Anoualerpeton
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''Anoualerpeton'' 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
lissamphibia The Lissamphibia (from Greek λισσός (lissós, "smooth") + ἀμφίβια (amphíbia), meaning "smooth amphibians") is a group of tetrapods that includes all modern amphibians. Lissamphibians consist of three living groups: the Salientia ( ...
n in the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Albanerpetontidae The Albanerpetontidae (also spelled Albanerpetidae and Albanerpetonidae) 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 ...
. It is the oldest and most primitive albanerpetontid known. Fossils have been found of two different
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
, ''Anoualerpeton priscus'' from the
Middle Jurassic The Middle Jurassic is the second Epoch (geology), epoch of the Jurassic Period (geology), Period. It lasted from about 174.1 to 161.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relativel ...
(
Bathonian In the geologic timescale the Bathonian is an age (geology), age and stage (stratigraphy), stage of the Middle Jurassic. It lasted from approximately 168.2 ±1.2 annum, Ma to around 165.3 ±1.1 Ma (million years ago). The Bathonian Age succeeds ...
) aged Forest Marble and Kilmaluag formations of England and Scotland, and ''Anoualerpeton unicus'' from
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
-
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
(
Tithonian In the geological timescale, the Tithonian is the latest age (geology), age of the Late Jurassic Epoch and the uppermost stage (stratigraphy), stage of the Upper Jurassic Series. It spans the time between 149.2 ±0.7 annum, Ma and 143.1 ±0.6 (mi ...
-
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 143.1 ±0.6 Ma and 137.05 ± 0.2 (million years ago) ...
) Ksar Metlili Formation of
Morocco Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines on the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to Algeria–Morocc ...
. ''A. unicus'' is the only named albanerpetontid from
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
.


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), but is characterized by a mosaic of features that occur either in ''
Albanerpeton ''Albanerpeton'' is an extinct genus of salamander-like Albanerpetontidae, albanerpetontid amphibian found in North America, Europe and Asia first appearing in Cretaceous-aged strata. There are eight described members of the genus, and one undiag ...
'' 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''. 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), 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 of the
Bathonian In the geologic timescale the Bathonian is an age (geology), age and stage (stratigraphy), stage of the Middle Jurassic. It lasted from approximately 168.2 ±1.2 annum, Ma to around 165.3 ±1.1 Ma (million years ago). The Bathonian Age succeeds ...
of Central England. The Kirtlington Mammal Bed is the most productive microvertebrate deposit of the Kirtlington Quarry, a limestone quarry west of Kirtlington, 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 of synapsid tetrapods that includes the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts during the Late Triassic. It is defined as ...
. 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 of the
Isle of Skye The Isle of Skye, or simply 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 by the Cuillin, the rocky slopes of which provide some of ...
, 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 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 143.1 ±0.6 Ma and 137.05 ± 0.2 (million years ago) ...
, but have also been considered
Tithonian In the geological timescale, the Tithonian is the latest age (geology), age of the Late Jurassic Epoch and the uppermost stage (stratigraphy), stage of the Upper Jurassic Series. It spans the time between 149.2 ±0.7 annum, Ma and 143.1 ±0.6 (mi ...
. 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