Casineria
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''Casineria'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of
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 t ...
which lived about 340-334 million years ago in the Mississippian epoch of the
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferou ...
period. Its generic name, ''Casineria'', is a latinization of Cheese Bay, the site near
Edinburgh Edinburgh ( ; gd, Dùn Èideann ) is the capital city of Scotland and one of its 32 Council areas of Scotland, council areas. Historically part of the county of Midlothian (interchangeably Edinburghshire before 1921), it is located in Lothian ...
,
Scotland Scotland (, ) is a country that is part of the United Kingdom. Covering the northern third of the island of Great Britain, mainland Scotland has a border with England to the southeast and is otherwise surrounded by the Atlantic Ocean to ...
where 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 seve ...
fossil was found. When originally described in 1999, it was identified as a
transitional fossil A transitional fossil is any fossilized remains of a life form that exhibits traits common to both an ancestral group and its derived descendant group. This is especially important where the descendant group is sharply differentiated by gross a ...
noted for its mix of basal (
amphibian Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbo ...
-like) and advanced (
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates ( lizards and snakes) and rhynchocephalia ...
-like) characteristics, putting it at or very near the origin of the
amniote Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are dis ...
s, the group containing all
mammals Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur o ...
,
birds Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
, modern reptiles, and other descendants of their reptile-like
common ancestor Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. All living beings are in fact descendants of a unique ancestor commonly referred to as the last universal comm ...
. However, the sole known fossil is lacking key elements such as a
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
, making exact analysis difficult. As a result, the classification of ''Casineria'' has been more controversial in analyses conducted since 1999. Other proposed affinities include a placement among the lepospondyls, seymouriamorphs, " gephyrostegids", or as a synonym of ''
Caerorhachis ''Caerorhachis'' (meaning "suitable spine" in Greek) is an extinct genus of early tetrapod from the Early Carboniferous of Scotland, probably from the Serpukhovian stage. Its placement within Tetrapoda is uncertain, but it is generally regarded a ...
'', another controversial tetrapod which may have been an early
temnospondyl Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carb ...
.


Description

''Casineria'' was a small animal with a
postcrania Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...
length estimated to have been 15 centimeters. Its small size would have made it ideal for hunting the invertebrates of the
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferou ...
. ''Casineria'' had five fingers, with the bones at the finger tip being tapering and curved. This spurred the initial describers to argue that it was the oldest known animal with claws on each hand, and marks the earliest clawed foot. Claws are extremely rare among amphibians, but ubiquitous among
amniotes Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are dis ...
, so their supposed presence in ''Casineria'' has been considered to be evidence towards its amniote identity. Claws are also a feature intimately bound to the formation of
keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up Scale (anatomy), scales, hair, Nail ...
ous scales in reptiles, so in life, ''Casineria'' would have borne scaly, reptilian-type skin, and would have resembled a small lizard (despite being completely unrelated to true
lizards Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia altho ...
). Under the hypothesis that it was among the first amniotes in the biological sense, it would have laid an amniotic egg not dependent on being laid in water to survive, possibly hiding them in damp vegetation or hollowed out tree stumps. This has been inferred from the fact that ''Casineria'' was found in rocks showing a rather dry environment. In the early Carboniferous period before the appearance of ''Casineria'', vertebrates were primarily aquatic, only spending part of their time on land. ''Casineria'' was believed to be among the first vertebrates to live and reproduce on land. However, later studies have not consistently placed ''Casineria'' as an early amniote. Even the presence of claws has been considered doubtful, as Marjanovic & Laurin (2019) noted that the finger tips were squared-off, rather than pointed.


Discovery

In 1992, an amateur fossil collector spotted the remnants of this four-legged creature on the shore of Cheese Bay, Scotland. For the next five years, the fossil languished at the
National Museum of Scotland The National Museum of Scotland in Edinburgh, Scotland, was formed in 2006 with the merger of the new Museum of Scotland, with collections relating to Scottish antiquities, culture and history, and the adjacent Royal Scottish Museum (opene ...
in Edinburgh while researchers focused on other projects. Consisting of a slab and counterslab ( compression fossil) of a single partial skeleton, the fossil was cataloged with the specimen designation NMS G.1993.54.1. Around 1997, work began to expose the remainder of the fossil from the surrounding matrix. The work revealed that the animal probably lived in an environment much drier than previously understood. The findings were first reported in the April 8, 1999 edition of ''Nature''.


Classification

While retaining a general build like those found in the amphibian reptiliomorph groups like
Seymouriamorpha Seymouriamorpha were a small but widespread group of limbed vertebrates (tetrapods). They have long been considered reptiliomorphs, and most paleontologists may still accept this point of view, but some analyses suggest that seymouriamorphs are s ...
and
Diadectomorpha Diadectomorpha is a clade of large tetrapods that lived in Euramerica during the Carboniferous and Early Permian periods and in Asia during Late Permian (Wuchiapingian), They have typically been classified as advanced reptiliomorphs (transitiona ...
, ''Casineria'' also shows features that tie it in with early reptiles, notably a generally gracile build with light leg bones, unfused ankles and toes terminating in claws. This would enable the animal to use their feet actively in traction, rather than as holdfasts, an indication of a primarily terrestrial lifestyle. These traits have been argued to show that it was more closely related to
amniote Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are dis ...
s than to other known reptiliomorph amphibians. With its advanced features, ''Casineria'' may have been one of the first true
amniote Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are dis ...
s, and thus the first
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates ( lizards and snakes) and rhynchocephalia ...
under traditional classification. In
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
parlance it would have been a
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
amniote, close to, but outside the
crown group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor ...
Amniota (the group consisting of the last common ancestor of
synapsids Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes rept ...
and
sauropsids Sauropsida ("lizard faces") is a clade of amniotes, broadly equivalent to the class Reptilia. Sauropsida is the sister taxon to Synapsida, the other clade of amniotes which includes mammals as its only modern representatives. Although early syn ...
and all its descendants). ''Casineria'' pushes back the origin of amniote lineages much earlier than was previously assumed. However, like with much of the basal tetrapod stock, the phylogenetic position is uncertain, and it has also been suggested ''Casineria'' is a lepospondyl,
seymouriamorph Seymouriamorpha were a small but widespread group of limbed vertebrates (tetrapods). They have long been considered reptiliomorphs, and most paleontologists may still accept this point of view, but some analyses suggest that seymouriamorphs are s ...
, synonymous with ''
Caerorhachis ''Caerorhachis'' (meaning "suitable spine" in Greek) is an extinct genus of early tetrapod from the Early Carboniferous of Scotland, probably from the Serpukhovian stage. Its placement within Tetrapoda is uncertain, but it is generally regarded a ...
'' (which is possibly a basal
temnospondyl Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carb ...
amphibian),Chapter 6: "Walking with early tetrapods: evolution of the postcranial skeleton and the phylogenetic affinities of the Temnospondyli (Vertebrata: Tetrapoda)." ''In'': Kat Pawley (2006).
The postcranial skeleton of temnospondyls (Tetrapoda: temnospondyli)
" PhD Thesis. La Trobe University, Melbourne.
or part of a grade of small tetrapods traditionally considered the family Gephyrostegidae.


See also

*''
Solenodonsaurus ''Solenodonsaurus'' ("single-tooth lizard") is an extinct genus of reptiliomorphs that lived in what is now Czech Republic, during the Westphalian stage. Description ''Solenondosaurus'' had snout-vent length with a skull length . ''Solenod ...
'' *''
Westlothiana ''Westlothiana'' ("animal from West Lothian") is a genus of reptile-like tetrapod that lived about 338 million years ago during the latest part of the Visean age of the Carboniferous. Members of the genus bore a superficial resemblance to modern ...
''


References


Further reading

* Richard Monastersky
"Out of the Swamps: How early vertebrates established a foothold—with all 10 toes—on land"
''The Weekly Newsmagazine of Science'', Volume 155, Number 21 (May 22, 1999) *The Encyclopedia of Dinosaurs & Prehistoric Creatures/Dougal Dixon {{Taxonbar, from=Q1094145 Carboniferous tetrapods of Europe Enigmatic vertebrate taxa Fossil taxa described in 1999