noting that some carried heavy loads of parasites, which is common in watersnakes. You can remember how to tell this species from its southern counterpart because the bands across its back fail to connect near the tail. OK, I'm stretching a bit for this one. Keep reading, it gets better.

8. Delaware. Copperhead (Agkistrodon contortrix)

Agkistrodon contortrix

Delaware was the first state, and the Copperhead is the first snake you should learn to identify if you live within its range. Delaware's Brandywine Valley is reminiscent of New England, whereas its middle and lower parts resemble the tidewater South. Both regions are inhabited by Copperheads, which are found from New York to Texas, where they are the most common venomous snakes in many suburban areas. Due to the high encounter rate, more than a third of U. S. venomous snakebites are from Copperheads. Fortunately, they seldom require antivenin, because they have the least potent venom of any North American viper, and the fatality rate is negligible. Furthermore, the vast majority occur when someone deliberately attempts to handle or kill the snake. In this way, Copperheads and their relatives are distinctly "antiwar," eponymous as they are of the vocal antebellum Democrats that once represented Delaware and other Union states in Congress.

9. Florida. Eastern Coralsnake (Micrurus fulvius)

Micrurus fulvius

Florida as a state is unlike any other U. S. state. Coralsnakes are unlike any other North American snakes - they are more closely related to cobras and sea snakes, which live on other continents or in the oceans. They're also brightly colored, like the iconic pink flamingos often seen on Sunshine State lawns. Coralsnakes spend most of their time underground and are difficult to find. When asked how to see one in the wild, most herpers reply: "Go to Florida." Although Coralsnakes are quite venomous, they are secretive and non-aggressive, so very few Coralsnake bites ever occur. Runners up: Short-tailed Snake (Lampropeltis [Stilosoma] extenuatum) or Rim Rock Crowned Snake (Tantilla oolitica), both endemic to the state.

10. Georgia. Eastern Indigo Snake (Drymarchon couperi)

Drymarchon couperi

The largest snakes in North America and literal "lords of the forest" (from the Greek drymos: forest and archos: commander), Eastern Indigo Snakes are in decline due to habitat loss and fragmentation. Georgia boasts some of the best remaining indigo habitat (both Fort Stewart and Fort Benning harbor large, unfragmented tracts of longleaf pine savanna with healthy populations of Gopher Tortoises, the actual Peach State reptile and the burrows of which these snakes depend upon) and the headquarters of the Orianne Society, a non-profit formed to help conserve indigo snakes and other rare reptiles.

11. Hawaii. Yellow-bellied Sea Snake (Pelamis platura)

Pelamis platura

This is the Aloha State's only native snake, making it a pretty easy choice. It's also awesome! Yellow-bellied Sea Snakes are the only pelagic snakes, and the most widely distributed and specialized of the sea snakes. They are most commonly seen in Hawaiian waters in El Niño years, when average water temperatures are in the upper 70s. Pelamis have fast-acting venom that they use to kill fishes. They shed their skin often to remove barnacles, using a knotting behavior to compensate for the absence of objects to brush against in the open ocean. Paying homage, a Scottish tidal energy company adopted the genus name of this snake for both their company and the offshore wave energy converter itself, although the company closed its doors in 2014.

12. Idaho. Rubber Boa (Charina bottae)

Charina bottae

I've written about Rubber Boas before, but man are they cool. Some of the best research on Rubber Boa thermal biology and behavior was conducted in southeastern Idaho, and Craters of the Moon National Monument is a great place to see them. Incredibly, these snakes are active at temperatures as low as 40°F, which is good news for them given that they live in Idaho's mountains. Roughly the same color as a potato, Rubber Boas mostly eat small mammals, which they kill by constriction similar to their larger relatives, which include Anacondas and Boa Constrictors. As is fitting for the Gem State, their small, smooth scales have a rubbery shine.

13. Illinois. Prairie Kingsnake (Lampropeltis calligaster)

Lampropeltis calligaster

Illinois, like Arkansas, contains many ecoregions but is not dominated by any one that isn't more iconic of elsewhere. There are actually more species of snakes in Illinois than in many southern and eastern states, due to its mixture of northern, southern, eastern, and western ecosystems. I lived here for a while, but it was hard to pick a snake for Illinois because its habitats are so varied. Prairie Kingsnakes are common along Illinois roads and lake shores in spring. Unfortunately, many are run over by vehicles during this time, and still more are mistaken for venomous snakes are maliciously killed. In fact, these large and harmless constrictors are highly beneficial, consuming mice and voles that would otherwise overrun farms, gardens, and homes. In the agricultural desert of the Prairie State, Prairie Kingsnakes are one species that seems to do reasonably well.

14. Indiana. Eastern Ribbonsnake (Thamnophis sauritus)

Thamnophis sauritus

The slender, long-tailed ribbonsnakes share a genus with gartersnakes, from which they are not terribly different. Ribbonsnakes show a preference for wetter habitats and eat predominantly fishes and amphibians. Indiana harbors two subspecies of Eastern Ribbonsnakes, which together are found essentially throughout the state where suitable habitat still exists. Because of the Hoosier State's intensive agricultural land use, a state snake that requires intact wetlands with healthy prey populations could encourage a shift towards sustainable agricultural practices that use minimal pesticide and fertilizer and promote healthy waterways.

15. Iowa. Western Foxsnake (Pantherophis ramspotti)

Pantherophis ramspotti

These large constrictors used to be considered the same species as Eastern Foxsnakes, Pantherophis vulpinus, but were elevated to full species status by Brian Crother and colleagues in 2011. Rapid northward expansion by Foxsnakes from refugia in various southern states following the last glacial maximum has resulted in their odd modern distribution: two species apparently separated not by the gap in the range of the eastern in Michigan but by the Mississippi River. Because they no longer occupy any of the habitat where their fossils are found, it seems that Foxsnakes simply filled in the new habitat made available by the retreat of the glaciers. They had few barriers since the glaciers flattened the topography, and few states beat the Hawkeye State for flat.

16. Kansas. Milksnake (Lampropeltis triangulum)

Lampropeltis triangulum

The Red Milksnakes of Kansas are like jewels. Essentially fancy kingsnakes, these beauties are easy to find underneath large, flat rocks in prairies and fields almost throughout the state. They eat lizards as well as small mammals and other snakes, which should be easy for them to find since they all live under the same rocks. The late herpetologist extraordinaire Henry Fitch undertook detailed natural history studies of nearly every Kansas snake during his long and productive career, publishing hundreds of articles of the highest caliber, and his 1970 paper on milksnake ecology and natural history is no exception. About as popular as a snake can get, the Red Milksnake (L. t. syspila) is an easy choice for the state snake of the Sunflower State. All those states within the range of the Scarlet Kingsnake (L. t. elapsoides) only wish they weren't also home to so many other strong contenders.

17. Kentucky. Plain-bellied Watersnake (Nerodia erythrogaster)

Nerodia erythrogaster

Common elsewhere, a threatened subspecies of Plain-bellied Watersnake, the Copper-bellied (N. e. neglecta), is found in isolated spots in the western part of the Bluegrass State, as well as in adjacent areas in Illinois and Indiana. It is the most terrestrial of the ten species of watersnakes in the US, in part because it prefers frogs and toads to the other, more aquatic prey of its congeners. Uniquely among watersnakes, this species may possess resistance to toad toxin. It has been found that, atypically for a watersnake, these snakes tend to choose hibernacula close to wetlands, such as crayfish burrows and muskrat lodges. During spring floods, they are often submerged, but somehow are not drowned or swept away. The presence of these amazing snakes helps justify wetland buffers and adjacent upland conservation as part of the innovative Copperbelly Watersnake Conservation Agreement made between coal mining companies and state environmental groups within the range of the snake, particularly in Kentucky.

18. Louisiana. Louisiana Pinesnake (Pituophis ruthveni)

Pituophis ruthveni

Notable for its large eggs and small clutch sizes, the Louisiana Pinesnake is indigenous to west-central Louisiana and eastern Texas, where it relies strongly on Baird's pocket gophers for burrows and food. The Louisiana Pinesnake is rarely seen in the wild and is considered to be one of the rarest snakes in North America. The demise of the species is due to its low fecundity coupled with the extensive loss and degradation of suitable habitat, the longleaf pine savannas of the Gulf coastal plain. Now nearly a lost cause, some positive PR could help ensure that the last fragments of longleaf pine are preserved in perpetuity, for Louisiana Pinesnakes as well as their other unique flora and fauna.

19. Maine. Smooth Greensnake (Opheodrys vernalis)

Opheodrys vernalis

Although the Pine Tree State has a reputation for being snowy and cold, it is actually replete with snakes, at least in summer along its rocky coast. In the crumbling foundations of homes in old fields live many Smooth Greensnakes, a species that is a good example of the ecosystem services that snakes provide because it eats so many insects and spiders. Only a few species of snakes eat insects, this one chief among them. Less arboreal than their rough-scaled congeners, Smooth Greensnakes are quite beautiful and most people seem to know that they are harmless. A ranger in Acadia National Park found them "common" from sea level to above 1000 feet in 1938. Smooth Greensnakes are even found on Isle au Haut, a remote island in the outer reaches of Maine's Penobscot Bay first settled by Europeans in 1792, so it is likely that this is one of the first North American snakes ever seen by Europeans. Isle au Haut is home to Microneta bowditchiae, an endemic spider, which is undoubtedly eaten by the local O. vernalis.

20. Maryland. Queensnake (Regina septemvittata)

Regina septemvittata

Named for Queen Henrietta Maria of France and crisscrossed by myriad freshwater habitats, including the cool crayfish-filled streams preferred by Queensnakes, Maryland is a beautiful but densely-populated state. From the Central Appalachians in the west to the tidal marshes and barrier islands of the Delmarva peninsula and Chesapeake Bay, Maryland seems to represent the entire gamut of eastern North American habitats. Queensnakes inhabit most of these, although they are absent from the lowest and highest elevations and from brackish water. Research has shown that Queensnakes are exquisitely sensitive to the crayfish molting hormone ecdysone. Because numerous stream contaminants are known to imitate ecdysone and disrupt the arthropod molting cycle, this finding may have important implications for the conservation of Queensnakes and their conegners.

21. Massachusetts. Red-bellied Snake (Storeria occipitomaculata)

Storeria occipitomaculata

These little slug-eaters are named for a Massachusetts herpetologist, David Humphreys Storer, who wrote Report on the fishes, reptiles and birds of Massachusetts in 1839. Being one of the original 13 colonies, Massachusetts should show a little appreciation for its herpetological history. In my experience, these snakes are more common in New England's forests than they are throughout the rest of their range, particularly in Maine, which was once a part of Massachusetts. The Bay State gets kudos for actually selecting a group of snakes as their state reptile: the genus Thamnophis, the gartersnakes, is not an inapt choice, although it is a little vague.

22. Michigan: Eastern Massasauga (Sistrurus catenatus)

Sistrurus catenatus

The Eastern Massasauga Rattlesnake has declined dramatically over the last century and is critically endangered in nearly every state where it occurs. Due to human persecution and the nearly complete conversion of native wet prairie habitats to agriculture, this formerly widespread and abundant species is seemingly relegated to just a few viable populations within its historic range in Wisconsin and Illinois. Michigan is its stronghold, and state snakehood might help keep it that way.

23. Minnesota. Plains Gartersnake (Thamnophis radix)

Thamnophis radix

These beautiful gartersnakes are common in the plains of southern Minnesota. In northwestern part of the Land of 10,000 Lakes, the prairie-forest ecotone is characterized by the westward extension of the forests, which terminate roughly along the Big Stone Moraine, from which scattered areas of relict prairie form "fingers" that extend eastward into the woodland. Both Plains and Common (T. sirtalis) Gartersnakes co-occur there, but are not usually found together in high densities, perhaps because the diets of both species consist essentially of the same items: earthworms, minnows, and frogs. Unfortunately, Minnesota's many lakes (the combined shorelines of which exceed those of California, Florida, and Hawaii combined) are too cold, young, and oligotrophic to support any endemic snakes.

24. Mississippi. Mudsnake (Farancia abacura)

Farancia abacura

The Mississippi Mudsnakes has a good ring to it. Found in swamps, which abound in the western part of Mississippi, these gorgeous snakes eat mostly giant aquatic salamanders such as sirens and amphiumas. Seldom seen, their name belies their fabulous coloration, which more than compensates for their clandestine nature. Any state in their range would be proud to honor the Mudsnake with a state snakehood. Mudsnakes prefer still, acidic waters with aquatic vegetation and bottom debris. They also occur along small, acidic streams with swampy edges. Runner up: The Mississippi Green Watersnake (Nerodia cyclopion) shares its name with the state.

25. Missouri. Lined Snake (Tropidoclonion lineatum)

Tropidoclonion lineatum

Little is known about the tiny Lined Snake, a small relative of watersnakes and gartersnakes whose scientific name is longer than it is. Little is left of their original habitat, wet prairies, now mostly converted to agriculture. Lined Snakes, often incongruously misheard as "lion snakes," are secretive and semifossorial. The Forest Park neighborhood of St. Louis, south of the zoo and near the wonderful Turtle Park, was built on a hill during the 1920s and harbors a dense population due to the relatively light disturbance to the area's soil, so many urban residents of this city and others in the Show-Me State could show themselves their state snake by turning over paving stones or garden gnomes in their yard or garden.

Stay tuned, faithful reader, for Part II!

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1 Life is Short but Snakes are Long does not officially endorse the views of interviewed persons.^↩

ACKNOWLEDGMENTS

Thanks to photographers Dave Irving, JD Willson, Todd Pierson, James Turek, Lost Springs Ranch, Jason Folt, David Jahn, Jonathan Mays, Atrox, Eric Vanderduys, and Nathanael Herrera.

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