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Saturday, August 30, 2014

Filesnakes, Wartsnakes, or Elephant Trunksnakes

Arafura Filesnake (Acrochordus arafurae)
In the swamps, marshes, streams, and estuaries of northern Australia and southeastern Asia live ancient snakes as thick as your arm, with tongues as thin as a thread, skin as rough as a file, and a disposition as gentle as a lamb. These snakes comprise the family Acrochordidae (from the Greek akrochordon, wart), and are known as filesnakes1, wartsnakes, or elephant trunksnakes. In Indonesian they are known as karung, which means 'sack'; in Thai, as ngū nguang-cĥāng, 'elephant-trunk snake'. There are three species, all in the genus Acrochordus: Javan (A. janavicus), Arafura (A. arafurae), and Little (A. granulatus). The largest, female Javans, grow up to 8 feet and over 20 pounds. Acrochordids are an old and highly distinct group of snakes, distantly related to colubroids, with which they share a common ancestor between 50 and 70, but possibly as long as 90 million years ago.

Close-up of A. arafurae scales
Filesnakes have strongly-keeled scales with the texture of sandpaper or a coarse file, after which they are named. They have very loose, baggy skin. I held one once and it felt like a human arm inside the sleeve of a very sturdy, very baggy rain poncho made out of chain mail. This loose skin is likely an adaptation that allows filesnakes to withstand the great force of the initial dash for freedom of their fish prey. Their sharp scales are used to help gain purchase on slimy fish skin during constriction. They have sensory, bristle-bearing tubercles on the skin between their scales, as well as sensory organs on the scales themselves, both of which presumably help them sense the underwater movements of nearby prey, analogous to the lateral line systems of fishes and some amphibians. Acrochordids sometimes ambush their prey, but more often they forage by searching slowly along the edges of mangroves, billabongs, and other water bodies at night, looking for sleeping fishes and crustaceans (although they don't tear them apart like some southeast Asian snakes). During the day, filesnakes hide in the shadows of overhanging trees, moving with them to remain concealed from predatory birds. They are nearly incapable of moving on land, and shed in the water using a knotting behavior similar to that of Pelamis platurus, the most completely aquatic sea snake.

Acrochordus arafurae regurgitates
an eel-tailed catfish (Tandanus tandanus)
whose spine has pierced its neck
Filesnakes occupy a unique phylogenetic position, not closely related to anything but somewhere in-between the colubroids ("advanced snakes") and the "henophidians" (boas, pythons, and uropeltoids). A few recent papers reanimate an old hypothesis that they might be closely related to dragonsnakes, but historically acrochordids have been considered the sister group to all colubroids, a group of >2,850 species (>80% of all living snakes) that includes dragonsnakes, asymmetrical slug-eaters, vipers, homalopsids, elapids, lamprophiids, and strict colubrids. Colubroidea and the three Acrochordus species together form the Caenophidia ("recent snakes"). Acrochordids share unspecialized head scales and undifferentiated ventral scales with boas and pythons, but they are united with colubroids in that they totally lack vestigial limbs and have spines on their hemipenes, a well-developed vomeronasal system, and several particular characteristics of skull morphology, including a coronoid bone. Other features of the skull and skeleton are unique to acrochordids, including the aforementioned skin sense organs, a passive joint between the frontal and parietal skull bones, the presence of certain holes in the vertebrae, the shape of the head of the ribs, and an ear region that most closely resembles the ears that other snakes have as embryos, but which forms in a different way. Acrochordids also have an unusual lung morphology, with a double row of holes leading from the trachea into individual small lunglets, and a more tangled intestinal tract than other snakes.

Acrochordus granulatus with algae growing on its back
Filesnakes have incredibly low metabolic rates, even for a snake, and cannot sustain rigorous physical activity for very long. In captivity, they "epitomize sluggishness in snakes", although radio telemetry has shown that in the wild they move around wetlands slowly but steadily, covering up to 450 feet per night. They can remain submerged for over an hour (record 2 h 20 min), and surface to take about 5 breaths, about one per minute. The first several of these breaths oxygenate the blood, and the last one fills the multi-chambered lung. In addition, Little Filesnakes have about twice as much blood as other snakes, and this voluminous blood is about twice as thick with red blood cells as even that of other diving snakes. Their hemoglobin has a very high affinity for oxygen, which results in their being able to store between three and fifteen times as much oxygen in their blood as a similarly-sized sea snake, and release it slowly over a long period of time. Many turtles also use this strategy. Also like turtles, filesnakes can both obtain oxygen from and release acidic carbon dioxide into the water through their skin, which helps prolong their dives.2 In fact, filesnakes are so well-adapted to sitting still that they are practically incapable of exercise, and get tired out quickly.

This slow theme carries over into filesnake life history. Male filesnakes mature around six years old, females around nine, and 8-10 years may elapse between consecutive births. Studies from northern Australia found that only a small proportion of females are reproductive in any given year, and that only the very largest females reproduce relatively frequently. Large Javan Filesnakes give birth to as many as 52 young at once, although the average is closer to 30. Arafura Filesnakes average about 16 (as few as 9 and as many as 25 have been reported), and Little Filesnakes about 6 (as few as 1 and as many as 12). Female filesnakes are courted by up to eight males at a time in shallow water. Their population dynamics are driven by rainfall in northern Australia. One captive filesnake gave birth to a single young after seven years of isolation, suggesting that filesnakes are either capable of parthenogenesis or of very prolonged sperm storage.

The Little Filesnake (Acrochordus granulatus) has
a banded pattern like a sea krait (Laticauda colubrina)
At first glance the three extant Acrochordus species seem quite similar, but in fact they exhibit striking differences in both anatomy and ecology. The Little Filesnake (Acrochordus granulatus) was described in 1799 and used to be classified in a separate, monotypic genus (Chersydrus). As its name suggests, it is the smallest acrochordid (~ 3 feet in total length) and the most widely distributed. It is found along the coast from northwestern India throughout southeast Asia and Indonesia, reaching east to the Solomon Islands. Its diverse habitats include freshwater lakes, rivers, mangroves, mudflats, reefs, and the open ocean, up to 6 miles offshore and over 60 feet deep. It is the most marine of the filesnakes, the most brightly patterned, and has a shorter, more laterally-compressed tail, more granular scales, more dorsally-oriented nostrils, and a salt excretion gland beneath its tongue.3

Acrochordus javanicus
The Javan Filesnake (Acrochordus javanicus) was the first to be described, in 1787, and is the largest and heaviest filesnake, sometimes reaching 8 feet and over 20 pounds. It is found in fresh and brackish water on the Malay Peninsula and on the islands of Sumatra, Java, and Borneo (and was introduced to Florida in the 1980s, although it does not appear to have established there). It is harvested for meat and for its skin, out of which is made fine leather; up to 2 million are exported from Indonesia annually. Unlike other filesnakes, the posterior-most teeth in its lower jaw have sharp edges. The Arafura Filesnake (Acrochordus arafurae) was thought to be the same species as the Javan until 1979. It grows as long but at the same size is only about half as heavy-bodied. It is found only in freshwater habitats in northern Australia and southern New Guinea. Surprisingly, A. arafurae is more closely related to A. granulatus than either is to A. javanicus, a relationship that is supported by genetics as well as morphology.

The long, thin tongue of Acrochordus javanicus
From Greene (1997)
Fossil Acrochordus have been found in Pakistan and Nepal, as well as within the extant range. These extinct filesnakes date from 5-20 million years ago during the Miocene, only a few million years after the Indian plate crashed into Asia to form the Himalayas. They grew larger than modern filesnakes, reaching at least 9 feet, and are the most well-represented snakes in the southern Asian fossil record, possibly because their habitat lends itself well to fossilization. The extinct species Acrochordus dehmi is represented by over 1000 fossils from over 100 different locations, and probably went extinct about 6 million years ago. Because it is so well-known, we can say with confidence that it is more closely related to A. javanicus than to the other two living species of AcrochordusMolecular clock methods suggest that the three modern species of Acrochordus and A. dehmi diverged from one another 16-20 million years ago, a timescale that usually justifies separation into family-level or higher categories. Despite their superficial similarities, the ecological and morphological differences among the three living Acrochordus species have been considered equivalent to differences among genera in other groups of snakes. Because no fossil acrochordids have been found in Australia, it is assumed that they evolved in Asia and spread to Australia in the last 5 million years. It is also likely that the ancestors of the Little Filesnake entered the ocean before sea snakes (~7 mya) and kraits (~13 mya) and just after marine homalopsids (~18 mya).

Acrochordus in contemporary aboriginal artwork by Chris Liddy (Moonggun),
showing the embryos inside the snake in the northern Australian style
One of the most interesting things about filesnakes is that Aboriginal Australians collect and eat them in some areas. Mostly this is done at the end of the Australian dry season, in November, when water levels are lowest and the snakes are easiest to find and capture. Although the snakes themselves don't generally put up much resistance, the old women who hunt them do so by wading into murky waters filled with crocodiles and feel under overhanging banks, weed beds, and logs, sometimes collecting over 30 snakes per person-hour. Often the snakes are killed immediately by biting their necks. The pregnant females are highly prized for their embryos, which are cooked on hot ashes, eaten like popcorn, and called 'cookies' by Aboriginal children.

1 Not to be confused with African Filesnakes (genus Mehelya), which are so-named not for their texture but for their cross-sectional shape, which resembles a triangular file.

2 Although the warm, shallow, slow-moving waters in which they live are fairly oxygen-poor and oxygen is difficult to extract out of salty water, so augmenting their ability to hold their breath using their massive blood oxygen reservoir is almost certainly of greater importance.

3 Little Filesnakes can excrete salt but gradually get dehydrated, so they must have a source of fresh water. They drink rain that falls on the ocean or migrate to areas where rivers flow into estuaries. This is because, like other marine reptiles, filesnakes "pee like a fish": they excrete nitrogen as ammonia, rather than as uric acid like other snakes or as urea like mammals. This is much more wasteful of water than the uric acid method, and it's not clear why they do this.


Thanks to Chris LiddyMatt Summerville, Darryl Houston, M. & P. Fogden, Jordan de Jong, Stephen Zozaya, Jason Isley, and Dick Bartlett for their photos, and to Rick Shine for information on tracking down Darryl Houston.


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Intertubercular papilla of Acrochordus granulatus
From Povel & Van Der Kooij 1996

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