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''Andersonerpeton'' is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
aïstopod Aistopoda (Greek for " avingnot-visible feet") is an order of highly specialised snake-like stegocephalians known from the Carboniferous and Early Permian of Europe and North America, ranging from tiny forms only , to nearly in length. They fi ...
from the
Bashkirian The Bashkirian is in the International Commission on Stratigraphy geologic timescale the lowest stage (stratigraphy), stage or oldest age (geology), age of the Pennsylvanian (geology), Pennsylvanian. The Bashkirian age lasted from to Mega annu ...
(early Pennsylvanian) of
Nova Scotia, Canada Nova Scotia is a Provinces and territories of Canada, province of Canada, located on its east coast. It is one of the three Maritime Canada, Maritime provinces and Population of Canada by province and territory, most populous province in Atlan ...
. It is known from a single jaw, which shares an unusual combination of features from both other aistopods and from
stem-tetrapod The Stem Tetrapoda are a cladistically defined group, consisting of all animals more closely related to extant four-legged vertebrates than to their closest extant relatives (the lungfish), but excluding the crown group Tetrapoda. They are thus ...
tetrapodomorph Tetrapodomorpha (also known as Choanata) is a clade of vertebrates consisting of tetrapods (four-limbed vertebrates) and their closest sarcopterygian relatives that are more closely related to living tetrapods than to living lungfish. Advanced for ...
fish. As a result, ''Andersonerpeton'' is significant for supporting a new classification scheme which states that aistopods evolved much earlier than previously expected. The genus contains a single species, ''A. longidentatum'', which was previously believed to have been a species of the
microsaur Microsauria is an extinct, possibly polyphyletic order of tetrapods from the late Carboniferous and early Permian periods. It is the most diverse and species-rich group of lepospondyls. Recently, Microsauria has been considered paraphyletic, as s ...
''
Hylerpeton ''Hylerpeton'' is an extinct genus of microsaurian tetrapods belonging to the family Gymnarthridae from the late Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic ...
.''


History

The type species, ''A. longidentatum'', was initially described by
John William Dawson Sir John William Dawson (1820–1899) was a Canadian geologist and university administrator. Life and work John William Dawson was born on 13 October 1820 in Pictou, Nova Scotia, where he attended and graduated from Pictou Academy. Of Sco ...
on the basis of RM 2.1129, a left mandible. This fossil hailed from the
Joggins fossil cliffs Joggins is a rural community located in western Cumberland County, Nova Scotia, Canada. On July 7, 2008 a 15-km length of the coast constituting the Joggins Fossil Cliffs was officially inscribed on the World Heritage List.Nova Scotia Nova Scotia is a Provinces and territories of Canada, province of Canada, located on its east coast. It is one of the three Maritime Canada, Maritime provinces and Population of Canada by province and territory, most populous province in Atlan ...
famous for fossil deposits dated to the
Bashkirian The Bashkirian is in the International Commission on Stratigraphy geologic timescale the lowest stage (stratigraphy), stage or oldest age (geology), age of the Pennsylvanian (geology), Pennsylvanian. The Bashkirian age lasted from to Mega annu ...
, the first stage of the Pennsylvanian subperiod of the
Carboniferous period The Carboniferous ( ) is a geologic period and system of the Paleozoic era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the Permian Period, Ma. It is the fifth and penultimate perio ...
. ''A. longidentatum'' was originally named ''Hylerpeton longidentatum'', as Dawson considered it a new species of the microsaur ''
Hylerpeton ''Hylerpeton'' is an extinct genus of microsaurian tetrapods belonging to the family Gymnarthridae from the late Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic ...
,'' distinct from ''Hylerpeton dawsoni'' (the type species of ''
Hylerpeton ''Hylerpeton'' is an extinct genus of microsaurian tetrapods belonging to the family Gymnarthridae from the late Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic ...
''). Steen (1934) and Carroll (1966) questioned the microsaurian placement of ''H. longidentatum'', but withheld judgment on its taxonomic status, although Carroll (1966) noted similarities to non-tetrapod sarcopterygians. Pardo and Mann (2018) demonstrated that ''H. longidentatum'' belongs to the limbless tetrapod clade
Aistopoda Aistopoda (Greek for " avingnot-visible feet") is an order of highly specialised snake-like stegocephalians known from the Carboniferous and Early Permian of Europe and North America, ranging from tiny forms only , to nearly in length. They fi ...
and renamed it ''Andersonerpeton'' in honor of Jason S. Anderson, a contributor to enhanced understanding of aïstopod morphology and phylogeny.


Description

The preserved portion of the jaw was smooth, shallow and relatively wide, curving upwards at the parasymphyseal (chin) region. The main bone of the jaw, the dentary, possessed a single row of long, curved teeth, known as marginal teeth. These teeth were weakly attached to the underlying bone, which allows them to fold backwards. This ability is also seen in certain
snakes Snakes are elongated Limbless vertebrate, limbless reptiles of the suborder Serpentes (). Cladistically Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales much like other members of ...
as well as oestocephalid aistopods, betraying ''Andersonerpeton'''s aistopod affinities. Unusually, the teeth also have a maze-like internal structure, formally known as labyrinthine infolding. This type of structure is known in large "
labyrinthodont "Labyrinthodontia" (Greek, 'maze-toothed') is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras (about 390 to 150 million years ago). Traditionally conside ...
" tetrapods and their tetrapodomorph ancestors, but unknown in any other aistopod. Lingually from the dentary tooth row (i.e. towards the tongue), the upper surface of the mandible has five plate-like bones: a parasymphyseal plate at the chin, an adsymphyseal bone directly behind it, and a string of three coronoid bones directly behind the adsymphyseal. All five of these bones are covered with tiny tooth-like bumps known as denticles. Larger fangs, similar to the marginal teeth, were also present on some of these bones; the first and second coronoids each have two of these fangs at their front outer corner and the parasympheseal has a single large fang in the middle. Additional denticles may have been present on the dentary, which has a small number of small bumps on the outer surface of the parasymphyseal region. If these were true denticles, then this is a surprisingly "primitive" feature present in ''Andersonerpeton'', as this feature is typically considered to have been lost very early in the evolution of tetrapods (specifically, after the evolution of ''
Elginerpeton ''Elginerpeton'' is a genus of stegocephalian ( stem-tetrapod), the fossils of which were recovered from Scat Craig, Morayshire in the UK, from rocks dating to the late Devonian Period (Late Frasnian stage, 375 million years ago). The type spe ...
''). The prominent parasympheseal and coronoid fangs are also more consistent with stem-tetrapods rather than other aistopods. ''Andersonerpeton'' does have one specific feature shared by both certain aistopods and certain stem-tetrapods: a longitudinal plate of bone directly below the coronoids, known as a "meckelian ossification".


Classification

The uniquely foldable teeth indicate that ''Andersonerpeton'' is an aistopod, as these teeth are also seen in '' Coloraderpeton'' and ''
Oestocephalus ''Oestocephalus'' is an extinct genus of Aïstopoda, aïstopod tetrapodomorphs that lived during the Carboniferous period. Fossils have been found in the Czech Republic, and in Ohio and Illinois in the United States. It is the type genus of the f ...
'', but no other tetrapods until the appearance of snakes in the Cretaceous period. In addition, the mandible was smooth and wide, similar to that of most aistopods. However, in many other features the jaw of ''Andersonerpeton'' closely resembles early tetrapod relatives such as ''Elginerpeton''. These include the arrangement of teeth and denticles on the parasympheseal and coronoid bones, as well as labyrinthine marginal teeth. The possible presence of denticles on the dentary next to the tooth row is particularly significant, as these are present in many
tetrapodomorph Tetrapodomorpha (also known as Choanata) is a clade of vertebrates consisting of tetrapods (four-limbed vertebrates) and their closest sarcopterygian relatives that are more closely related to living tetrapods than to living lungfish. Advanced for ...
fish but completely lacking in all but the earliest four-limbed vertebrates, with ''Elginerpeton'' believed to have been the last known animal to have possessed such a feature. Overall, ''Andersonerpeton'' seems to bridge a gap between stem-tetrapods and aistopods. This classification scheme contrasts with traditional interpretations of aistopods forming a group called
Lepospondyli Lepospondyli is a diverse clade of early tetrapods. With the exception of one late-surviving lepospondyl from the Late Permian of Morocco ('' Diplocaulus minimus''), lepospondyls lived from the Visean stage of the Early Carboniferous to the Ea ...
with animals like '' Diplocaulus'' and microsaurs, which are almost always considered true
crown A crown is a traditional form of head adornment, or hat, worn by monarchs as a symbol of their power and dignity. A crown is often, by extension, a symbol of the monarch's government or items endorsed by it. The word itself is used, parti ...
-
tetrapods A tetrapod (; from Ancient Greek τετρα- ''(tetra-)'' 'four' and πούς ''(poús)'' 'foot') is any four- limbed vertebrate animal of the clade Tetrapoda (). Tetrapods include all extant and extinct amphibians and amniotes, with the lat ...
. However, the idea that aistopods branched off from the tetrapodomorph family tree much earlier than other lepospondyls has been supported by some analyses, such as a 2017 study on the
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, brain-pan, or brainbox, is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calv ...
of ''
Lethiscus ''Lethiscus'' is the earliest known representative of the Aistopoda, a group of very specialised snake-like tetrapodomorphs known from the early Carboniferous ( Mississippian). ''Lethiscus'' is known from only a single specimen from the Holkeri ...
'', which was considered to have been the most "primitive" aistopod prior to the naming of ''Andersonerpeton''.


References

{{Taxonbar, from=Q60789065 Aistopoda Bashkirian life Pennsylvanian sarcopterygians of North America Paleozoic life of Nova Scotia Fossil taxa described in 2018