Atretochoana
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''Atretochoana eiselti'' is a species of
caecilian Caecilians (; ) are a group of limbless, vermiform or serpentine amphibians. They mostly live hidden in the ground and in stream substrates, making them the least familiar order of amphibians. Caecilians are mostly distributed in the tropics of ...
originally known only from two preserved specimens discovered by Sir Graham Hales in the Brazilian rainforest, while on an expedition with Sir Brian Doll in the late 1800s, but rediscovered in 2011 by engineers working on a hydroelectric dam project in Brazil. Until 1998, it was known only from the
type specimen In biology, a type is a particular wiktionary:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to a ...
in the
Naturhistorisches Museum The Natural History Museum Vienna (german: Naturhistorisches Museum Wien) is a large natural history museum located in Vienna, Austria. It is one of the most important natural history museums worldwide. The NHM Vienna is one of the largest museum ...
, Vienna. Originally placed in the genus ''
Typhlonectes ''Typhlonectes'' (from el, τῠφλός , 'blind' and el, νηκτῆς , 'swimmer') is a genus of caecilians in the family Typhlonectidae. These fully aquatic amphibians are found in the Amazon Basin and Northern South America, and typically ...
'' in 1968, it was reclassified into its own
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
genus, ''Atretochoana'', in 1996. It was also found to be more closely related to the genus ''
Potamotyphlus ''Potamotyphlus kaupii'' (also known as Kaup's caecilian) is a species of amphibian in the family Typhlonectidae. It is monotypic within the genus ''Potamotyphlus''. It is found widely in the Amazon Basin and The Guianas in South America.Oliveira ...
'' than ''Typholonectes''. The species is the largest of the few known lungless
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
s, and the only known lungless caecilian.


Description

''A. eiselti'' is the largest
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
to lack
lung The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side of t ...
s, double the size of the next largest. Caecilians such as ''Atretochoana'' are limbless amphibians with snake-like bodies, marked with rings like those of
earthworm An earthworm is a terrestrial invertebrate that belongs to the phylum Annelida. They exhibit a tube-within-a-tube body plan; they are externally segmented with corresponding internal segmentation; and they usually have setae on all segments. Th ...
s. It has significant morphological differences from other caecilians, even the genera most closely related to it, even though those genera are aquatic. The skull is very different from those of other caecilians, giving the animal a broad, flat head. Its nostrils are sealed, and it has an enlarged mouth with a mobile cheek. Its body has a fleshy dorsal fin. Most caecilians have a well-developed right lung and a
vestigial Vestigiality is the retention, during the process of evolution, of genetically determined structures or attributes that have lost some or all of the ancestral function in a given species. Assessment of the vestigiality must generally rely on co ...
left lung. Some, such as ''Atretochoana''s relatives, have two well-developed lungs. ''Atretochoana'', however, entirely lacks lungs, and has a number of other features associated with lunglessness, including sealed
choanae The choanae (singular choana), posterior nasal apertures or internal nostrils are two openings found at the back of the nasal passage between the nasal cavity and the throat in tetrapods, including humans and other mammals (as well as crocodilia ...
, and an absence of
pulmonary arteries A pulmonary artery is an artery in the pulmonary circulation that carries deoxygenated blood from the right side of the heart to the lungs. The largest pulmonary artery is the ''main pulmonary artery'' or ''pulmonary trunk'' from the heart, and t ...
. Its skin is filled with capillaries that penetrate the
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water rele ...
, allowing gas exchange. Its skull shows evidence of muscles not found in any other organism. The Vienna specimen of ''Atretochoana'' is a large caecilian at a length of , while the Brasília specimen is larger still at . By comparison, caecilians in general range in length from . Although it is not a
snake Snakes are elongated, Limbless vertebrate, limbless, carnivore, carnivorous reptiles of the suborder Serpentes . Like all other Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales. Ma ...
, it has been called various
common names In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often contrast ...
in the media such as "penis snake", "man-aconda", and "floppy snake", owing to its visual similarity to the
human penis The human penis is an external male intromittent organ that additionally serves as the urinary duct. The main parts are the root (radix); the body (corpus); and the epithelium of the penis including the shaft skin and the foreskin (prepuce) co ...
.


History

The specimen in the Vienna museum was known only to have originated from somewhere in South America, at least before 1945, most likely in the 19th century. Its lack of lungs was not known at this time, and the specimen was assigned to the species ''
Typhlonectes compressicauda ''Typhlonectes compressicauda'', the Cayenne caecilian, is a species of amphibian in the family Typhlonectidae that lives in water. It is found in Amazonian Brazil, Peru, and Colombia as well as in Guyana and French Guiana, and likely Suriname, ...
''. The Vienna specimen was the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
for this species when it was first described by
Edward Harrison Taylor Edward Harrison Taylor (April 23, 1889 – June 16, 1978) was an American herpetologist from Missouri. Family Taylor was born in Maysville, Missouri, to George and Loretta Taylor. He had an older brother, Eugene. Education Taylor studied at th ...
in his 1968 monograph ''Caecilians of the World''. He named it ''Typhlonectes eiselti'', in honour of Viennese herpetologist
Josef Eiselt Josef may refer to *Josef (given name) *Josef (surname) * ''Josef'' (film), a 2011 Croatian war film *Musik Josef Musik Josef is a Japanese manufacturer of musical instruments. It was founded by Yukio Nakamura, and is the only company in Japan spe ...
. Taylor considered it to be similar to the aquatic caecilians of the genus ''
Typhlonectes ''Typhlonectes'' (from el, τῠφλός , 'blind' and el, νηκτῆς , 'swimmer') is a genus of caecilians in the family Typhlonectidae. These fully aquatic amphibians are found in the Amazon Basin and Northern South America, and typically ...
'' and ''
Potamotyphlus ''Potamotyphlus kaupii'' (also known as Kaup's caecilian) is a species of amphibian in the family Typhlonectidae. It is monotypic within the genus ''Potamotyphlus''. It is found widely in the Amazon Basin and The Guianas in South America.Oliveira ...
'', and placed it in the former genus, taking much note only of its large size and high number of
splenial The splenial is a small bone in the lower jaw of reptiles, amphibians and birds, usually located on the lingual side (closest to the tongue) between the angular and surangular The suprangular or surangular is a jaw bone found in most land verteb ...
teeth. Taylor did not inform the curators of the Naturhistorisches Museum that he designated the specimen a holotype, so it was not mentioned in the museum's catalogue of type specimens, and was placed beneath glass in a public display. There, it was noticed by the visiting English herpetologist Mark Wilkinson, who then borrowed the specimen to examine it with his American colleague Ronald A. Nussbaum. Examination of the specimen showed it to have a number of unusual features, including the large number of splenial teeth observed by Taylor, but most unusually, closed
choana The choanae (singular choana), posterior nasal apertures or internal nostrils are two openings found at the back of the nasal passage between the nasal cavity and the throat in tetrapods, including humans and other mammals (as well as crocodilia ...
e, which showed it could not fill any lungs it might have. Because of these and other distinctive features, Nussbaum and Wilkinson gave this species its own genus when they reported on the results of their research in a 1995 issue of the ''
Proceedings of the Royal Society B ''Proceedings of the Royal Society'' is the main research journal of the Royal Society. The journal began in 1831 and was split into two series in 1905: * Series A: for papers in physical sciences and mathematics. * Series B: for papers in life s ...
''. The name they gave this genus was ''Atretochoana'', from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
word ''atretos'', meaning "imperforate", and the Greek word ''choana'', referring to a funnel or tube. Nussbaum and Wilkinson published further studies in 1997 describing in detail the caecilian's anatomy and morphology. In 1998, they discovered the second specimen in the Universidade de Brasilia, although the origin of this specimen was also unknown. In 1999, they determined ''Atretochoana'' was a sister taxon of ''
Potamotyphlus ''Potamotyphlus kaupii'' (also known as Kaup's caecilian) is a species of amphibian in the family Typhlonectidae. It is monotypic within the genus ''Potamotyphlus''. It is found widely in the Amazon Basin and The Guianas in South America.Oliveira ...
'', and in 2011, grouped it in the family
Typhlonectidae Typhlonectidae, also known as aquatic caecilians or rubber eels, are a family of caecilians found east of the Andes in South America. They are viviparous animals, giving birth to young that possess external gills. Of the five genera in the fami ...
. Both of these specimens are mature females.


Biology

Most caecilians are burrowers, but some, including ''Atretochoana''s relatives, are largely aquatic. ''Atretochoana'' is thought to be aquatic since its relatives and
lungless salamander Plethodontidae, or lungless salamanders, are a family of salamanders. Most species are native to the Western Hemisphere, from British Columbia to Brazil, although a few species are found in Sardinia, Europe south of the Alps, and South Korea. In ...
s, some of the few other lungless tetrapods, are aquatic as well. It was postulated to inhabit fast-flowing water. Due to the lack of information, it is classified as "Data Deficient" by the
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
. It is thought to be uncommon, with a limited distribution. It is likely a predator or scavenger, and is thought to be
viviparous Among animals, viviparity is development of the embryo inside the body of the parent. This is opposed to oviparity which is a reproductive mode in which females lay developing eggs that complete their development and hatch externally from the m ...
. In June 2011, an amphibian was photographed near Praia de Marahú on the island of Mosqueiro (near
Belém Belém (; Portuguese for Bethlehem; initially called Nossa Senhora de Belém do Grão-Pará, in English Our Lady of Bethlehem of Great Pará) often called Belém of Pará, is a Brazilian city, capital and largest city of the state of Pará in t ...
, Brazil) that appeared to be ''A. eiselti'', but was not positively identified. In 2011, six individual organisms were found in the
Madeira River The Madeira River ( pt, Rio Madeira, link=no ) is a major waterway in South America. It is estimated to be in length, while the Madeira-Mamoré is estimated near or in length depending on the measuring party and their methods. The Madeira is ...
. Neither have cold, fast-flowing water, as was originally thought, as there is less oxygen in warmer water. This makes its lack of lungs even more unusual, and the question of how it breathes has not yet been resolved.


References


External links


1998 paper reporting discovery of second specimen

2011 paper reporting the discovery of live specimens
{{Taxonbar, from1=Q1916885, from2=Q15727328 Typhlonectidae Monotypic amphibian genera Amphibians described in 1968 Taxonomy articles created by Polbot