Albanerpetontidae

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Albanerpetontidae
Temporal range: Middle JurassicPleistocene Bathonian–Gelasian
Celtedens ibericus.svg
Fossil of Celtedens ibericus, showing the remains of scales surrounding the body in grey
Yaksha drawing no photo.svg
Skull of Yaksha peretti
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Amphibia
Order: Allocaudata
Fox and Naylor, 1982
Family: Albanerpetontidae
Fox and Naylor, 1982
Genera
Synonyms
  • Albanerpetonidae
  • Albanerpetidae

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 be allied with living amphibians belonging to Lissamphibia. Despite a superficially salamander-like bodyform, their anatomy is strongly divergent from modern amphibians in numerous aspects. The fossil record of albanerpetontids spans over 160 million years from the Middle Jurassic to the beginning of the Pleistocene, about 2.13–2 million years ago.

Contents

History of research

Holotype fossil of Celtedens megacephalus from Italy Celtedens megacephalus.JPG
Holotype fossil of Celtedens megacephalus from Italy

The earliest specimen of an albanerpetontid to be discovered was that of Celtedens megacephalus from the Early Cretaceous (Albian) Pietraroja Plattenkalk of Italy, described by Oronzio Gabriele Costa in 1864, and originally placed in the genus Triton, a junior synonym of the salamander genus Triturus . [1] Jaw elements of albanerpetontids from the Cretaceous of North America were assigned to the salamander genus Prosiren by Richard Estes in 1969, erecting the family Prosirenidae to accommodate the genus. [2] Prosiren was originally described by Coleman J. Goin and Walter Auffenberg in 1958, based on vertebrae found in Cretaceous aged deposits in Texas. [3] Albanerpeton , the type genus of the family was first named by Estes and Robert Hoffstetter in 1976 for the species of A. inexpectatum described from a large number of jaws and frontal bones from a Miocene aged fissure fill deposit near Saint-Alban-de-Roche in France, and was initially classified as a salamander, and placed in the family Prosirenidae alongside Prosiren due to the morphological similarity with the jaw fragments attributed to Prosiren by Estes (1969). [4] Richard Fox and Bruce Naylor in 1982 realised that Albanerpeton was not a salamander, noting that the holotype vertebra of Prosiren was different to those of albanerpetontids, concluding that Albanerpeton was "well isolated from salamanders" and that it "seems no nearer phyletically to any other known amphibians, from Devonian to Recent" erecting the family Albanerpetontidae and the order Allocaudata to accommodate it. [5]

Description

Bones of the only articulated albanerpetontid skull, Yaksha peretti, vomer not shown Yaksha perettii skull diagram.svg
Bones of the only articulated albanerpetontid skull, Yaksha peretti, vomer not shown

Albanerpetontids were small (several cm to several tens of centimetres in length) and superficially lizard-like. The skin of albanerpetontids was embedded with bony, fish-like scales. The forelimbs only had four digits, while retaining five digits on the hindlimbs. The morphology of the complete three-dimensionally preserved skull of Yaksha peretti suggests that albanerpetontids had ballistic tongues akin to those of chameleons and plethodontid salamanders, as evidenced by the presence of an elongated rod shaped bone in the jaw cavity, dubbed the hyoid entoglossal process, which in life was embedded within the tongue. Analogous bones exists in chameleons and plethodontids, which allow rapid propulsion of the tongue. [6] A hyoid entoglossal process is also known from Celtedens megacephalus, suggesting that the presence of a ballistic tongue is characteristic for the group. [7] [6] Distinguishing apomorphic traits characteristic of albanerpetontids include a complex mortise and tenon–like joint connecting the dentary bones at the front of the jaw, teeth which are non-pedicellate and slightly tricuspid (bearing three cusps), the frontal bones of the skull display raised polygonal sculpturing, and three anterior cervical components form an 'atlasaxis' complex, similar to that of amniotes. [8]

Paleobiology

Life restoration of Albanerpeton inexpectatum Albanerpeton inexpectatum.jpg
Life restoration of Albanerpeton inexpectatum

The morphology of albanerpetontids suggests that they were sit-and-wait terrestrial predators and fed on invertebrates, similar to living plethodontids. The fact that the skull of the juvenile paratype of Yaksha was around 1/4 of the size of the adult suggests that albanerpetontids grew by direct development and did not have a metamorphic larval stage. [6] It has been suggested that albanerpetontids absorbed oxygen entirely through the skin via cutaneous respiration and lacked lungs like plethodontid salamanders, due to the length of the hyoid entoglossal process, which may have made normal breathing difficult. [6] This proposal is supported by the internal vascularisation and lack of Sharpey's fibres in the frontal bones. [9] Albanerpetontids are associated with both wet and dry environments, but it is unclear how tolerant they were of dry habitats, and they may have been confined to wet microhabitats in dry areas. [10] Some authors have suggested that they were likely fossorial, using their heads to burrow, but this has been questioned by other authors. [11]

Distribution

The distribution of albanerpetontids is largely confined to Eurasia and North America, with remains also known from Morocco in North Africa. [12] [13] The first albanerpetontids are known from the western Palearctic (Europe and North Africa) in the Middle Jurassic (Bathonian ~168–166 million years ago), with the oldest records of the group in North America and Asia dating to the Early Cretaceous. The last known remains of albanerpetontids in North America are from the Paskapoo Formation in Canada, dating to the Paleocene. All other Cenozoic members of the family, belonging to the genus Albanerpeton, are known from Europe and Anatolia, from the Oligocene onwards (there is no fossil record of albanerpetontids during the Eocene) until their final appearance in Northern Italy during the Early Pleistocene, around 2.13-2 million years ago. [13] [14] [15] [8] Another possible late record is known from northern Spain, dating to around 2.2-2.6 million years ago. [16]

Classification

Albanerpetontids were long thought to be salamanders because of their small size and generalized body plans. [17] However, these features are now thought to be ancestral for lissamphibians and not indicative of close relationships between the two groups. [18] Albanerpetontids share with living lissamphibians an atlanto-occipital joint with two cotyles, a four fingered forelimb (manus), ectochordal (spoon shaped with open centra) vertebrae with cylindrical centra, ribs that do not encircle the body, and a salamander-like quadratesquamosal articulation, but are distinguished from the three living groups of lissamphibians by their possession of keratinized claw sheaths and their retention of skull bones lost in other lissamphibians, including epipterygoids, supraoccipitals and large palatines, as well as the absence of pedicellate teeth or a wide parasphenoid cultriform process. [6] Albanerpetontids are now recognized as a distinct clade of lissamphibians separate from the three living orders of amphibians – Anura (frogs), Caudata (salamanders), and Gymnophiona (caecilians). Many studies show them as more closely related to frogs and salamanders than to caecilians, [19] but bootstrap and Bayesian analyses show that this result is not robust and that they could also be sister-group of the Lissamphibia, [20] or as most closely related to caecillians. [21] The presence of epipterygoids and a separate supraoccipital at least argues against a position within Batrachia. [8] A phylogenetic analysis in 2020 among lissamphibian relationships using multiple methods found no consensus for the position of Albanerpetontidae in relation to other lissamphibians, but they were always placed closer to lissamphibians than to other extinct groups of amphibians, such as lepospondyls and temnospondyls. [6]

Taxonomy

Phylogeny

From Daza et al 2020. [6]

Anoualerpeton

Celtedens

Uña taxon

Wesserpeton

Albanerpeton arthridion

Albanerpeton gracilis + Albanerpeton galaktion

Albanerpeton cifellii

Yaksha

Shirerpeton

Albanerpeton nexuosum

Albanerpeton pannonicum

Paskapoo sp

Albanerpeton inexpectatum

Related Research Articles

<span class="mw-page-title-main">Lissamphibia</span> Subclass of amphibians

The Lissamphibia is a group of tetrapods that includes all modern amphibians. Lissamphibians consist of three living groups: the Salientia, the Caudata, and the Gymnophiona.

<span class="mw-page-title-main">Caecilian</span> Order of amphibians

Caecilians are a group of limbless, vermiform (worm-shaped) or serpentine (snake-shaped) amphibians with small or sometimes nonexistent eyes. They mostly live hidden in soil or in streambeds, and this cryptic lifestyle renders caecilians among the least familiar amphibians. Modern caecilians live in the tropics of South and Central America, Africa, and southern Asia. Caecilians feed on small subterranean creatures such as earthworms. The body is cylindrical and often darkly coloured, and the skull is bullet-shaped and strongly built. Caecilian heads have several unique adaptations, including fused cranial and jaw bones, a two-part system of jaw muscles, and a chemosensory tentacle in front of the eye. The skin is slimy and bears ringlike markings or grooves and may contain scales.

<span class="mw-page-title-main">Batrachia</span> Clade of amphibians

The Batrachia are a clade of amphibians that includes frogs and salamanders, but not caecilians nor the extinct allocaudates. The name Batrachia was first used by French zoologist Pierre André Latreille in 1800 to refer to frogs, but has more recently been defined in a phylogenetic sense as a node-based taxon that includes the last common ancestor of frogs and salamanders and all of its descendants. The idea that frogs and salamanders are more closely related to each other than either is to caecilians is strongly supported by morphological and molecular evidence; they are, for instance, the only vertebrates able to raise and lower their eyes. However, an alternative hypothesis exists in which salamanders and caecilians are each other's closest relatives as part of a clade called the Procera, with frogs positioned as the sister taxon of this group.

<span class="mw-page-title-main">Dissorophoidea</span> Extinct superfamily of amphibians

Dissorophoidea is a clade of medium-sized, temnospondyl amphibians that appeared during the Moscovian in Euramerica, and continued through to the Late Permian and the Early Triassic of Gondwana. They are distinguished by various details of the skull, and many species seem to have been well adapted for life on land.

<span class="mw-page-title-main">Cryptobranchoidea</span> Suborder of salamanders

The Cryptobranchoidea are a suborder of salamanders found in Asia, European Russia, and the United States. They are known as primitive salamanders, in contrast to Salamandroidea, the advanced salamanders. It has two living subdivisions, Cryptobranchidae, and Hynobiidae, commonly known as Asian salamanders.

<i>Albanerpeton</i> Extinct genus of amphibians

Albanerpeton is an extinct genus of salamander-like 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 undiagnosed species from the Paskapoo Formation, making it by far the most speciose genus in the family. Members of the genus had a robust head and neck which likely allowed them to actively burrow, characteristic of fossorial species, and they lived in a wide range of environments. This genus of amphibian was the last of its order, surviving until into the Early Pleistocene (Gelasian) of northern Italy, and possibly northern Spain, until around 2 million years ago. It likely became extinct when the region developed its present Mediterranean-type climate, having preferred one that was cold and humid. The monophyly of Albanerpeton has recently been questioned, with some authors regarding the genus as paraphyletic.

Apodops is an extinct genus of early caecilians from the Early Eocene Itaboraí Formation of Brazil. The type species of the genus is A. pricei, described based on an isolated and broken trunk vertebra.

<i>Anoualerpeton</i> Extinct genus of amphibians

Anoualerpeton is an extinct genus of lissamphibian in the family Albanerpetontidae. It is the oldest and most primitive albanerpetontid known. Fossils have been found of two different species, Anoualerpeton priscus from the Middle Jurassic (Bathonian) aged Forest Marble and Kilmaluag formations of England and Scotland, and Anoualerpeton unicus from Late Jurassic-Early Cretaceous (Tithonian-Berriasian) Ksar Metlili Formation of Morocco. A. unicus is the only named albanerpetontid from Gondwana.

<i>Celtedens</i> Extinct genus of amphibians

Celtedens is an extinct genus of albanerpetontid amphibian from the Early Cretaceous of England, Spain, Sweden and Italy, and the Late Jurassic of Portugal.

Horezmia is an extinct genus of Mesozoic salamanders. The fossils have been found in Russia. It is comparable to modern advanced salamanders, though its phylogenetic placement within Salamandroidea is uncertain.

Sinerpeton is an extinct genus of salamander from the Late Jurassic of China. It is monotypic and consists of one species, S. fengshanense.

<span class="mw-page-title-main">Salientia</span> Order of amphibians

The Salientia are a total group of amphibians that includes the order Anura, the frogs and toads, and various extinct proto-frogs that are more closely related to the frogs than they are to the Urodela, the salamanders and newts. The oldest fossil "proto-frog" appeared in the early Triassic of Madagascar, but molecular clock dating suggests their origins may extend further back to the Permian, 265 million years ago.

<i>Wesserpeton</i> Extinct genus of amphibians

Wesserpeton is an extinct genus of albanerpetontid amphibian known from the Isle of Wight, southern England.

<span class="mw-page-title-main">Karauridae</span> Extinct family of amphibians

The Karauridae are a family of stem-group salamanders (Caudata) that are known from the Middle Jurassic to Early Cretaceous in Central Asia, Northern Asia and Western Europe. The family includes four members: Karaurus from the Middle-Late Jurassic Karabastau Formation of Kazakhstan, Kokartus from the Middle Jurassic Balabansai Formation of Kyrgyzstan, Marmorerpeton from the Middle Jurassic Forest Marble Formation of England and Kilmaluag Formation of Scotland, and Kuzbassia from the Early Cretaceous (Aptian) Ilek Formation. The members are some of the oldest known salamanders. The family is united by several morphological characters, including sculptured skull roof bones. Like some modern salamanders, karaurids were neotenic. Members of the family likely fed via suction feeding on small fish and invertebrates. The Early Cretaceous Siberian Kulgeriherpeton has been suggested to be a karaurid by some authors.

<i>Shirerpeton</i> Extinct genus of amphibians

Shirerpeton is an extinct genus of albanerpetontid amphibian from the Early Cretaceous Kuwajima Formation, which is located in Japan. The type species is Shirerpeton isajii, which was described by Masumoto & Evans in 2018. Shirerpeton represents the first record of Albanerpetontidae in East Asia and the holotype is SBEI 2459, a small block bearing most of a disarticulated but associated skull with some postcranial elements present as well.

Chinlestegophis is a diminutive Late Triassic stereospondyl that has been interpreted as a putative stem caecilian, a living group of legless burrowing amphibians. If Chinlestegophis is indeed both an advanced stereospondyl and a relative of caecilians, this means that stereospondyls survived to the present day; historically the group was thought to have gone extinct by the early Cretaceous. Chinlestegophis jenkinsi, the type and only species, is known from two partial skulls discovered in the Chinle Formation in Colorado.

The Ksar Metlili Formation is a geological formation in eastern High Atlas of Morocco, it is late Tithonian to Berriasian in age. It is approximately 80 metres (260 ft) thick and primarily consists of mudstone and sandstone, with thin calcareous beds. One of these calcareous beds near the middle of the sequence is an important microvertebrate locality. Subsequent to the original site, several other localities have been sampled. The depositional environment is thought to be near shore deltaic.

<i>Yaksha perettii</i> Extinct species of amphibian

Yaksha perettii is an extinct species of albanerpetontid amphibian, and the only species in the genus Yaksha. It is known from three specimens found in Cenomanian aged Burmese amber from Myanmar. The remains of Yaksha perettii are the best preserved of all albanerpetontids, which usually consist of isolated fragments or crushed flat, and have provided significant insights in the morphology and lifestyle of the group.

This list of fossil amphibians described in 2020 is a list of new taxa of fossil amphibians that were described during the year 2020, as well as other significant discoveries and events related to amphibian paleontology that occurred in 2020.

<i>Funcusvermis</i> Extinct genus of amphibians

Funcusvermis is an extinct genus of stem-caecilian from the Late Triassic of Arizona. It is based on a large sample of jaws and other skull and postcranial fragments, discovered in an approximately 220 million years old layer of rock in the Blue Mesa Member of the Chinle Formation at Petrified Forest National Park. There is a single species, called Funcusvermis gilmorei.

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