Anoualerpeton
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''Anoualerpeton'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
lissamphibia The Lissamphibia is a group of tetrapods that includes all modern amphibians. Lissamphibians consist of three living groups: the Salientia (frogs, toads, and their extinct relatives), the Caudata (salamanders, newts, and their extinct relatives), ...
n in the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Albanerpetontidae The Albanerpetontidae are an extinct family of small amphibians, native to the Northern Hemisphere during the Mesozoic and Cenozoic. The only members of the order Allocaudata, they are thought to be allied with living amphibians belonging to L ...
. It is the oldest and most primitive albanerpetontid known. Fossils have been found of two different
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
, ''Anoualerpeton priscus'' from the
Middle Jurassic The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations co ...
(
Bathonian In the geologic timescale the Bathonian is an age and stage of the Middle Jurassic. It lasted from approximately 168.3 Ma to around 166.1 Ma (million years ago). The Bathonian Age succeeds the Bajocian Age and precedes the Callovian Age. Strat ...
) aged
Forest Marble The Forest Marble is a geological formation in England. Part of the Great Oolite Group, it dates to the late Bathonian stage of the Middle Jurassic.Weishampel, David B; et al. (2004). "Dinosaur distribution (Middle Jurassic, Europe)." In: Weisha ...
and Kilmaluag formations of England and Scotland, and ''Anoualerpeton unicus'' from
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
-
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
(
Tithonian In the geological timescale, the Tithonian is the latest age of the Late Jurassic Epoch and the uppermost stage of the Upper Jurassic Series. It spans the time between 152.1 ± 4 Ma and 145.0 ± 4 Ma (million years ago). It is preceded by the K ...
-
Berriasian In the geological timescale, the Berriasian is an age/stage of the Early/Lower Cretaceous. It is the oldest subdivision in the entire Cretaceous. It has been taken to span the time between 145.0 ± 4.0 Ma and 139.8 ± 3.0 Ma (million years ago) ...
)
Ksar Metlili Formation The Ksar Metlili Formation is a geological formation in eastern High Atlas of Morocco, it is late Tithonian to Berriasian In the geological timescale, the Berriasian is an age/stage of the Early/Lower Cretaceous. It is the oldest subdivision ...
of
Morocco Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
. ''A. unicus'' is the only named albanerpetontid from
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...
.


Etymology

The name "''Anoualerpeton''" is composed of Anoual, the name of a city in the eastern High Atlas (Morocco), in whose relative proximity the type material of the type species ''Anoualerpeton unicus'' was found, and the ancient Greek word ἑρπετόν herpeton, which means 'creeping animal" and is often used in the generic names of fossil amphibians. The epithets "priscus" and "unicus" mean 'old' and 'unique', which refers to the fact that ''Anoualerpeton'' ''priscus'' is geologically the oldest albanerpetontid and ''Anoualerpeton unicus'' the only albanerpetontid that occurs on the former large southern continent of Gondwana.Gardner, J.D., Evans, S.E., and Sigogneau−Russell, D. (2003). New albanerpetontid amphibians from the Early Cretaceous of Morocco and Middle Jurassic of England. ''Acta Palaeontologica Polonica'' 48(2): 301–319.


Morphology

''Anoualerpeton'' does not have any unique features (
autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...
), but is characterized by a mosaic of features that occur either in ''
Albanerpeton ''Albanerpeton'' is an extinct genus of salamander-like lissamphibian found in North America and Europe, first appearing in Cretaceous-aged strata. There are eight described members of the genus, and one undiagnosed species from the Paskapoo Form ...
'' but not in ''
Celtedens ''Celtedens'' is an extinct genus of albanerpetontid amphibian from the Early Cretaceous of England, Spain, Sweden and Italy, and the Late Jurassic of Portugal. Taxonomy * †''Celtedens ibericus'' McGowan and Evans 1995 La Huérguina Format ...
'' or in ''Celtedens'' but not in ''Albanerpeton''. Thus, the occlusal margin of maxillary and dental teeth is convex in labial and lingual view, respectively, and the teeth in the middle third of the row are the longest, both of which are otherwise only known from the species ''
Albanerpeton nexuosus ''Albanerpeton'' is an extinct genus of salamander-like lissamphibian found in North America and Europe, first appearing in Cretaceous-aged strata. There are eight described members of the genus, and one undiagnosed species from the Paskapoo For ...
''. The processus internasalis of the azygic frontal, i.e. the end of the bone resulting from the fusion of the originally paired frontal structures, which is flanked by the nasal bone and points towards the tip of the snout, tapers to a point. The lateral edges of this process are designed as a kind of groove into which the edges of the nasalia engage (
tongue-and-groove Tongue and groove is a method of fitting similar objects together, edge to edge, used mainly with wood, in flooring, parquetry, panelling, and similar constructions. Tongue and groove joints allow two flat pieces to be joined strongly together t ...
), both of which occur in ''Albanerpeton'' but not in ''Celtedens''. In addition, the length-width ratio of the frontal is relatively large and the frontal is bell-shaped rather than triangular in dorsal and ventral view, features which ''Anoualerpeton'' shares only with ''Celtedens'' but not with ''Albanerpeton''. The two ''Anoualerpeton'' species differ primarily in the location of the suprapalatal pit. This is a depression in the inner side (lingual side) of that part of the premaxillary which reaches up to the roof of the skull (pars dorsalis). In ''Anoualerpeton priscus'', the suprapalatal fossa lies in the medial edge of the external nasal opening. In ''Anoualerpeton unicus'', however, it lies further mediad, outside the border of the external nasal opening.


Locality

As is common for Albanerpetontids, the remains of ''Anoualerpeton'' all come from so-called microvertebrate localities, i.e. isolated, relatively small bones obtained by slurrying and sieving samples of the sedimentary rock in which they were embedded. The older species ''Anoualerpeton priscus'' originates from the Kirtlington Mammal Bed ("Kirtlington mammal layer") in the lowest part of the
Forest Marble Formation The Forest Marble is a geological formation in England. Part of the Great Oolite Group, it dates to the late Bathonian stage of the Middle Jurassic.Weishampel, David B; et al. (2004). "Dinosaur distribution (Middle Jurassic, Europe)." In: Weisham ...
of the
Bathonian In the geologic timescale the Bathonian is an age and stage of the Middle Jurassic. It lasted from approximately 168.3 Ma to around 166.1 Ma (million years ago). The Bathonian Age succeeds the Bajocian Age and precedes the Callovian Age. Strat ...
of Central England. The Kirtlington Mammal Bed is the most productive microvertebrate deposit of the
Kirtlington Quarry Kirtlington Quarry is a geological Site of Special Scientific Interest west of Kirtlington in Oxfordshire. It is a Geological Conservation Review site, and it is part of the Kirtlington Quarry Local Nature Reserve. Kirtlington Quarry is one of ...
, a limestone quarry west of
Kirtlington Kirtlington is a village and civil parish in Oxfordshire about west of Bicester. The parish includes the hamlet of Northbrook. The 2011 Census recorded the parish's population as 988. The parish measures nearly north–south and about east ...
, about 15 kilometers north of Oxford. It consists of a weakly consolidated clayey marl, which was probably deposited in a swampy environment and contains, among other things, teeth of dinosaurs and early
mammaliaformes Mammaliaformes ("mammalian forms") is a clade that contains the crown group mammals and their closest Extinction, extinct relatives; the group adaptive radiation, radiated from earlier probainognathian cynodonts. It is defined as the clade origin ...
. The material from Kirtlington, classified as ''Anoualerpeton priscus'', was formerly assigned to the genera ''Albanerpeton'' or ''Celtedens''. In 2020 a connected pair of premaxilla similar to those of ''A. priscus'' were reported from the
Kilmaluag Formation The Kilmaluag Formation is a Middle Jurassic geologic formation in Scotland. It was formerly known as the Ostracod Limestone for the abundance of fossil freshwater ostracods within it. The Kilmaluag Formation is very fossiliferous, with ostracods ...
of the
Isle of Skye The Isle of Skye, or simply Skye (; gd, An t-Eilean Sgitheanach or ; sco, Isle o Skye), is the largest and northernmost of the major islands in the Inner Hebrides of Scotland. The island's peninsulas radiate from a mountainous hub dominated ...
, Scotland. The deposit is equivalent in age to the Kirtlington Mammal Bed, and is thought to have been deposited in a coastal lagoon. The younger species ''Anoualerpeton unicus'', which is also a type species of the genus, was identified in the microvertebrate fauna of the
Ksar Metlili Formation The Ksar Metlili Formation is a geological formation in eastern High Atlas of Morocco, it is late Tithonian to Berriasian In the geological timescale, the Berriasian is an age/stage of the Early/Lower Cretaceous. It is the oldest subdivision ...
in the eastern High Atlas. The locality is located in the south of the Talsint province of Morocco, about 100 kilometers east of the city of Anoual. The deposit is located in a lens of non-marine limestone embedded in marine sediments in the upper part of the so-called Couches Rouges ("red layers"). These marine sediments have been uncertainly dated by coccolithophorids to the
Berriasian In the geological timescale, the Berriasian is an age/stage of the Early/Lower Cretaceous. It is the oldest subdivision in the entire Cretaceous. It has been taken to span the time between 145.0 ± 4.0 Ma and 139.8 ± 3.0 Ma (million years ago) ...
, but have also been considered
Tithonian In the geological timescale, the Tithonian is the latest age of the Late Jurassic Epoch and the uppermost stage of the Upper Jurassic Series. It spans the time between 152.1 ± 4 Ma and 145.0 ± 4 Ma (million years ago). It is preceded by the K ...
. Also in Ksar Met Lili dinosaurs can be found and, due to the lower geological age, also representatives of the mammalian crown group (Mammalia).


Phylogeny

From Daza et al. 2020.


References

{{Taxonbar, from=Q4770421 Albanerpetontidae Prehistoric amphibian genera Late Jurassic amphibians Middle Jurassic amphibians Jurassic amphibians of Europe Cretaceous amphibians of Africa Fossil taxa described in 2003