Caseidae - Revision history

Bazza 7: Updated short description

2024-10-15T07:21:25Z

Updated short description

← Previous revision		Revision as of 07:21, 15 October 2024
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	{{Short description\|Extinct family of ~~synapsids~~}}		{{Short description\|Extinct family of vertebrate animals}}
	{{Automatic taxobox		{{Automatic taxobox
	\| fossil_range = [[Late Carboniferous]] - [[Middle Permian]], {{fossil_range\|300\|265\|}}		\| fossil_range = [[Late Carboniferous]] - [[Middle Permian]], {{fossil_range\|300\|265\|}}

Dimadick: /* External links */

2024-10-06T09:11:22Z

External links

← Previous revision		Revision as of 09:11, 6 October 2024
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	{{Taxonbar\|from=Q132725}}		{{Taxonbar\|from=Q132725}}

			[[Category:Caseidae\| ]]
	[[Category:Caseasaurs]]		[[Category:Caseasaurs]]
	[[Category:Prehistoric synapsid families]]		[[Category:Prehistoric synapsid families]]

Crystalespeon: no comma needed

2024-08-22T15:11:12Z

no comma needed

← Previous revision		Revision as of 15:11, 22 August 2024
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	\|footer=''Cotylorhynchus romeri''. Top : skull reconstruction in left lateral view, and medial and lateral views of mandible; bottom : skull reconstruction in dorsal, ventral, posterior and anterior views.}}[[File:Euromycter rutenus 6778 retouched.jpg\|thumb\|right\|Skull of ''[[Euromycter\|Euromycter rutenus]]'' in ventral view allowing to see the numerous small teeth adorning the bones of the palate (the [[parasphenoid]], triangular in shape in the center, and the very elongated [[Pterygoid bone\|pterygoids]] on each side). The hyoid apparatus is not present here because it was removed during the preparation of the palate.]]		\|footer=''Cotylorhynchus romeri''. Top : skull reconstruction in left lateral view, and medial and lateral views of mandible; bottom : skull reconstruction in dorsal, ventral, posterior and anterior views.}}[[File:Euromycter rutenus 6778 retouched.jpg\|thumb\|right\|Skull of ''[[Euromycter\|Euromycter rutenus]]'' in ventral view allowing to see the numerous small teeth adorning the bones of the palate (the [[parasphenoid]], triangular in shape in the center, and the very elongated [[Pterygoid bone\|pterygoids]] on each side). The hyoid apparatus is not present here because it was removed during the preparation of the palate.]]

	Caseids measured from less than {{convert\|1\|m}} to {{convert\|7\|m}} in length.<ref name="Romano2017">{{cite journal\|last1=Romano \|first1=M. \|year=2017 \|title=Long bone scaling of caseid synapsids: a combined morphometric and cladistic approach \|journal=Lethaia \|volume=50 \|issue=4 \|pages=511–526 \|doi=10.1111/let.12207}}</ref><ref name="Angielczyk&Kammerer2018">{{cite book\|last=Angielczyk\|first=K.D. \|author2=Kammerer, C.F. \|year=2018 \|chapter=Non-Mammalian synapsids : the deep roots of the mammalian family tree \|pages=117–198 \|title=Handbook of Zoology : Mammalian Evolution, Diversity and Systematics \|editor1-last=Zachos\|editor1-first=F.E. \|editor2=Asher, R.J. \|publisher=de Gruyter \|location=Berlin \|isbn=978-3-11-027590-2}}</ref><ref name="Romano&al.2018">{{cite journal\|last1=Romano\|first1=M. \|author2=Citton, P. \|author3=Maganuco, S. \|author4=Sacchi, E. \|author5=Caratelli, M. \|author6=Ronchi, A. \|author7=Nicosia, U. \|year=2018 \|title=New basal synapsid discovery at the Permian outcrop of Torre del Porticciolo (Alghero, Italy) \|journal=Geological Journal \|volume=54 \|issue=3 \|pages=1554–1566 \|doi=10.1002/gj.3250\|s2cid=133755506 }}</ref> They had a small head wider than high and with a forward-inclined snout, a very short neck, a long tail, robust forelimbs, and a body of variable proportions depending on their diet. Small insectivorous species like ''[[Eocasea]]'' had an unexpanded trunk.<ref name="Reisz&Fröbisch2014">{{cite journal\|last1=Reisz\|first1=R.R. \|author2=Fröbisch, J. \|year=2014 \|title=Oldest caseid synapsid from the late Pennsylvanian of Kansas, and the evolution of herbivory in terrestrial vertebrates \|journal=PLOS ONE \|volume=9 \|issue=4 \|pages=e94518 \|doi=10.1371/journal.pone.0094518 \|pmid=24739998 \|pmc=3989228\|bibcode=2014PLoSO...994518R \|doi-access=free }}</ref> Others with an omnivorous diet like ''[[Martensius]]'' had a barely enlarged rib cage, a more elongated skull, smaller [[nostril]]s, and a snout less inclined forward than in herbivorous caseids.<ref name="Berman&al.2020">{{cite journal\|last1=Berman \|first1=D.S. \|author2=Maddin, H.C. \|author3=Henrici, A.C. \|author4=Sumida, S.S. \|author5=Scott, D. \|author6=Reisz, R.R. \|title=New primitive Caseid (Synapsida, Caseasauria) from the Early Permian of Germany \|journal=Annals of Carnegie Museum \|year=2020 \|volume=86 \|issue=1 \|pages=43–75 \|doi=10.2992/007.086.0103\|s2cid=216027787 }}</ref> The latter were characterized by their disproportionately small [[skull]] compared to the size of the body. The postcranial skeleton indeed shows a spectacular increase in the volume of the [[rib cage]], which becomes very wide and barrel-shaped, probably to accommodate a particularly developed [[Gastrointestinal tract#Lower gastrointestinal tract\|intestine]], necessary for the digestion of high-fiber rich plants. In these forms, the skull has very large external nostrils and a very short facial region with a strong forward inclination of the end of the snout which clearly overhangs the dental row. The [[Skull#Fenestrae\|temporal fenestrae]] are also relatively large (especially in ''[[Ennatosaurus]]''), the [[Supratemporal bone\|supratemporals]] are large in size, and, on the [[Occipital bone\|occipital]] surface, the paroccipital [[Process (anatomy)\|processes]] are massively developed, establishing strong supporting contacts with the [[Squamosal bone\|squamosals]].<ref name="Kemp1982">{{cite book\|last=Kemp \|first=T.S. \|year=1982 \|chapter=Pelycosaurs \|pages=39 \|title=Mammal-like reptiles and the origin of Mammals \|editor1-last=Kemp\|editor1-first=T.S. \|publisher=Academic Press \|location=London \|isbn=978-0124041202}}</ref> The dorsal surface of the skull is covered with numerous small pits. These suggest the presence of large [[Reptile scale\|scales]] on the head of these animals.<ref name="Romano&al.2017a">{{cite journal\|last1=Romano \|first1=M. \|author2=Brocklehurst, N. \|author3=Fröbisch, J. \|year=2017 \|title=The postcranial skeleton of ''Ennatosaurus tecton'' (Synapsida, Caseidae) \|journal=Journal of Systematic Palaeontology \|volume=16 \|issue= 13\|pages=1097–1122 \|doi=10.1080/14772019.2017.1367729\|s2cid=89922565 }}</ref> Numerous [[Lip\|labial]] [[Foramen\|foramina]] running parallel to the ventral edge of the [[premaxilla]] and [[maxilla]], as well as along the dorsal edge of the [[Mandible#Other vertebrates\|dentary]], suggest the presence of scaly "lips" which must have concealed the dentition when the jaws were closed.<ref name="Reisz&al.2022">{{cite journal\|last1=Reisz \|first1=R.R. \|author2=Scott, D. \|author3=Modesto, S.P. \|year=2022 \|title=Cranial Anatomy of the Caseid Synapsid ''Cotylorhynchus romeri'', a Large Terrestrial Herbivore From the Lower Permian of Oklahoma, U.S.A \|journal=Frontiers in Earth Science \|volume=10 \|issue= \|pages=1–19 \|doi=10.3389/feart.2022.847560\|doi-access=free }}</ref><ref name="Romano&al.2017a"/> The [[Tooth\|teeth]], simply conical and pointed in insectivorous species, adopt in herbivorous species a leafy or spatulate morphology and are provided with more or less numerous cuspules. Numerous small teeth also adorned several bones of the palate. Herbivorous species do not show a simple evolutionary trend towards increasing tooth complexity.<ref name="Maddin&al.2008">{{cite journal \|last1=Maddin \|first1=H.C. \|author2=Sidor, C.A. \|author3=Reisz, R.R. \|year=2008 \|title=Cranial anatomy of ''Ennatosaurus tecton'' (Synapsida: Caseidae) from the Middle Permian of Russia and the evolutionary relationships of Caseidae \|journal=Journal of Vertebrate Paleontology \|volume=28 \|issue=1 \|pages=160–180 \|doi=10.1671/0272-4634(2008)28[160:CAOETS]2.0.CO;2\|s2cid=44064927 }}</ref> Thus, the teeth of the basal taxa ''[[Casea]]'' and ''[[Arisierpeton]]'' have three cuspules<ref name="Olson1968">{{cite journal\|last1=Olson \|first1=E.C. \|year=1968 \|title=The family Caseidae \|journal=Fieldiana: Geology \|volume=17 \|issue= \|pages=225–349\| doi= }}</ref><ref name="Reisz2019">{{cite journal \|last1=Reisz \|first1=R. \|year=2019 \|title=A small caseid synapsid, ''Arisierpeton simplex'' gen. et sp. nov., from the early Permian of Oklahoma, with a discussion of synapsid diversity at the classic Richards Spur locality \|journal=PeerJ \|volume=7e6615 \|issue= \|pages=1–20 \|doi=10.7717/peerj.6615 \|pmid=30997285 \|pmc=6462398 \|doi-access=free }}</ref> just like in the more derived forms ''[[Cotylorhynchus]]'' and ''[[Caseopsis]]''.<ref name="Olson1968"/> ''Ennatosaurus'' and ''[[Euromycter]]'', which occupy an intermediate [[Phylogenetics\|phylogenetic]] position, have teeth bearing 5 to 7 cuspules and 5 to 8 cuspules respectively.<ref name="Maddin&al.2008"/><ref name="sigogneau-russell1974">{{cite journal \|last1=Sigoneau-Russell \|first1=D. \|author2=Russel, D.E. \|year=1974 \|title=Étude du premier caséidé (Reptilia, Pelycosauria) d'Europe occidentale \|journal=Bulletin du Muséum National d'Histoire Naturelle \|series=Série 3 \|volume=38 \|issue=230 \|pages=145–215 \|doi= \|url=http://bibliotheques.mnhn.fr/EXPLOITATION/infodoc/digitalCollections/viewerpopup.aspx?seid=BMSCT_S003_1974_T230_N038}}</ref> ''[[Angelosaurus]]'', one of the most derived caseids, has teeth with 5 cuspules.<ref name="Olson1968"/> In ''Angelosaurus'' the teeth have a bulbous morphology with very short and wide crowns. Their sturdiness and the significant wear they show, indicate that ''Angelosaurus'' must have fed on tougher plants than those on which most other herbivorous caseids fed.<ref name="Olson1968"/> Herbivorous caseids also show very different dietary adaptations from those seen in another group of basal synapsids, the [[Edaphosauridae]]. The latter had, in addition to the marginal dentition, a dental battery made up of numerous teeth located both on the [[palate]] and on the inner surface of the lower jaws. In herbivorous caseids, the palatal teeth are smaller, and the inner surface of the lower jaws bears no teeth. Instead of a dental battery, they had a massive [[tongue]] (as indicated by the presence of a highly developed [[hyoid apparatus]] found in ''Ennatosaurus'' and ''Euromycter'') perhaps rough, with which they had to compress food against the palatal teeth.<ref name="Olson1968"/><ref name="Reisz&al.2022"/>		Caseids measured from less than {{convert\|1\|m}} to {{convert\|7\|m}} in length.<ref name="Romano2017">{{cite journal\|last1=Romano \|first1=M. \|year=2017 \|title=Long bone scaling of caseid synapsids: a combined morphometric and cladistic approach \|journal=Lethaia \|volume=50 \|issue=4 \|pages=511–526 \|doi=10.1111/let.12207}}</ref><ref name="Angielczyk&Kammerer2018">{{cite book\|last=Angielczyk\|first=K.D. \|author2=Kammerer, C.F. \|year=2018 \|chapter=Non-Mammalian synapsids : the deep roots of the mammalian family tree \|pages=117–198 \|title=Handbook of Zoology : Mammalian Evolution, Diversity and Systematics \|editor1-last=Zachos\|editor1-first=F.E. \|editor2=Asher, R.J. \|publisher=de Gruyter \|location=Berlin \|isbn=978-3-11-027590-2}}</ref><ref name="Romano&al.2018">{{cite journal\|last1=Romano\|first1=M. \|author2=Citton, P. \|author3=Maganuco, S. \|author4=Sacchi, E. \|author5=Caratelli, M. \|author6=Ronchi, A. \|author7=Nicosia, U. \|year=2018 \|title=New basal synapsid discovery at the Permian outcrop of Torre del Porticciolo (Alghero, Italy) \|journal=Geological Journal \|volume=54 \|issue=3 \|pages=1554–1566 \|doi=10.1002/gj.3250\|s2cid=133755506 }}</ref> They had a small head wider than high and with a forward-inclined snout, a very short neck, a long tail, robust forelimbs, and a body of variable proportions depending on their diet. Small insectivorous species like ''[[Eocasea]]'' had an unexpanded trunk.<ref name="Reisz&Fröbisch2014">{{cite journal\|last1=Reisz\|first1=R.R. \|author2=Fröbisch, J. \|year=2014 \|title=Oldest caseid synapsid from the late Pennsylvanian of Kansas, and the evolution of herbivory in terrestrial vertebrates \|journal=PLOS ONE \|volume=9 \|issue=4 \|pages=e94518 \|doi=10.1371/journal.pone.0094518 \|pmid=24739998 \|pmc=3989228\|bibcode=2014PLoSO...994518R \|doi-access=free }}</ref> Others with an omnivorous diet like ''[[Martensius]]'' had a barely enlarged rib cage, a more elongated skull, smaller [[nostril]]s, and a snout less inclined forward than in herbivorous caseids.<ref name="Berman&al.2020">{{cite journal\|last1=Berman \|first1=D.S. \|author2=Maddin, H.C. \|author3=Henrici, A.C. \|author4=Sumida, S.S. \|author5=Scott, D. \|author6=Reisz, R.R. \|title=New primitive Caseid (Synapsida, Caseasauria) from the Early Permian of Germany \|journal=Annals of Carnegie Museum \|year=2020 \|volume=86 \|issue=1 \|pages=43–75 \|doi=10.2992/007.086.0103\|s2cid=216027787 }}</ref> The latter were characterized by their disproportionately small [[skull]] compared to the size of the body. The postcranial skeleton indeed shows a spectacular increase in the volume of the [[rib cage]], which becomes very wide and barrel-shaped, probably to accommodate a particularly developed [[Gastrointestinal tract#Lower gastrointestinal tract\|intestine]], necessary for the digestion of high-fiber rich plants. In these forms, the skull has very large external nostrils and a very short facial region with a strong forward inclination of the end of the snout which clearly overhangs the dental row. The [[Skull#Fenestrae\|temporal fenestrae]] are also relatively large (especially in ''[[Ennatosaurus]]''), the [[Supratemporal bone\|supratemporals]] are large in size, and, on the [[Occipital bone\|occipital]] surface, the paroccipital [[Process (anatomy)\|processes]] are massively developed, establishing strong supporting contacts with the [[Squamosal bone\|squamosals]].<ref name="Kemp1982">{{cite book\|last=Kemp \|first=T.S. \|year=1982 \|chapter=Pelycosaurs \|pages=39 \|title=Mammal-like reptiles and the origin of Mammals \|editor1-last=Kemp\|editor1-first=T.S. \|publisher=Academic Press \|location=London \|isbn=978-0124041202}}</ref> The dorsal surface of the skull is covered with numerous small pits. These suggest the presence of large [[Reptile scale\|scales]] on the head of these animals.<ref name="Romano&al.2017a">{{cite journal\|last1=Romano \|first1=M. \|author2=Brocklehurst, N. \|author3=Fröbisch, J. \|year=2017 \|title=The postcranial skeleton of ''Ennatosaurus tecton'' (Synapsida, Caseidae) \|journal=Journal of Systematic Palaeontology \|volume=16 \|issue= 13\|pages=1097–1122 \|doi=10.1080/14772019.2017.1367729\|s2cid=89922565 }}</ref> Numerous [[Lip\|labial]] [[Foramen\|foramina]] running parallel to the ventral edge of the [[premaxilla]] and [[maxilla]], as well as along the dorsal edge of the [[Mandible#Other vertebrates\|dentary]], suggest the presence of scaly "lips" which must have concealed the dentition when the jaws were closed.<ref name="Reisz&al.2022">{{cite journal\|last1=Reisz \|first1=R.R. \|author2=Scott, D. \|author3=Modesto, S.P. \|year=2022 \|title=Cranial Anatomy of the Caseid Synapsid ''Cotylorhynchus romeri'', a Large Terrestrial Herbivore From the Lower Permian of Oklahoma, U.S.A \|journal=Frontiers in Earth Science \|volume=10 \|issue= \|pages=1–19 \|doi=10.3389/feart.2022.847560\|doi-access=free }}</ref><ref name="Romano&al.2017a"/> The [[Tooth\|teeth]], simply conical and pointed in insectivorous species, adopt in herbivorous species a leafy or spatulate morphology and are provided with more or less numerous cuspules. Numerous small teeth also adorned several bones of the palate. Herbivorous species do not show a simple evolutionary trend towards increasing tooth complexity.<ref name="Maddin&al.2008">{{cite journal \|last1=Maddin \|first1=H.C. \|author2=Sidor, C.A. \|author3=Reisz, R.R. \|year=2008 \|title=Cranial anatomy of ''Ennatosaurus tecton'' (Synapsida: Caseidae) from the Middle Permian of Russia and the evolutionary relationships of Caseidae \|journal=Journal of Vertebrate Paleontology \|volume=28 \|issue=1 \|pages=160–180 \|doi=10.1671/0272-4634(2008)28[160:CAOETS]2.0.CO;2\|s2cid=44064927 }}</ref> Thus, the teeth of the basal taxa ''[[Casea]]'' and ''[[Arisierpeton]]'' have three cuspules<ref name="Olson1968">{{cite journal\|last1=Olson \|first1=E.C. \|year=1968 \|title=The family Caseidae \|journal=Fieldiana: Geology \|volume=17 \|issue= \|pages=225–349\| doi= }}</ref><ref name="Reisz2019">{{cite journal \|last1=Reisz \|first1=R. \|year=2019 \|title=A small caseid synapsid, ''Arisierpeton simplex'' gen. et sp. nov., from the early Permian of Oklahoma, with a discussion of synapsid diversity at the classic Richards Spur locality \|journal=PeerJ \|volume=7e6615 \|issue= \|pages=1–20 \|doi=10.7717/peerj.6615 \|pmid=30997285 \|pmc=6462398 \|doi-access=free }}</ref> just like in the more derived forms ''[[Cotylorhynchus]]'' and ''[[Caseopsis]]''.<ref name="Olson1968"/> ''Ennatosaurus'' and ''[[Euromycter]]'', which occupy an intermediate [[Phylogenetics\|phylogenetic]] position, have teeth bearing 5 to 7 cuspules and 5 to 8 cuspules respectively.<ref name="Maddin&al.2008"/><ref name="sigogneau-russell1974">{{cite journal \|last1=Sigoneau-Russell \|first1=D. \|author2=Russel, D.E. \|year=1974 \|title=Étude du premier caséidé (Reptilia, Pelycosauria) d'Europe occidentale \|journal=Bulletin du Muséum National d'Histoire Naturelle \|series=Série 3 \|volume=38 \|issue=230 \|pages=145–215 \|doi= \|url=http://bibliotheques.mnhn.fr/EXPLOITATION/infodoc/digitalCollections/viewerpopup.aspx?seid=BMSCT_S003_1974_T230_N038}}</ref> ''[[Angelosaurus]]'', one of the most derived caseids, has teeth with 5 cuspules.<ref name="Olson1968"/> In ''Angelosaurus'' the teeth have a bulbous morphology with very short and wide crowns. Their sturdiness and the significant wear they show indicate that ''Angelosaurus'' must have fed on tougher plants than those on which most other herbivorous caseids fed.<ref name="Olson1968"/> Herbivorous caseids also show very different dietary adaptations from those seen in another group of basal synapsids, the [[Edaphosauridae]]. The latter had, in addition to the marginal dentition, a dental battery made up of numerous teeth located both on the [[palate]] and on the inner surface of the lower jaws. In herbivorous caseids, the palatal teeth are smaller, and the inner surface of the lower jaws bears no teeth. Instead of a dental battery, they had a massive [[tongue]] (as indicated by the presence of a highly developed [[hyoid apparatus]] found in ''Ennatosaurus'' and ''Euromycter'') perhaps rough, with which they had to compress food against the palatal teeth.<ref name="Olson1968"/><ref name="Reisz&al.2022"/>

	[[File:Arisierpeton simplex, premaxillae.png\|thumb\|right\|Premaxillae of ''[[Arisierpeton\|Arisierpeton simplex]]''. (A) Specimen GAA 00242 in anterior, labial, posterior, ventral, and dorsal views; (B) Specimen GAA 00239 in labial (showing a complete tooth with tricuspid distal end), partial labial, posterior and ventral views.]][[File:Cotylorhynchus romeria from Norman, Oklahoma.jpg\|thumb\|left\|Skeleton of ''Cotylorhynchus romeri'' on display at the [[Sam Noble Oklahoma Museum of Natural History]].]]		[[File:Arisierpeton simplex, premaxillae.png\|thumb\|right\|Premaxillae of ''[[Arisierpeton\|Arisierpeton simplex]]''. (A) Specimen GAA 00242 in anterior, labial, posterior, ventral, and dorsal views; (B) Specimen GAA 00239 in labial (showing a complete tooth with tricuspid distal end), partial labial, posterior and ventral views.]][[File:Cotylorhynchus romeria from Norman, Oklahoma.jpg\|thumb\|left\|Skeleton of ''Cotylorhynchus romeri'' on display at the [[Sam Noble Oklahoma Museum of Natural History]].]]

OAbot: Open access bot: hdl updated in citation with #oabot.

2024-01-04T01:08:47Z

Open access bot: hdl updated in citation with #oabot.

← Previous revision		Revision as of 01:08, 4 January 2024
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	}}		}}

	In 2017 Romano and colleagues published the first phylogenetic analysis including the genus ''[[Alierasaurus]]''. The latter is recovered as the sister taxon of the genus ''Cotylorhynchus''.<ref name="Romano&al.2017b">{{cite journal\|last1=Romano \|first1=M. \|author2=Ronchi, A. \|author3=Maganuco, S. \|author4=Nicosia, U. \|year=2017 \|title=New material of ''Alierasaurus ronchii'' (Synapsida, Caseidae) from the Permian of Sardinia (Italy), and its phylogenetic affinities \|journal=Palaeontologia Electronica \|volume=20.2.26A \|issue= \|pages=1–27 \|doi=10.26879/684\|s2cid=73577681 \|doi-access=free }}</ref>		In 2017 Romano and colleagues published the first phylogenetic analysis including the genus ''[[Alierasaurus]]''. The latter is recovered as the sister taxon of the genus ''Cotylorhynchus''.<ref name="Romano&al.2017b">{{cite journal\|last1=Romano \|first1=M. \|author2=Ronchi, A. \|author3=Maganuco, S. \|author4=Nicosia, U. \|year=2017 \|title=New material of ''Alierasaurus ronchii'' (Synapsida, Caseidae) from the Permian of Sardinia (Italy), and its phylogenetic affinities \|journal=Palaeontologia Electronica \|volume=20.2.26A \|issue= \|pages=1–27 \|doi=10.26879/684\|s2cid=73577681 \|doi-access=free \|hdl=11573/1045550 \|hdl-access=free }}</ref>

	Below is the phylogenetic analysis published by Romano and colleagues in 2017.<ref name="Romano&al.2017b"/>		Below is the phylogenetic analysis published by Romano and colleagues in 2017.<ref name="Romano&al.2017b"/>

Anaxial: Undid revision 1192793414 by AB-Babayo (talk) - link is already free to read

2023-12-31T13:04:28Z

Undid revision 1192793414 by AB-Babayo (talk) - link is already free to read

← Previous revision		Revision as of 13:04, 31 December 2023
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	The family Caseidae was erected by [[Samuel Wendell Williston]] in 1911.<ref name="Williston1911">{{cite journal\|last1=Williston \|first1=S.W. \|year=1911 \|title=Permian Reptiles \|journal=Sciences \|volume=33 \|issue=851 \|pages=631–632 \|doi=10.1126/science.33.851.631\|pmid=17798180 \|bibcode=1911Sci....33..631W \|url=https://zenodo.org/record/2413774 }}</ref> In 1940, [[Alfred Romer\|Alfred Sherwood Romer]] and L.W. Price placed Caseidae and [[Edaphosauridae]] in the same suborder, which they named Edaphosauria.<ref name="Romer&Price1940">{{cite journal\|last1=Romer \|first1=A.S. \|author2=Price, L.I. \|year=1940 \|title=Review of the pelycosauria \|journal=Geological Society of America Special Paper \|series=Geological Society of America Special Papers \|volume=28 \|issue= \|pages=1–538 \| doi=10.1130/SPE28-p1}}</ref> This group is now abandoned, the edaphosaurids being closer to the [[Sphenacodontia]] (with which they form the [[clade]] [[Eupelycosauria\|Sphenacomorpha]]) than to the caseids.<ref name="Spindler&al.2019"/> The latter are grouped with the [[Eothyrididae]] in the clade [[Caseasauria]], which represent the most [[Basal (phylogenetics)\|basal]] [[synapsid]]s.<ref name="Brocklehurst&al.2016">{{cite journal\|last1=Broklehurst \|first1=N. \|author2=Reisz, R. \|author3=Fernandez, V. \|author4=Fröbisch, J. \|year=2016 \|title=A Re-Description of 'Mycterosaurus' smithae, an Early Permian Eothyridid, and Its Impact on the Phylogeny of Pelycosaurian-Grade Synapsids \|journal=PLOS ONE \|volume=11 \|issue=6 \|pages=e0156810 \|doi=10.1371/journal.pone.0156810 \|pmid=27333277 \|pmc=4917111 \|bibcode=2016PLoSO..1156810B \|doi-access=free }}</ref>		The family Caseidae was erected by [[Samuel Wendell Williston]] in 1911.<ref name="Williston1911">{{cite journal\|last1=Williston \|first1=S.W. \|year=1911 \|title=Permian Reptiles \|journal=Sciences \|volume=33 \|issue=851 \|pages=631–632 \|doi=10.1126/science.33.851.631\|pmid=17798180 \|bibcode=1911Sci....33..631W \|url=https://zenodo.org/record/2413774 }}</ref> In 1940, [[Alfred Romer\|Alfred Sherwood Romer]] and L.W. Price placed Caseidae and [[Edaphosauridae]] in the same suborder, which they named Edaphosauria.<ref name="Romer&Price1940">{{cite journal\|last1=Romer \|first1=A.S. \|author2=Price, L.I. \|year=1940 \|title=Review of the pelycosauria \|journal=Geological Society of America Special Paper \|series=Geological Society of America Special Papers \|volume=28 \|issue= \|pages=1–538 \| doi=10.1130/SPE28-p1}}</ref> This group is now abandoned, the edaphosaurids being closer to the [[Sphenacodontia]] (with which they form the [[clade]] [[Eupelycosauria\|Sphenacomorpha]]) than to the caseids.<ref name="Spindler&al.2019"/> The latter are grouped with the [[Eothyrididae]] in the clade [[Caseasauria]], which represent the most [[Basal (phylogenetics)\|basal]] [[synapsid]]s.<ref name="Brocklehurst&al.2016">{{cite journal\|last1=Broklehurst \|first1=N. \|author2=Reisz, R. \|author3=Fernandez, V. \|author4=Fröbisch, J. \|year=2016 \|title=A Re-Description of 'Mycterosaurus' smithae, an Early Permian Eothyridid, and Its Impact on the Phylogeny of Pelycosaurian-Grade Synapsids \|journal=PLOS ONE \|volume=11 \|issue=6 \|pages=e0156810 \|doi=10.1371/journal.pone.0156810 \|pmid=27333277 \|pmc=4917111 \|bibcode=2016PLoSO..1156810B \|doi-access=free }}</ref>

	The first [[Phylogenetics\|phylogenetic]] analysis of caseids was published in 2008 by Hillary C. Maddin and colleagues. In this analysis, the genus ''[[Oromycter]]'' occupies the most basal position within the clade. ''[[Ennatosaurus]]'' is the [[sister group]] of a clade containing ''[[Cotylorhynchus\|Cotylorhynchus romeri]]'' and ''[[Angelosaurus\|Angelosaurus dolani]]''. This analysis reveals for the first time the [[paraphyly]] of the [[genus]] ''[[Casea]]'', the species ''“Casea” rutena'' representing a distinct genus which will be named in 2011 ''[[Euromycter]]''.<ref name="Maddin&al.2008"/><ref name=Reisz&al.2011>{{cite journal\|last1=Reisz \|first1=R.R. \|author2=Maddin, H.C. \|author3=Fröbisch, J. \|author4=Falconnet, J. \|year=2011 \|title=A new large caseid (Synapsida, Caseasauria) from the Permian of Rodez (France), including a reappraisal of ''"Casea" rutena'' Sigogneau-Russell & Russell, 1974 \|journal=Geodiversitas \|volume=33 \|issue=2 \|pages=227–246 \| doi= 10.5252/g2011n2a2\|s2cid=129458820 \|url=~~https~~://~~zenodo~~.~~org~~/~~record~~/~~4597072~~/~~files~~/~~source.pdf~~}}</ref>		The first [[Phylogenetics\|phylogenetic]] analysis of caseids was published in 2008 by Hillary C. Maddin and colleagues. In this analysis, the genus ''[[Oromycter]]'' occupies the most basal position within the clade. ''[[Ennatosaurus]]'' is the [[sister group]] of a clade containing ''[[Cotylorhynchus\|Cotylorhynchus romeri]]'' and ''[[Angelosaurus\|Angelosaurus dolani]]''. This analysis reveals for the first time the [[paraphyly]] of the [[genus]] ''[[Casea]]'', the species ''“Casea” rutena'' representing a distinct genus which will be named in 2011 ''[[Euromycter]]''.<ref name="Maddin&al.2008"/><ref name=Reisz&al.2011>{{cite journal\|last1=Reisz \|first1=R.R. \|author2=Maddin, H.C. \|author3=Fröbisch, J. \|author4=Falconnet, J. \|year=2011 \|title=A new large caseid (Synapsida, Caseasauria) from the Permian of Rodez (France), including a reappraisal of ''"Casea" rutena'' Sigogneau-Russell & Russell, 1974 \|journal=Geodiversitas \|volume=33 \|issue=2 \|pages=227–246 \| doi= 10.5252/g2011n2a2\|s2cid=129458820 \|url=http://sciencepress.mnhn.fr/fr/periodiques/geodiversitas/33/2/un-nouveau-grand-caseide-synapsida-caseasauria-du-permien-du-bassin-de-rodez-france-et-reevaluation-de-casea-rutena-sigogneau-russell-russell-1974}}</ref>

	Below is the first [[cladogram]] of caseids published by Maddin et al. in 2008.<ref name="Maddin&al.2008"/>		Below is the first [[cladogram]] of caseids published by Maddin et al. in 2008.<ref name="Maddin&al.2008"/>

AB-Babayo: Added free to read link in citations with OAbot #oabot

2023-12-31T10:09:43Z

Added free to read link in citations with OAbot #oabot

← Previous revision		Revision as of 10:09, 31 December 2023
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	The family Caseidae was erected by [[Samuel Wendell Williston]] in 1911.<ref name="Williston1911">{{cite journal\|last1=Williston \|first1=S.W. \|year=1911 \|title=Permian Reptiles \|journal=Sciences \|volume=33 \|issue=851 \|pages=631–632 \|doi=10.1126/science.33.851.631\|pmid=17798180 \|bibcode=1911Sci....33..631W \|url=https://zenodo.org/record/2413774 }}</ref> In 1940, [[Alfred Romer\|Alfred Sherwood Romer]] and L.W. Price placed Caseidae and [[Edaphosauridae]] in the same suborder, which they named Edaphosauria.<ref name="Romer&Price1940">{{cite journal\|last1=Romer \|first1=A.S. \|author2=Price, L.I. \|year=1940 \|title=Review of the pelycosauria \|journal=Geological Society of America Special Paper \|series=Geological Society of America Special Papers \|volume=28 \|issue= \|pages=1–538 \| doi=10.1130/SPE28-p1}}</ref> This group is now abandoned, the edaphosaurids being closer to the [[Sphenacodontia]] (with which they form the [[clade]] [[Eupelycosauria\|Sphenacomorpha]]) than to the caseids.<ref name="Spindler&al.2019"/> The latter are grouped with the [[Eothyrididae]] in the clade [[Caseasauria]], which represent the most [[Basal (phylogenetics)\|basal]] [[synapsid]]s.<ref name="Brocklehurst&al.2016">{{cite journal\|last1=Broklehurst \|first1=N. \|author2=Reisz, R. \|author3=Fernandez, V. \|author4=Fröbisch, J. \|year=2016 \|title=A Re-Description of 'Mycterosaurus' smithae, an Early Permian Eothyridid, and Its Impact on the Phylogeny of Pelycosaurian-Grade Synapsids \|journal=PLOS ONE \|volume=11 \|issue=6 \|pages=e0156810 \|doi=10.1371/journal.pone.0156810 \|pmid=27333277 \|pmc=4917111 \|bibcode=2016PLoSO..1156810B \|doi-access=free }}</ref>		The family Caseidae was erected by [[Samuel Wendell Williston]] in 1911.<ref name="Williston1911">{{cite journal\|last1=Williston \|first1=S.W. \|year=1911 \|title=Permian Reptiles \|journal=Sciences \|volume=33 \|issue=851 \|pages=631–632 \|doi=10.1126/science.33.851.631\|pmid=17798180 \|bibcode=1911Sci....33..631W \|url=https://zenodo.org/record/2413774 }}</ref> In 1940, [[Alfred Romer\|Alfred Sherwood Romer]] and L.W. Price placed Caseidae and [[Edaphosauridae]] in the same suborder, which they named Edaphosauria.<ref name="Romer&Price1940">{{cite journal\|last1=Romer \|first1=A.S. \|author2=Price, L.I. \|year=1940 \|title=Review of the pelycosauria \|journal=Geological Society of America Special Paper \|series=Geological Society of America Special Papers \|volume=28 \|issue= \|pages=1–538 \| doi=10.1130/SPE28-p1}}</ref> This group is now abandoned, the edaphosaurids being closer to the [[Sphenacodontia]] (with which they form the [[clade]] [[Eupelycosauria\|Sphenacomorpha]]) than to the caseids.<ref name="Spindler&al.2019"/> The latter are grouped with the [[Eothyrididae]] in the clade [[Caseasauria]], which represent the most [[Basal (phylogenetics)\|basal]] [[synapsid]]s.<ref name="Brocklehurst&al.2016">{{cite journal\|last1=Broklehurst \|first1=N. \|author2=Reisz, R. \|author3=Fernandez, V. \|author4=Fröbisch, J. \|year=2016 \|title=A Re-Description of 'Mycterosaurus' smithae, an Early Permian Eothyridid, and Its Impact on the Phylogeny of Pelycosaurian-Grade Synapsids \|journal=PLOS ONE \|volume=11 \|issue=6 \|pages=e0156810 \|doi=10.1371/journal.pone.0156810 \|pmid=27333277 \|pmc=4917111 \|bibcode=2016PLoSO..1156810B \|doi-access=free }}</ref>

	The first [[Phylogenetics\|phylogenetic]] analysis of caseids was published in 2008 by Hillary C. Maddin and colleagues. In this analysis, the genus ''[[Oromycter]]'' occupies the most basal position within the clade. ''[[Ennatosaurus]]'' is the [[sister group]] of a clade containing ''[[Cotylorhynchus\|Cotylorhynchus romeri]]'' and ''[[Angelosaurus\|Angelosaurus dolani]]''. This analysis reveals for the first time the [[paraphyly]] of the [[genus]] ''[[Casea]]'', the species ''“Casea” rutena'' representing a distinct genus which will be named in 2011 ''[[Euromycter]]''.<ref name="Maddin&al.2008"/><ref name=Reisz&al.2011>{{cite journal\|last1=Reisz \|first1=R.R. \|author2=Maddin, H.C. \|author3=Fröbisch, J. \|author4=Falconnet, J. \|year=2011 \|title=A new large caseid (Synapsida, Caseasauria) from the Permian of Rodez (France), including a reappraisal of ''"Casea" rutena'' Sigogneau-Russell & Russell, 1974 \|journal=Geodiversitas \|volume=33 \|issue=2 \|pages=227–246 \| doi= 10.5252/g2011n2a2\|s2cid=129458820 \|url=~~http~~://~~sciencepress~~.~~mnhn.fr~~/fr/~~periodiques~~/~~geodiversitas~~/~~33/2/un-nouveau-grand-caseide-synapsida-caseasauria-du-permien-du-bassin-de-rodez-france-et-reevaluation-de-casea-rutena-sigogneau-russell-russell-1974~~}}</ref>		The first [[Phylogenetics\|phylogenetic]] analysis of caseids was published in 2008 by Hillary C. Maddin and colleagues. In this analysis, the genus ''[[Oromycter]]'' occupies the most basal position within the clade. ''[[Ennatosaurus]]'' is the [[sister group]] of a clade containing ''[[Cotylorhynchus\|Cotylorhynchus romeri]]'' and ''[[Angelosaurus\|Angelosaurus dolani]]''. This analysis reveals for the first time the [[paraphyly]] of the [[genus]] ''[[Casea]]'', the species ''“Casea” rutena'' representing a distinct genus which will be named in 2011 ''[[Euromycter]]''.<ref name="Maddin&al.2008"/><ref name=Reisz&al.2011>{{cite journal\|last1=Reisz \|first1=R.R. \|author2=Maddin, H.C. \|author3=Fröbisch, J. \|author4=Falconnet, J. \|year=2011 \|title=A new large caseid (Synapsida, Caseasauria) from the Permian of Rodez (France), including a reappraisal of ''"Casea" rutena'' Sigogneau-Russell & Russell, 1974 \|journal=Geodiversitas \|volume=33 \|issue=2 \|pages=227–246 \| doi= 10.5252/g2011n2a2\|s2cid=129458820 \|url=https://zenodo.org/record/4597072/files/source.pdf}}</ref>

	Below is the first [[cladogram]] of caseids published by Maddin et al. in 2008.<ref name="Maddin&al.2008"/>		Below is the first [[cladogram]] of caseids published by Maddin et al. in 2008.<ref name="Maddin&al.2008"/>

AB-Babayo: Added free to read link in citations with OAbot #oabot

2023-12-28T13:15:37Z

Added free to read link in citations with OAbot #oabot

← Previous revision		Revision as of 13:15, 28 December 2023
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	==Footprints==		==Footprints==
	Many vertebrate tracks have been proposed as belonging to Caseidae. In the early 2000s, large footprints known as ''[[Brontopus\|Brontopus giganteus]]'' from the Permian [[Lodève]] basin in southern [[France]] were considered to belong either to a caseid or [[dinocephalia]]n [[therapsid]].<ref name="Gand&al.2000">{{cite journal\|last1=Gand \|first1=G. \|author2=Garric, J. \|author3=Demathieu, G. \|author4=Ellenberger, P. \|title=La palichnofaune de vertébrés tétrapodes du Permien supérieur du bassin de Lodève (Languedoc-France) \|journal=Palaevertebrata \|year=2000 \|volume=29 \|issue=1\|pages=1–82 \|doi= \|url=https://palaeovertebrata.com/issues/view/103}}</ref> In 2019, Lorenzo Marchetti and colleagues, however, determined that dinocephalians were most likely the trackmakers of the ichnogenus ''Brontopus''.<ref name="Marchetti&al.2019a">{{cite journal\|last1=Marchetti \|first1=L. \|author2=Klein, H. \|author3=Buchwitz, M. \|author4=Ronchi, A. \|author5=Smith, R.M.H. \|author6=De Klerk, W.J. \|author7=Sciscio, L. \|author8=Groenewald, G.H. \|title=Permian-Triassic vertebrate footprints from South Africa: Ichnotaxonomy, producers and biostratigraphy through two major faunal crises \|journal=Gondwana Research \|year=2019 \|volume=72 \|issue= \|pages=139–168 \|doi=10.1016/j.gr.2019.03.009\|bibcode=2019GondR..72..139M \|s2cid=133781923 \|doi-access=free }}</ref> In 2012, Rafael Costa da Silva and colleagues proposed that the ichnogenus ''[[Chelichnus]]'', widely distributed in Permian desert facies (fossil [[dune]]s) of [[Europe]], [[North America]] and [[South America]], could represent caseids footprints.<ref name="Silva&al.2012">{{cite journal\|last1=Silva \|first1=R.C. \|author2=Sedor, F.A. \|author3=Fernandes, A.C.S. \|title=Fossil footprints from the Late Permian of Brazil: An example of hidden biodiversity \|journal=Journal of South American Earth Sciences \|year=2012 \|volume=38 \|issue= \|pages=31–43 \|doi=10.1016/j.jsames.2012.05.001\|bibcode=2012JSAES..38...31D }}</ref> In 2019, Marchetti and colleagues, however, reinterpreted ''Chelichnus'' as a ''[[nomen dubium]]'' and a [[taphotaxon]], this type of track showing false anatomical features generated by locomotion on sandy inclined paleosurface.<ref name="Marchetti&al.2019b">{{cite journal\|last1=Marchetti \|first1=L. \|author2=Voigt, S. \|author3=Lucas, S.G. \|title=An anatomy-consistent study of the Lopingian eolian tracks of Germany and Scotland reveals the first evidence of the end-Guadalupian mass extinction at low paleolatitudes of Pangea \|journal=Gondwana Research \|year=2019 \|volume=73 \|issue= \|pages=32–53 \|doi=10.1016/j.gr.2019.03.013\|bibcode=2019GondR..73...32M \|s2cid=146780505 }}</ref> In 2014, Eva Sacchi and colleagues described the ichnotaxon ''[[Dimetropus\|Dimetropus osageorum]]'' from about a hundred isolated footprints and several trackways from the early Permian [[Wellington Formation]] in [[Oklahoma]]. The morphological study of these footprints and their comparison with tetrapod skeletons revealed that they probably belong to a large caseid comparable in size to ''[[Cotylorhynchus\|Cotylorhynchus romeri]]'' (the latter from more recent strata).<ref name="Sacchi&al.2014">{{cite journal\|last1=Sacchi \|first1=E. \|author2=Cifelli, R. \|author3=Citton, P. \|author4=Nicosia, U. \|author5=Romano, M. \|title=''Dimetropus osageorum'' n. isp. from the Early Permian of Oklahoma (USA): A Trace and its Trackmaker \|journal=Ichnos: An International Journal for Plant and Animal Traces \|year=2014 \|volume=21 \|issue=3 \|pages=175–192 \|doi=10.1080/10420940.2014.933070\|s2cid=129567990 }}</ref> According to Sacchi and colleagues, some footprints from the Lodève Basin (unspecified geological formation) are similar to those from the Wellington Formation and these authors designate them as ''Dimetropus'' cf. ''osageorum''.<ref name="Sacchi&al.2014"/> The study by Sacchi et al. also demonstrates that the ichnogenus ''Dimetropus'' exhibits great morphological variation and that its producers can be assigned to different zoological groups among non-therapsid synapsids and not just [[Sphenacodontidae]] as once thought.<ref name="Sacchi&al.2014"/> In 2021, Rafel Matamales-Andreu and colleagues assigned footprints found in the Lower Permian ([[Artinskian]]-[[Kungurian]]) Port des Canonge Formation in [[Mallorca]] to a caseid. These footprints resemble those of ''D. osageorum'' but they also have differences. These footprints are left in open nomenclature as cf. ''Dimetropus'' sp.. They are, however, smaller than those of ''D. osageorum'' and would have been produced by a caseid of modest size with proportions comparable to ''[[Ennatosaurus]]''.<ref name="Matamales-Andreu&al.2021">{{cite journal\|last1=Matamales-Andreu \|first1=R. \|author2=Mujal, E. \|author3=Galobart, À. \|author4=Fortuny, J. \|title=Insight on the evolution of synapsid locomotion based on tetrapod tracks from the lower Permian of Mallorca (Balearic Islands, western Mediterranean) \|journal=Palaeogeography, Palaeoclimatology, Palaeoecology \|year=2021 \|volume=579 \|issue= \|pages=110589 \|doi=10.1016/j.palaeo.2021.110589\|bibcode=2021PPP...579k0589M }}</ref>		Many vertebrate tracks have been proposed as belonging to Caseidae. In the early 2000s, large footprints known as ''[[Brontopus\|Brontopus giganteus]]'' from the Permian [[Lodève]] basin in southern [[France]] were considered to belong either to a caseid or [[dinocephalia]]n [[therapsid]].<ref name="Gand&al.2000">{{cite journal\|last1=Gand \|first1=G. \|author2=Garric, J. \|author3=Demathieu, G. \|author4=Ellenberger, P. \|title=La palichnofaune de vertébrés tétrapodes du Permien supérieur du bassin de Lodève (Languedoc-France) \|journal=Palaevertebrata \|year=2000 \|volume=29 \|issue=1\|pages=1–82 \|doi= \|url=https://palaeovertebrata.com/issues/view/103}}</ref> In 2019, Lorenzo Marchetti and colleagues, however, determined that dinocephalians were most likely the trackmakers of the ichnogenus ''Brontopus''.<ref name="Marchetti&al.2019a">{{cite journal\|last1=Marchetti \|first1=L. \|author2=Klein, H. \|author3=Buchwitz, M. \|author4=Ronchi, A. \|author5=Smith, R.M.H. \|author6=De Klerk, W.J. \|author7=Sciscio, L. \|author8=Groenewald, G.H. \|title=Permian-Triassic vertebrate footprints from South Africa: Ichnotaxonomy, producers and biostratigraphy through two major faunal crises \|journal=Gondwana Research \|year=2019 \|volume=72 \|issue= \|pages=139–168 \|doi=10.1016/j.gr.2019.03.009\|bibcode=2019GondR..72..139M \|s2cid=133781923 \|doi-access=free }}</ref> In 2012, Rafael Costa da Silva and colleagues proposed that the ichnogenus ''[[Chelichnus]]'', widely distributed in Permian desert facies (fossil [[dune]]s) of [[Europe]], [[North America]] and [[South America]], could represent caseids footprints.<ref name="Silva&al.2012">{{cite journal\|last1=Silva \|first1=R.C. \|author2=Sedor, F.A. \|author3=Fernandes, A.C.S. \|title=Fossil footprints from the Late Permian of Brazil: An example of hidden biodiversity \|journal=Journal of South American Earth Sciences \|year=2012 \|volume=38 \|issue= \|pages=31–43 \|doi=10.1016/j.jsames.2012.05.001\|bibcode=2012JSAES..38...31D }}</ref> In 2019, Marchetti and colleagues, however, reinterpreted ''Chelichnus'' as a ''[[nomen dubium]]'' and a [[taphotaxon]], this type of track showing false anatomical features generated by locomotion on sandy inclined paleosurface.<ref name="Marchetti&al.2019b">{{cite journal\|last1=Marchetti \|first1=L. \|author2=Voigt, S. \|author3=Lucas, S.G. \|title=An anatomy-consistent study of the Lopingian eolian tracks of Germany and Scotland reveals the first evidence of the end-Guadalupian mass extinction at low paleolatitudes of Pangea \|journal=Gondwana Research \|year=2019 \|volume=73 \|issue= \|pages=32–53 \|doi=10.1016/j.gr.2019.03.013\|bibcode=2019GondR..73...32M \|s2cid=146780505 }}</ref> In 2014, Eva Sacchi and colleagues described the ichnotaxon ''[[Dimetropus\|Dimetropus osageorum]]'' from about a hundred isolated footprints and several trackways from the early Permian [[Wellington Formation]] in [[Oklahoma]]. The morphological study of these footprints and their comparison with tetrapod skeletons revealed that they probably belong to a large caseid comparable in size to ''[[Cotylorhynchus\|Cotylorhynchus romeri]]'' (the latter from more recent strata).<ref name="Sacchi&al.2014">{{cite journal\|last1=Sacchi \|first1=E. \|author2=Cifelli, R. \|author3=Citton, P. \|author4=Nicosia, U. \|author5=Romano, M. \|title=''Dimetropus osageorum'' n. isp. from the Early Permian of Oklahoma (USA): A Trace and its Trackmaker \|journal=Ichnos: An International Journal for Plant and Animal Traces \|year=2014 \|volume=21 \|issue=3 \|pages=175–192 \|doi=10.1080/10420940.2014.933070\|s2cid=129567990 }}</ref> According to Sacchi and colleagues, some footprints from the Lodève Basin (unspecified geological formation) are similar to those from the Wellington Formation and these authors designate them as ''Dimetropus'' cf. ''osageorum''.<ref name="Sacchi&al.2014"/> The study by Sacchi et al. also demonstrates that the ichnogenus ''Dimetropus'' exhibits great morphological variation and that its producers can be assigned to different zoological groups among non-therapsid synapsids and not just [[Sphenacodontidae]] as once thought.<ref name="Sacchi&al.2014"/> In 2021, Rafel Matamales-Andreu and colleagues assigned footprints found in the Lower Permian ([[Artinskian]]-[[Kungurian]]) Port des Canonge Formation in [[Mallorca]] to a caseid. These footprints resemble those of ''D. osageorum'' but they also have differences. These footprints are left in open nomenclature as cf. ''Dimetropus'' sp.. They are, however, smaller than those of ''D. osageorum'' and would have been produced by a caseid of modest size with proportions comparable to ''[[Ennatosaurus]]''.<ref name="Matamales-Andreu&al.2021">{{cite journal\|last1=Matamales-Andreu \|first1=R. \|author2=Mujal, E. \|author3=Galobart, À. \|author4=Fortuny, J. \|title=Insight on the evolution of synapsid locomotion based on tetrapod tracks from the lower Permian of Mallorca (Balearic Islands, western Mediterranean) \|journal=Palaeogeography, Palaeoclimatology, Palaeoecology \|year=2021 \|volume=579 \|issue= \|pages=110589 \|doi=10.1016/j.palaeo.2021.110589\|bibcode=2021PPP...579k0589M \|url=https://ddd.uab.cat/pub/artpub/2021/248974/palpalpal_a2021v579a110589.pdf }}</ref>

	==Paleobiology==		==Paleobiology==

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2023-12-03T22:10:41Z

Add: doi-access, authors 1-1. Removed proxy/dead URL that duplicated identifier. Removed parameters. Some additions/deletions were parameter name changes. | Use this bot. Report bugs. | #UCB_CommandLine

← Previous revision		Revision as of 22:10, 3 December 2023
Line 41:		Line 41:
	\|footer=''Cotylorhynchus romeri''. Top : skull reconstruction in left lateral view, and medial and lateral views of mandible; bottom : skull reconstruction in dorsal, ventral, posterior and anterior views.}}[[File:Euromycter rutenus 6778 retouched.jpg\|thumb\|right\|Skull of ''[[Euromycter\|Euromycter rutenus]]'' in ventral view allowing to see the numerous small teeth adorning the bones of the palate (the [[parasphenoid]], triangular in shape in the center, and the very elongated [[Pterygoid bone\|pterygoids]] on each side). The hyoid apparatus is not present here because it was removed during the preparation of the palate.]]		\|footer=''Cotylorhynchus romeri''. Top : skull reconstruction in left lateral view, and medial and lateral views of mandible; bottom : skull reconstruction in dorsal, ventral, posterior and anterior views.}}[[File:Euromycter rutenus 6778 retouched.jpg\|thumb\|right\|Skull of ''[[Euromycter\|Euromycter rutenus]]'' in ventral view allowing to see the numerous small teeth adorning the bones of the palate (the [[parasphenoid]], triangular in shape in the center, and the very elongated [[Pterygoid bone\|pterygoids]] on each side). The hyoid apparatus is not present here because it was removed during the preparation of the palate.]]

	Caseids measured from less than {{convert\|1\|m}} to {{convert\|7\|m}} in length.<ref name="Romano2017">{{cite journal\|last1=Romano \|first1=M. \|year=2017 \|title=Long bone scaling of caseid synapsids: a combined morphometric and cladistic approach \|journal=Lethaia \|volume=50 \|issue=4 \|pages=511–526 \|doi=10.1111/let.12207}}</ref><ref name="Angielczyk&Kammerer2018">{{cite book\|last=Angielczyk\|first=K.D. \|author2=Kammerer, C.F. \|year=2018 \|chapter=Non-Mammalian synapsids : the deep roots of the mammalian family tree \|pages=117–198 \|title=Handbook of Zoology : Mammalian Evolution, Diversity and Systematics \|editor1-last=Zachos\|editor1-first=F.E. \|editor2=Asher, R.J. \|publisher=de Gruyter \|location=Berlin \|isbn=978-3-11-027590-2}}</ref><ref name="Romano&al.2018">{{cite journal\|last1=Romano\|first1=M. \|author2=Citton, P. \|author3=Maganuco, S. \|author4=Sacchi, E. \|author5=Caratelli, M. \|author6=Ronchi, A. \|author7=Nicosia, U. \|year=2018 \|title=New basal synapsid discovery at the Permian outcrop of Torre del Porticciolo (Alghero, Italy) \|journal=Geological Journal \|volume=54 \|issue=3 \|pages=1554–1566 \|doi=10.1002/gj.3250\|s2cid=133755506 }}</ref> They had a small head wider than high and with a forward-inclined snout, a very short neck, a long tail, robust forelimbs, and a body of variable proportions depending on their diet. Small insectivorous species like ''[[Eocasea]]'' had an unexpanded trunk.<ref name="Reisz&Fröbisch2014">{{cite journal\|last1=Reisz\|first1=R.R. \|author2=Fröbisch, J. \|year=2014 \|title=Oldest caseid synapsid from the late Pennsylvanian of Kansas, and the evolution of herbivory in terrestrial vertebrates \|journal=PLOS ONE \|volume=9 \|issue=4 \|pages=e94518 \|doi=10.1371/journal.pone.0094518 \|pmid=24739998 \|pmc=3989228\|bibcode=2014PLoSO...994518R \|doi-access=free }}</ref> Others with an omnivorous diet like ''[[Martensius]]'' had a barely enlarged rib cage, a more elongated skull, smaller [[nostril]]s, and a snout less inclined forward than in herbivorous caseids.<ref name="Berman&al.2020">{{cite journal\|last1=Berman \|first1=D.S. \|author2=Maddin, H.C. \|author3=Henrici, A.C. \|author4=Sumida, S.S. \|author5=Scott, D. \|author6=Reisz, R.R. \|title=New primitive Caseid (Synapsida, Caseasauria) from the Early Permian of Germany \|journal=Annals of Carnegie Museum \|year=2020 \|volume=86 \|issue=1 \|pages=43–75 \|doi=10.2992/007.086.0103\|s2cid=216027787 }}</ref> The latter were characterized by their disproportionately small [[skull]] compared to the size of the body. The postcranial skeleton indeed shows a spectacular increase in the volume of the [[rib cage]], which becomes very wide and barrel-shaped, probably to accommodate a particularly developed [[Gastrointestinal tract#Lower gastrointestinal tract\|intestine]], necessary for the digestion of high-fiber rich plants. In these forms, the skull has very large external nostrils and a very short facial region with a strong forward inclination of the end of the snout which clearly overhangs the dental row. The [[Skull#Fenestrae\|temporal fenestrae]] are also relatively large (especially in ''[[Ennatosaurus]]''), the [[Supratemporal bone\|supratemporals]] are large in size, and, on the [[Occipital bone\|occipital]] surface, the paroccipital [[Process (anatomy)\|processes]] are massively developed, establishing strong supporting contacts with the [[Squamosal bone\|squamosals]].<ref name="Kemp1982">{{cite book\|last=Kemp \|first=T.S. \|year=1982 \|chapter=Pelycosaurs \|pages=39 \|title=Mammal-like reptiles and the origin of Mammals \|editor1-last=Kemp\|editor1-first=T.S. \|publisher=Academic Press \|location=London \|isbn=978-0124041202}}</ref> The dorsal surface of the skull is covered with numerous small pits. These suggest the presence of large [[Reptile scale\|scales]] on the head of these animals.<ref name="Romano&al.2017a">{{cite journal\|last1=Romano \|first1=M. \|author2=Brocklehurst, N. \|author3=Fröbisch, J. \|year=2017 \|title=The postcranial skeleton of ''Ennatosaurus tecton'' (Synapsida, Caseidae) \|journal=Journal of Systematic Palaeontology \|volume=16 \|issue= 13\|pages=1097–1122 \|doi=10.1080/14772019.2017.1367729\|s2cid=89922565 }}</ref> Numerous [[Lip\|labial]] [[Foramen\|foramina]] running parallel to the ventral edge of the [[premaxilla]] and [[maxilla]], as well as along the dorsal edge of the [[Mandible#Other vertebrates\|dentary]], suggest the presence of scaly "lips" which must have concealed the dentition when the jaws were closed.<ref name="Reisz&al.2022">{{cite journal\|last1=Reisz \|first1=R.R. \|author2=Scott, D. \|author3=Modesto, S.P. \|year=2022 \|title=Cranial Anatomy of the Caseid Synapsid ''Cotylorhynchus romeri'', a Large Terrestrial Herbivore From the Lower Permian of Oklahoma, U.S.A \|journal=Frontiers in Earth Science \|volume=10 \|issue= \|pages=1–19 \|doi=10.3389/feart.2022.847560\|doi-access=free }}</ref><ref name="Romano&al.2017a"/> The [[Tooth\|teeth]], simply conical and pointed in insectivorous species, adopt in herbivorous species a leafy or spatulate morphology and are provided with more or less numerous cuspules. Numerous small teeth also adorned several bones of the palate. Herbivorous species do not show a simple evolutionary trend towards increasing tooth complexity.<ref name="Maddin&al.2008">{{cite journal \|last1=Maddin \|first1=H.C. \|author2=Sidor, C.A. \|author3=Reisz, R.R. \|year=2008 \|title=Cranial anatomy of ''Ennatosaurus tecton'' (Synapsida: Caseidae) from the Middle Permian of Russia and the evolutionary relationships of Caseidae \|journal=Journal of Vertebrate Paleontology \|volume=28 \|issue=1 \|pages=160–180 \|doi=10.1671/0272-4634(2008)28[160:CAOETS]2.0.CO;2\|s2cid=44064927 }}</ref> Thus, the teeth of the basal taxa ''[[Casea]]'' and ''[[Arisierpeton]]'' have three cuspules<ref name="Olson1968">{{cite journal\|last1=Olson \|~~first~~=E.C. \|year=1968 \|title=The family Caseidae \|journal=Fieldiana: Geology \|volume=17 \|issue= \|pages=225–349\| doi= }}</ref><ref name="Reisz2019">{{cite journal \|last1=Reisz \|first1=R. \|year=2019 \|title=A small caseid synapsid, ''Arisierpeton simplex'' gen. et sp. nov., from the early Permian of Oklahoma, with a discussion of synapsid diversity at the classic Richards Spur locality \|journal=PeerJ \|volume=7e6615 \|issue= \|pages=1–20 \|doi=10.7717/peerj.6615 \|pmid=30997285 \|pmc=6462398 \|~~url~~=~~http://doi.org/10.7717/peerj.6615~~}}</ref> just like in the more derived forms ''[[Cotylorhynchus]]'' and ''[[Caseopsis]]''.<ref name="Olson1968"/> ''Ennatosaurus'' and ''[[Euromycter]]'', which occupy an intermediate [[Phylogenetics\|phylogenetic]] position, have teeth bearing 5 to 7 cuspules and 5 to 8 cuspules respectively.<ref name="Maddin&al.2008"/><ref name="sigogneau-russell1974">{{cite journal \|last1=Sigoneau-Russell \|first1=D. \|author2=Russel, D.E. \|year=1974 \|title=Étude du premier caséidé (Reptilia, Pelycosauria) d'Europe occidentale \|journal=Bulletin du Muséum National d'Histoire Naturelle \|series=Série 3 \|volume=38 \|issue=230 \|pages=145–215 \|doi= \|url=http://bibliotheques.mnhn.fr/EXPLOITATION/infodoc/digitalCollections/viewerpopup.aspx?seid=BMSCT_S003_1974_T230_N038}}</ref> ''[[Angelosaurus]]'', one of the most derived caseids, has teeth with 5 cuspules.<ref name="Olson1968"/> In ''Angelosaurus'' the teeth have a bulbous morphology with very short and wide crowns. Their sturdiness and the significant wear they show, indicate that ''Angelosaurus'' must have fed on tougher plants than those on which most other herbivorous caseids fed.<ref name="Olson1968"/> Herbivorous caseids also show very different dietary adaptations from those seen in another group of basal synapsids, the [[Edaphosauridae]]. The latter had, in addition to the marginal dentition, a dental battery made up of numerous teeth located both on the [[palate]] and on the inner surface of the lower jaws. In herbivorous caseids, the palatal teeth are smaller, and the inner surface of the lower jaws bears no teeth. Instead of a dental battery, they had a massive [[tongue]] (as indicated by the presence of a highly developed [[hyoid apparatus]] found in ''Ennatosaurus'' and ''Euromycter'') perhaps rough, with which they had to compress food against the palatal teeth.<ref name="Olson1968"/><ref name="Reisz&al.2022"/>		Caseids measured from less than {{convert\|1\|m}} to {{convert\|7\|m}} in length.<ref name="Romano2017">{{cite journal\|last1=Romano \|first1=M. \|year=2017 \|title=Long bone scaling of caseid synapsids: a combined morphometric and cladistic approach \|journal=Lethaia \|volume=50 \|issue=4 \|pages=511–526 \|doi=10.1111/let.12207}}</ref><ref name="Angielczyk&Kammerer2018">{{cite book\|last=Angielczyk\|first=K.D. \|author2=Kammerer, C.F. \|year=2018 \|chapter=Non-Mammalian synapsids : the deep roots of the mammalian family tree \|pages=117–198 \|title=Handbook of Zoology : Mammalian Evolution, Diversity and Systematics \|editor1-last=Zachos\|editor1-first=F.E. \|editor2=Asher, R.J. \|publisher=de Gruyter \|location=Berlin \|isbn=978-3-11-027590-2}}</ref><ref name="Romano&al.2018">{{cite journal\|last1=Romano\|first1=M. \|author2=Citton, P. \|author3=Maganuco, S. \|author4=Sacchi, E. \|author5=Caratelli, M. \|author6=Ronchi, A. \|author7=Nicosia, U. \|year=2018 \|title=New basal synapsid discovery at the Permian outcrop of Torre del Porticciolo (Alghero, Italy) \|journal=Geological Journal \|volume=54 \|issue=3 \|pages=1554–1566 \|doi=10.1002/gj.3250\|s2cid=133755506 }}</ref> They had a small head wider than high and with a forward-inclined snout, a very short neck, a long tail, robust forelimbs, and a body of variable proportions depending on their diet. Small insectivorous species like ''[[Eocasea]]'' had an unexpanded trunk.<ref name="Reisz&Fröbisch2014">{{cite journal\|last1=Reisz\|first1=R.R. \|author2=Fröbisch, J. \|year=2014 \|title=Oldest caseid synapsid from the late Pennsylvanian of Kansas, and the evolution of herbivory in terrestrial vertebrates \|journal=PLOS ONE \|volume=9 \|issue=4 \|pages=e94518 \|doi=10.1371/journal.pone.0094518 \|pmid=24739998 \|pmc=3989228\|bibcode=2014PLoSO...994518R \|doi-access=free }}</ref> Others with an omnivorous diet like ''[[Martensius]]'' had a barely enlarged rib cage, a more elongated skull, smaller [[nostril]]s, and a snout less inclined forward than in herbivorous caseids.<ref name="Berman&al.2020">{{cite journal\|last1=Berman \|first1=D.S. \|author2=Maddin, H.C. \|author3=Henrici, A.C. \|author4=Sumida, S.S. \|author5=Scott, D. \|author6=Reisz, R.R. \|title=New primitive Caseid (Synapsida, Caseasauria) from the Early Permian of Germany \|journal=Annals of Carnegie Museum \|year=2020 \|volume=86 \|issue=1 \|pages=43–75 \|doi=10.2992/007.086.0103\|s2cid=216027787 }}</ref> The latter were characterized by their disproportionately small [[skull]] compared to the size of the body. The postcranial skeleton indeed shows a spectacular increase in the volume of the [[rib cage]], which becomes very wide and barrel-shaped, probably to accommodate a particularly developed [[Gastrointestinal tract#Lower gastrointestinal tract\|intestine]], necessary for the digestion of high-fiber rich plants. In these forms, the skull has very large external nostrils and a very short facial region with a strong forward inclination of the end of the snout which clearly overhangs the dental row. The [[Skull#Fenestrae\|temporal fenestrae]] are also relatively large (especially in ''[[Ennatosaurus]]''), the [[Supratemporal bone\|supratemporals]] are large in size, and, on the [[Occipital bone\|occipital]] surface, the paroccipital [[Process (anatomy)\|processes]] are massively developed, establishing strong supporting contacts with the [[Squamosal bone\|squamosals]].<ref name="Kemp1982">{{cite book\|last=Kemp \|first=T.S. \|year=1982 \|chapter=Pelycosaurs \|pages=39 \|title=Mammal-like reptiles and the origin of Mammals \|editor1-last=Kemp\|editor1-first=T.S. \|publisher=Academic Press \|location=London \|isbn=978-0124041202}}</ref> The dorsal surface of the skull is covered with numerous small pits. These suggest the presence of large [[Reptile scale\|scales]] on the head of these animals.<ref name="Romano&al.2017a">{{cite journal\|last1=Romano \|first1=M. \|author2=Brocklehurst, N. \|author3=Fröbisch, J. \|year=2017 \|title=The postcranial skeleton of ''Ennatosaurus tecton'' (Synapsida, Caseidae) \|journal=Journal of Systematic Palaeontology \|volume=16 \|issue= 13\|pages=1097–1122 \|doi=10.1080/14772019.2017.1367729\|s2cid=89922565 }}</ref> Numerous [[Lip\|labial]] [[Foramen\|foramina]] running parallel to the ventral edge of the [[premaxilla]] and [[maxilla]], as well as along the dorsal edge of the [[Mandible#Other vertebrates\|dentary]], suggest the presence of scaly "lips" which must have concealed the dentition when the jaws were closed.<ref name="Reisz&al.2022">{{cite journal\|last1=Reisz \|first1=R.R. \|author2=Scott, D. \|author3=Modesto, S.P. \|year=2022 \|title=Cranial Anatomy of the Caseid Synapsid ''Cotylorhynchus romeri'', a Large Terrestrial Herbivore From the Lower Permian of Oklahoma, U.S.A \|journal=Frontiers in Earth Science \|volume=10 \|issue= \|pages=1–19 \|doi=10.3389/feart.2022.847560\|doi-access=free }}</ref><ref name="Romano&al.2017a"/> The [[Tooth\|teeth]], simply conical and pointed in insectivorous species, adopt in herbivorous species a leafy or spatulate morphology and are provided with more or less numerous cuspules. Numerous small teeth also adorned several bones of the palate. Herbivorous species do not show a simple evolutionary trend towards increasing tooth complexity.<ref name="Maddin&al.2008">{{cite journal \|last1=Maddin \|first1=H.C. \|author2=Sidor, C.A. \|author3=Reisz, R.R. \|year=2008 \|title=Cranial anatomy of ''Ennatosaurus tecton'' (Synapsida: Caseidae) from the Middle Permian of Russia and the evolutionary relationships of Caseidae \|journal=Journal of Vertebrate Paleontology \|volume=28 \|issue=1 \|pages=160–180 \|doi=10.1671/0272-4634(2008)28[160:CAOETS]2.0.CO;2\|s2cid=44064927 }}</ref> Thus, the teeth of the basal taxa ''[[Casea]]'' and ''[[Arisierpeton]]'' have three cuspules<ref name="Olson1968">{{cite journal\|last1=Olson \|first1=E.C. \|year=1968 \|title=The family Caseidae \|journal=Fieldiana: Geology \|volume=17 \|issue= \|pages=225–349\| doi= }}</ref><ref name="Reisz2019">{{cite journal \|last1=Reisz \|first1=R. \|year=2019 \|title=A small caseid synapsid, ''Arisierpeton simplex'' gen. et sp. nov., from the early Permian of Oklahoma, with a discussion of synapsid diversity at the classic Richards Spur locality \|journal=PeerJ \|volume=7e6615 \|issue= \|pages=1–20 \|doi=10.7717/peerj.6615 \|pmid=30997285 \|pmc=6462398 \|doi-access=free }}</ref> just like in the more derived forms ''[[Cotylorhynchus]]'' and ''[[Caseopsis]]''.<ref name="Olson1968"/> ''Ennatosaurus'' and ''[[Euromycter]]'', which occupy an intermediate [[Phylogenetics\|phylogenetic]] position, have teeth bearing 5 to 7 cuspules and 5 to 8 cuspules respectively.<ref name="Maddin&al.2008"/><ref name="sigogneau-russell1974">{{cite journal \|last1=Sigoneau-Russell \|first1=D. \|author2=Russel, D.E. \|year=1974 \|title=Étude du premier caséidé (Reptilia, Pelycosauria) d'Europe occidentale \|journal=Bulletin du Muséum National d'Histoire Naturelle \|series=Série 3 \|volume=38 \|issue=230 \|pages=145–215 \|doi= \|url=http://bibliotheques.mnhn.fr/EXPLOITATION/infodoc/digitalCollections/viewerpopup.aspx?seid=BMSCT_S003_1974_T230_N038}}</ref> ''[[Angelosaurus]]'', one of the most derived caseids, has teeth with 5 cuspules.<ref name="Olson1968"/> In ''Angelosaurus'' the teeth have a bulbous morphology with very short and wide crowns. Their sturdiness and the significant wear they show, indicate that ''Angelosaurus'' must have fed on tougher plants than those on which most other herbivorous caseids fed.<ref name="Olson1968"/> Herbivorous caseids also show very different dietary adaptations from those seen in another group of basal synapsids, the [[Edaphosauridae]]. The latter had, in addition to the marginal dentition, a dental battery made up of numerous teeth located both on the [[palate]] and on the inner surface of the lower jaws. In herbivorous caseids, the palatal teeth are smaller, and the inner surface of the lower jaws bears no teeth. Instead of a dental battery, they had a massive [[tongue]] (as indicated by the presence of a highly developed [[hyoid apparatus]] found in ''Ennatosaurus'' and ''Euromycter'') perhaps rough, with which they had to compress food against the palatal teeth.<ref name="Olson1968"/><ref name="Reisz&al.2022"/>

	[[File:Arisierpeton simplex, premaxillae.png\|thumb\|right\|Premaxillae of ''[[Arisierpeton\|Arisierpeton simplex]]''. (A) Specimen GAA 00242 in anterior, labial, posterior, ventral, and dorsal views; (B) Specimen GAA 00239 in labial (showing a complete tooth with tricuspid distal end), partial labial, posterior and ventral views.]][[File:Cotylorhynchus romeria from Norman, Oklahoma.jpg\|thumb\|left\|Skeleton of ''Cotylorhynchus romeri'' on display at the [[Sam Noble Oklahoma Museum of Natural History]].]]		[[File:Arisierpeton simplex, premaxillae.png\|thumb\|right\|Premaxillae of ''[[Arisierpeton\|Arisierpeton simplex]]''. (A) Specimen GAA 00242 in anterior, labial, posterior, ventral, and dorsal views; (B) Specimen GAA 00239 in labial (showing a complete tooth with tricuspid distal end), partial labial, posterior and ventral views.]][[File:Cotylorhynchus romeria from Norman, Oklahoma.jpg\|thumb\|left\|Skeleton of ''Cotylorhynchus romeri'' on display at the [[Sam Noble Oklahoma Museum of Natural History]].]]
Line 66:		Line 66:
	This hypothesis is however disputed by Kenneth Angielczyk and Christian Kammerer as well as by Robert Reisz and colleagues based on [[Paleontology\|paleontological]] and [[Taphonomy\|taphonomic]] data combined with the absence in these large caseids of morphological adaptations to an aquatic lifestyle. According to Angielczyk and Kammerer, the low bone density of caseids identified by Lambertz et al. does not resemble that of semiaquatic animals, which tend to have a more strongly ossified skeleton to provide passive [[buoyancy]] control and increased stability against current and wave action. ''Cotylorhynchus'' bone microstructure is more similar to what is seen in animals living in the [[Pelagic zone\|open ocean]], such as [[cetacea]]ns and [[pinniped]]s, which emphasize high maneuverability, rapid acceleration and hydrodynamic control of buoyancy. However, the caseid morphology was totally incompatible with a [[Pelagic zone\|pelagic]] lifestyle. Thus, due to these unusual data, Angielczyk and Kammerer consider that the available evidence is still insufficient to question the more widely assumed terrestrial lifestyle of caseids.<ref name="Angielczyk&Kammerer2018"/> Robert Reisz and colleagues also dispute the supposed semiaquatic lifestyle of the caseids on the fact that the latter possess no morphological adaptations to an aquatic lifestyle and, in the case of the species ''Cotylorhynchus romeri'', on the interpretation that this animal lived in a dry environment for part of the year as indicated by the presence of numerous skeletons of the amphibian ''[[Brachydectes]]'' preserved in [[aestivation]] and of the [[lungfish]] ''[[Gnathorhiza]]'', another well-known aestivator.<ref name="Reisz&al.2022"/>		This hypothesis is however disputed by Kenneth Angielczyk and Christian Kammerer as well as by Robert Reisz and colleagues based on [[Paleontology\|paleontological]] and [[Taphonomy\|taphonomic]] data combined with the absence in these large caseids of morphological adaptations to an aquatic lifestyle. According to Angielczyk and Kammerer, the low bone density of caseids identified by Lambertz et al. does not resemble that of semiaquatic animals, which tend to have a more strongly ossified skeleton to provide passive [[buoyancy]] control and increased stability against current and wave action. ''Cotylorhynchus'' bone microstructure is more similar to what is seen in animals living in the [[Pelagic zone\|open ocean]], such as [[cetacea]]ns and [[pinniped]]s, which emphasize high maneuverability, rapid acceleration and hydrodynamic control of buoyancy. However, the caseid morphology was totally incompatible with a [[Pelagic zone\|pelagic]] lifestyle. Thus, due to these unusual data, Angielczyk and Kammerer consider that the available evidence is still insufficient to question the more widely assumed terrestrial lifestyle of caseids.<ref name="Angielczyk&Kammerer2018"/> Robert Reisz and colleagues also dispute the supposed semiaquatic lifestyle of the caseids on the fact that the latter possess no morphological adaptations to an aquatic lifestyle and, in the case of the species ''Cotylorhynchus romeri'', on the interpretation that this animal lived in a dry environment for part of the year as indicated by the presence of numerous skeletons of the amphibian ''[[Brachydectes]]'' preserved in [[aestivation]] and of the [[lungfish]] ''[[Gnathorhiza]]'', another well-known aestivator.<ref name="Reisz&al.2022"/>

	In 2022, Werneburg and colleagues proposed a somewhat different semiaquatic lifestyle, in which large caseids like ''[[Lalieudorhynchus]]'' (whose bone texture is even more osteoporotic than in ''Cotylorhynchus'') would be ecological equivalents of modern [[Hippopotamus\|hippos]], passing part of their time in the water (being underwater walkers rather than swimming animals) but coming on dry land for food.<ref name="Werneburg&al.2022">{{cite journal\|last1=Werneburg \|~~first~~=R. \|author2=Spindler, F. \|author3=Falconnet, J. \|author4=Steyer, J.-S. \|author5=Vianey-Liaud, M. \|author6=Schneider, J.W. \|title=A new caseid synapsid from the Permian (Guadalupian) of the Lodève basin (Occitanie, France) \|journal=Palaeovertebrata \|year=2022 \|volume=45(2)-e2 \|issue= 2\|pages=e2 \|doi=10.18563/pv.45.2.e2\|s2cid=253542331 \|url=https://hal.archives-ouvertes.fr/hal-03851911/file/Werneburg%20et%20al%202022%20A%20new%20caseid%20synapsid%20from%20the%20Permian%20of%20the%20Lod%C3%A8ve%20Basin.pdf }}</ref>		In 2022, Werneburg and colleagues proposed a somewhat different semiaquatic lifestyle, in which large caseids like ''[[Lalieudorhynchus]]'' (whose bone texture is even more osteoporotic than in ''Cotylorhynchus'') would be ecological equivalents of modern [[Hippopotamus\|hippos]], passing part of their time in the water (being underwater walkers rather than swimming animals) but coming on dry land for food.<ref name="Werneburg&al.2022">{{cite journal\|last1=Werneburg \|first1=R. \|author2=Spindler, F. \|author3=Falconnet, J. \|author4=Steyer, J.-S. \|author5=Vianey-Liaud, M. \|author6=Schneider, J.W. \|title=A new caseid synapsid from the Permian (Guadalupian) of the Lodève basin (Occitanie, France) \|journal=Palaeovertebrata \|year=2022 \|volume=45(2)-e2 \|issue= 2\|pages=e2 \|doi=10.18563/pv.45.2.e2\|s2cid=253542331 \|url=https://hal.archives-ouvertes.fr/hal-03851911/file/Werneburg%20et%20al%202022%20A%20new%20caseid%20synapsid%20from%20the%20Permian%20of%20the%20Lod%C3%A8ve%20Basin.pdf }}</ref>

	==Evolution==		==Evolution==

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2023-08-12T09:35:49Z

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← Previous revision		Revision as of 09:35, 12 August 2023
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	[[File:Arisierpeton simplex, premaxillae.png\|thumb\|right\|Premaxillae of ''[[Arisierpeton\|Arisierpeton simplex]]''. (A) Specimen GAA 00242 in anterior, labial, posterior, ventral, and dorsal views; (B) Specimen GAA 00239 in labial (showing a complete tooth with tricuspid distal end), partial labial, posterior and ventral views.]][[File:Cotylorhynchus romeria from Norman, Oklahoma.jpg\|thumb\|left\|Skeleton of ''Cotylorhynchus romeri'' on display at the [[Sam Noble Oklahoma Museum of Natural History]].]]		[[File:Arisierpeton simplex, premaxillae.png\|thumb\|right\|Premaxillae of ''[[Arisierpeton\|Arisierpeton simplex]]''. (A) Specimen GAA 00242 in anterior, labial, posterior, ventral, and dorsal views; (B) Specimen GAA 00239 in labial (showing a complete tooth with tricuspid distal end), partial labial, posterior and ventral views.]][[File:Cotylorhynchus romeria from Norman, Oklahoma.jpg\|thumb\|left\|Skeleton of ''Cotylorhynchus romeri'' on display at the [[Sam Noble Oklahoma Museum of Natural History]].]]

	The forelimbs of caseids are often more robust than the hindlimbs. It has indeed been observed that the bones of the forelimbs gained in robustness from the beginning of the evolution of the group, before the appearance of large species, while the hindlimbs remained slenderer. These characteristics suggest that the initial strengthening of the forelimbs was probably related to a particular function such as digging, and that this trait was later [[Exaptation\|exapted]] by more derived and larger caseids to support their weights of up to several hundred kilograms.<ref name="Romano2017"/> During their evolutionary history, caseids also show a reduction in their phalangeal formula.{{refn\|group=nb\|The phalangeal formula corresponds to the number of phalanges constituting each digits of the manus and pes of [[Tetrapoda\|tetrapods]]. It is listed starting from digit I (corresponding in men to the thumb and big toe) to digit V (the equivalent of the little finger and little toe).}} The most basal caseids like ''Eoasea'', ''[[Callibrachion]]'', and ''Martensius'' possessed the [[Plesiomorphy and symplesiomorphy\|plesiomorphic]] condition of early [[amniote]]s with a [[Phalanx bone#Development\|phalangeal formula]] of 2-3-4-5-3 for [[Manus (anatomy)\|manus]] and 2-3-4-5-4 for [[Pes (anatomy)\|pes]].<ref name="Reisz&Fröbisch2014"/><ref name="Spindler&al2015">{{cite journal\|last1=Spindler \|first1=F. \|author2=Falconnet, J. \|author3=Fröbisch, J. \|title=''Callibrachion'' and ''Datheosaurus'', two historical and previously mistaken basal caseasaurian synapsids from Europe \|journal=Acta Palaeontologica Polonica \|year=2015 \|volume=361 \|issue=3 \|pages=597–616 \|doi=10.4202/app.00221.2015 \|url=http://app.pan.pl/article/item/app002212015.html \|doi-access=free}}</ref><ref name="Berman&al.2020"/> In ''Euromycter'' the manus has a formula of 2-3-4-4-3 (the pes is unknown).<ref name="sigogneau-russell1974"/> In the more derived forms like ''Cotylorhynchus'' the manus and pes show a phalangeal formula of 2-2-3-3-2.{{refn\|group=nb\|In 1962 Olson and Barghusen had reported in the species ''Cotylorhynchus bransoni'' a foot with the phalangeal formula of 2-2-2-3-2, i.e. a smaller formula than in the type species ''C. romeri''. In 2015 Romano and Nicosia however demonstrated that the third digit of the foot of this species indeed had three phalanges and not two. Consequently the phalangeal formula of the foot of ''C. bransoni'' was 2-2-3-3-2 like ''C. romeri''.}}<ref name="Stovall&al.1966">{{cite journal\|last1=Stovall \|first1=J.W. \|author2=Price, L.I. \|author3=Romer, A.S. \|year=1966 \|title=The Postcranial Skeleton of the Giant Permian Pelycosaur ''Cotylorhynchus romeri'' \|journal=Bulletin of the Museum of Comparative Zoology \|volume=135 \|issue=1 \|pages=1–30 \|doi=}}</ref><ref name="Olson1962">{{cite journal\|last1=Olson \|first1=E.C. \|year=1962 \|title=Late Permian terrestrial vertebrates, U.S.A and U.S.S.R. \|journal=Transactions of the American Philosophical Society \|series=New Series \|volume=52 \|issue= 2\|pages=1–224 \|doi=10.2307/1005904\|jstor=1005904 }}</ref><ref name="Olson1968"/><ref name="Romano&Nicosia2015">{{cite journal\|last1=Romano \|first1=M. \|author2=Nicosia, U. \|year=2015 \|title=Cladistic analysis of Caseidae (Caseasauria, Synapsida): using the gap-weighting method to include taxa based on incomplete specimens \|journal=Palaeontology \|volume=58 \|issue=6 \|pages=1109–1130 \|doi=10.1111/pala.12197\|bibcode=2015Palgy..58.1109R \|s2cid=86489484 }}</ref> Along with this reduction in the number of phalanges, the proportions of the autopods also change in derived caseids with [[Metacarpal bones\|metacarpals]], [[Metatarsal bones\|metatarsals]], and [[Phalanx bone\|phalanges]] becoming shorter and broader. At the extreme of this specialization the genus ''Angelosaurus'' has short, broad, and smooth [[ungual]] phalanges which resemble [[Hoof\|hooves]] rather than [[claw]]s.<ref name="Olson1968"/><ref name="Ronchi&al.2011">{{cite journal\|last1=Ronchi \|first1=A. \|author2=Sacchi, E. \|author3=Romano, M. \|author4=Nicosia, U. \|year=2011 \|title=A huge caseid pelycosaur from north-western Sardinia and its bearing on European Permian stratigraphy and palaeobiogeography \|journal=Acta Palaeontologica Polonica \|volume=56 \|issue=4 \|pages=723–738 \|doi=10.4202/app.2010.0087\|s2cid=55085495 \|doi-access=free }}</ref>		The forelimbs of caseids are often more robust than the hindlimbs. It has indeed been observed that the bones of the forelimbs gained in robustness from the beginning of the evolution of the group, before the appearance of large species, while the hindlimbs remained slenderer. These characteristics suggest that the initial strengthening of the forelimbs was probably related to a particular function such as digging, and that this trait was later [[Exaptation\|exapted]] by more derived and larger caseids to support their weights of up to several hundred kilograms.<ref name="Romano2017"/> During their evolutionary history, caseids also show a reduction in their phalangeal formula.{{refn\|group=nb\|The phalangeal formula corresponds to the number of phalanges constituting each digits of the manus and pes of [[Tetrapoda\|tetrapods]]. It is listed starting from digit I (corresponding in men to the thumb and big toe) to digit V (the equivalent of the little finger and little toe).}} The most basal caseids like ''Eoasea'', ''[[Callibrachion]]'', and ''Martensius'' possessed the [[Plesiomorphy and symplesiomorphy\|plesiomorphic]] condition of early [[amniote]]s with a [[Phalanx bone#Development\|phalangeal formula]] of 2-3-4-5-3 for [[Manus (anatomy)\|manus]] and 2-3-4-5-4 for [[Pes (anatomy)\|pes]].<ref name="Reisz&Fröbisch2014"/><ref name="Spindler&al2015">{{cite journal\|last1=Spindler \|first1=F. \|author2=Falconnet, J. \|author3=Fröbisch, J. \|title=''Callibrachion'' and ''Datheosaurus'', two historical and previously mistaken basal caseasaurian synapsids from Europe \|journal=Acta Palaeontologica Polonica \|year=2015 \|volume=361 \|issue=3 \|pages=597–616 \|doi=10.4202/app.00221.2015 \|url=http://app.pan.pl/article/item/app002212015.html \|doi-access=free}}</ref><ref name="Berman&al.2020"/> In ''Euromycter'' the manus has a formula of 2-3-4-4-3 (the pes is unknown).<ref name="sigogneau-russell1974"/> In the more derived forms like ''Cotylorhynchus'' the manus and pes show a phalangeal formula of 2-2-3-3-2.{{refn\|group=nb\|In 1962 Olson and Barghusen had reported in the species ''Cotylorhynchus bransoni'' a foot with the phalangeal formula of 2-2-2-3-2, i.e. a smaller formula than in the type species ''C. romeri''. In 2015 Romano and Nicosia however demonstrated that the third digit of the foot of this species indeed had three phalanges and not two. Consequently the phalangeal formula of the foot of ''C. bransoni'' was 2-2-3-3-2 like ''C. romeri''.}}<ref name="Stovall&al.1966">{{cite journal\|last1=Stovall \|first1=J.W. \|author2=Price, L.I. \|author3=Romer, A.S. \|year=1966 \|title=The Postcranial Skeleton of the Giant Permian Pelycosaur ''Cotylorhynchus romeri'' \|journal=Bulletin of the Museum of Comparative Zoology \|volume=135 \|issue=1 \|pages=1–30 \|doi=}}</ref><ref name="Olson1962">{{cite journal\|last1=Olson \|first1=E.C. \|year=1962 \|title=Late Permian terrestrial vertebrates, U.S.A and U.S.S.R. \|journal=Transactions of the American Philosophical Society \|series=New Series \|volume=52 \|issue= 2\|pages=1–224 \|doi=10.2307/1005904\|jstor=1005904 }}</ref><ref name="Olson1968"/><ref name="Romano&Nicosia2015">{{cite journal\|last1=Romano \|first1=M. \|author2=Nicosia, U. \|year=2015 \|title=Cladistic analysis of Caseidae (Caseasauria, Synapsida): using the gap-weighting method to include taxa based on incomplete specimens \|journal=Palaeontology \|volume=58 \|issue=6 \|pages=1109–1130 \|doi=10.1111/pala.12197\|bibcode=2015Palgy..58.1109R \|s2cid=86489484 \|doi-access=free }}</ref> Along with this reduction in the number of phalanges, the proportions of the autopods also change in derived caseids with [[Metacarpal bones\|metacarpals]], [[Metatarsal bones\|metatarsals]], and [[Phalanx bone\|phalanges]] becoming shorter and broader. At the extreme of this specialization the genus ''Angelosaurus'' has short, broad, and smooth [[ungual]] phalanges which resemble [[Hoof\|hooves]] rather than [[claw]]s.<ref name="Olson1968"/><ref name="Ronchi&al.2011">{{cite journal\|last1=Ronchi \|first1=A. \|author2=Sacchi, E. \|author3=Romano, M. \|author4=Nicosia, U. \|year=2011 \|title=A huge caseid pelycosaur from north-western Sardinia and its bearing on European Permian stratigraphy and palaeobiogeography \|journal=Acta Palaeontologica Polonica \|volume=56 \|issue=4 \|pages=723–738 \|doi=10.4202/app.2010.0087\|s2cid=55085495 \|doi-access=free }}</ref>

	==Footprints==		==Footprints==
	Many vertebrate tracks have been proposed as belonging to Caseidae. In the early 2000s, large footprints known as ''[[Brontopus\|Brontopus giganteus]]'' from the Permian [[Lodève]] basin in southern [[France]] were considered to belong either to a caseid or [[dinocephalia]]n [[therapsid]].<ref name="Gand&al.2000">{{cite journal\|last1=Gand \|first1=G. \|author2=Garric, J. \|author3=Demathieu, G. \|author4=Ellenberger, P. \|title=La palichnofaune de vertébrés tétrapodes du Permien supérieur du bassin de Lodève (Languedoc-France) \|journal=Palaevertebrata \|year=2000 \|volume=29 \|issue=1\|pages=1–82 \|doi= \|url=https://palaeovertebrata.com/issues/view/103}}</ref> In 2019, Lorenzo Marchetti and colleagues, however, determined that dinocephalians were most likely the trackmakers of the ichnogenus ''Brontopus''.<ref name="Marchetti&al.2019a">{{cite journal\|last1=Marchetti \|first1=L. \|author2=Klein, H. \|author3=Buchwitz, M. \|author4=Ronchi, A. \|author5=Smith, R.M.H. \|author6=De Klerk, W.J. \|author7=Sciscio, L. \|author8=Groenewald, G.H. \|title=Permian-Triassic vertebrate footprints from South Africa: Ichnotaxonomy, producers and biostratigraphy through two major faunal crises \|journal=Gondwana Research \|year=2019 \|volume=72 \|issue= \|pages=139–168 \|doi=10.1016/j.gr.2019.03.009\|bibcode=2019GondR..72..139M \|s2cid=133781923 }}</ref> In 2012, Rafael Costa da Silva and colleagues proposed that the ichnogenus ''[[Chelichnus]]'', widely distributed in Permian desert facies (fossil [[dune]]s) of [[Europe]], [[North America]] and [[South America]], could represent caseids footprints.<ref name="Silva&al.2012">{{cite journal\|last1=Silva \|first1=R.C. \|author2=Sedor, F.A. \|author3=Fernandes, A.C.S. \|title=Fossil footprints from the Late Permian of Brazil: An example of hidden biodiversity \|journal=Journal of South American Earth Sciences \|year=2012 \|volume=38 \|issue= \|pages=31–43 \|doi=10.1016/j.jsames.2012.05.001\|bibcode=2012JSAES..38...31D }}</ref> In 2019, Marchetti and colleagues, however, reinterpreted ''Chelichnus'' as a ''[[nomen dubium]]'' and a [[taphotaxon]], this type of track showing false anatomical features generated by locomotion on sandy inclined paleosurface.<ref name="Marchetti&al.2019b">{{cite journal\|last1=Marchetti \|first1=L. \|author2=Voigt, S. \|author3=Lucas, S.G. \|title=An anatomy-consistent study of the Lopingian eolian tracks of Germany and Scotland reveals the first evidence of the end-Guadalupian mass extinction at low paleolatitudes of Pangea \|journal=Gondwana Research \|year=2019 \|volume=73 \|issue= \|pages=32–53 \|doi=10.1016/j.gr.2019.03.013\|bibcode=2019GondR..73...32M \|s2cid=146780505 }}</ref> In 2014, Eva Sacchi and colleagues described the ichnotaxon ''[[Dimetropus\|Dimetropus osageorum]]'' from about a hundred isolated footprints and several trackways from the early Permian [[Wellington Formation]] in [[Oklahoma]]. The morphological study of these footprints and their comparison with tetrapod skeletons revealed that they probably belong to a large caseid comparable in size to ''[[Cotylorhynchus\|Cotylorhynchus romeri]]'' (the latter from more recent strata).<ref name="Sacchi&al.2014">{{cite journal\|last1=Sacchi \|first1=E. \|author2=Cifelli, R. \|author3=Citton, P. \|author4=Nicosia, U. \|author5=Romano, M. \|title=''Dimetropus osageorum'' n. isp. from the Early Permian of Oklahoma (USA): A Trace and its Trackmaker \|journal=Ichnos: An International Journal for Plant and Animal Traces \|year=2014 \|volume=21 \|issue=3 \|pages=175–192 \|doi=10.1080/10420940.2014.933070\|s2cid=129567990 }}</ref> According to Sacchi and colleagues, some footprints from the Lodève Basin (unspecified geological formation) are similar to those from the Wellington Formation and these authors designate them as ''Dimetropus'' cf. ''osageorum''.<ref name="Sacchi&al.2014"/> The study by Sacchi et al. also demonstrates that the ichnogenus ''Dimetropus'' exhibits great morphological variation and that its producers can be assigned to different zoological groups among non-therapsid synapsids and not just [[Sphenacodontidae]] as once thought.<ref name="Sacchi&al.2014"/> In 2021, Rafel Matamales-Andreu and colleagues assigned footprints found in the Lower Permian ([[Artinskian]]-[[Kungurian]]) Port des Canonge Formation in [[Mallorca]] to a caseid. These footprints resemble those of ''D. osageorum'' but they also have differences. These footprints are left in open nomenclature as cf. ''Dimetropus'' sp.. They are, however, smaller than those of ''D. osageorum'' and would have been produced by a caseid of modest size with proportions comparable to ''[[Ennatosaurus]]''.<ref name="Matamales-Andreu&al.2021">{{cite journal\|last1=Matamales-Andreu \|first1=R. \|author2=Mujal, E. \|author3=Galobart, À. \|author4=Fortuny, J. \|title=Insight on the evolution of synapsid locomotion based on tetrapod tracks from the lower Permian of Mallorca (Balearic Islands, western Mediterranean) \|journal=Palaeogeography, Palaeoclimatology, Palaeoecology \|year=2021 \|volume=579 \|issue= \|pages=110589 \|doi=10.1016/j.palaeo.2021.110589\|bibcode=2021PPP...579k0589M }}</ref>		Many vertebrate tracks have been proposed as belonging to Caseidae. In the early 2000s, large footprints known as ''[[Brontopus\|Brontopus giganteus]]'' from the Permian [[Lodève]] basin in southern [[France]] were considered to belong either to a caseid or [[dinocephalia]]n [[therapsid]].<ref name="Gand&al.2000">{{cite journal\|last1=Gand \|first1=G. \|author2=Garric, J. \|author3=Demathieu, G. \|author4=Ellenberger, P. \|title=La palichnofaune de vertébrés tétrapodes du Permien supérieur du bassin de Lodève (Languedoc-France) \|journal=Palaevertebrata \|year=2000 \|volume=29 \|issue=1\|pages=1–82 \|doi= \|url=https://palaeovertebrata.com/issues/view/103}}</ref> In 2019, Lorenzo Marchetti and colleagues, however, determined that dinocephalians were most likely the trackmakers of the ichnogenus ''Brontopus''.<ref name="Marchetti&al.2019a">{{cite journal\|last1=Marchetti \|first1=L. \|author2=Klein, H. \|author3=Buchwitz, M. \|author4=Ronchi, A. \|author5=Smith, R.M.H. \|author6=De Klerk, W.J. \|author7=Sciscio, L. \|author8=Groenewald, G.H. \|title=Permian-Triassic vertebrate footprints from South Africa: Ichnotaxonomy, producers and biostratigraphy through two major faunal crises \|journal=Gondwana Research \|year=2019 \|volume=72 \|issue= \|pages=139–168 \|doi=10.1016/j.gr.2019.03.009\|bibcode=2019GondR..72..139M \|s2cid=133781923 \|doi-access=free }}</ref> In 2012, Rafael Costa da Silva and colleagues proposed that the ichnogenus ''[[Chelichnus]]'', widely distributed in Permian desert facies (fossil [[dune]]s) of [[Europe]], [[North America]] and [[South America]], could represent caseids footprints.<ref name="Silva&al.2012">{{cite journal\|last1=Silva \|first1=R.C. \|author2=Sedor, F.A. \|author3=Fernandes, A.C.S. \|title=Fossil footprints from the Late Permian of Brazil: An example of hidden biodiversity \|journal=Journal of South American Earth Sciences \|year=2012 \|volume=38 \|issue= \|pages=31–43 \|doi=10.1016/j.jsames.2012.05.001\|bibcode=2012JSAES..38...31D }}</ref> In 2019, Marchetti and colleagues, however, reinterpreted ''Chelichnus'' as a ''[[nomen dubium]]'' and a [[taphotaxon]], this type of track showing false anatomical features generated by locomotion on sandy inclined paleosurface.<ref name="Marchetti&al.2019b">{{cite journal\|last1=Marchetti \|first1=L. \|author2=Voigt, S. \|author3=Lucas, S.G. \|title=An anatomy-consistent study of the Lopingian eolian tracks of Germany and Scotland reveals the first evidence of the end-Guadalupian mass extinction at low paleolatitudes of Pangea \|journal=Gondwana Research \|year=2019 \|volume=73 \|issue= \|pages=32–53 \|doi=10.1016/j.gr.2019.03.013\|bibcode=2019GondR..73...32M \|s2cid=146780505 }}</ref> In 2014, Eva Sacchi and colleagues described the ichnotaxon ''[[Dimetropus\|Dimetropus osageorum]]'' from about a hundred isolated footprints and several trackways from the early Permian [[Wellington Formation]] in [[Oklahoma]]. The morphological study of these footprints and their comparison with tetrapod skeletons revealed that they probably belong to a large caseid comparable in size to ''[[Cotylorhynchus\|Cotylorhynchus romeri]]'' (the latter from more recent strata).<ref name="Sacchi&al.2014">{{cite journal\|last1=Sacchi \|first1=E. \|author2=Cifelli, R. \|author3=Citton, P. \|author4=Nicosia, U. \|author5=Romano, M. \|title=''Dimetropus osageorum'' n. isp. from the Early Permian of Oklahoma (USA): A Trace and its Trackmaker \|journal=Ichnos: An International Journal for Plant and Animal Traces \|year=2014 \|volume=21 \|issue=3 \|pages=175–192 \|doi=10.1080/10420940.2014.933070\|s2cid=129567990 }}</ref> According to Sacchi and colleagues, some footprints from the Lodève Basin (unspecified geological formation) are similar to those from the Wellington Formation and these authors designate them as ''Dimetropus'' cf. ''osageorum''.<ref name="Sacchi&al.2014"/> The study by Sacchi et al. also demonstrates that the ichnogenus ''Dimetropus'' exhibits great morphological variation and that its producers can be assigned to different zoological groups among non-therapsid synapsids and not just [[Sphenacodontidae]] as once thought.<ref name="Sacchi&al.2014"/> In 2021, Rafel Matamales-Andreu and colleagues assigned footprints found in the Lower Permian ([[Artinskian]]-[[Kungurian]]) Port des Canonge Formation in [[Mallorca]] to a caseid. These footprints resemble those of ''D. osageorum'' but they also have differences. These footprints are left in open nomenclature as cf. ''Dimetropus'' sp.. They are, however, smaller than those of ''D. osageorum'' and would have been produced by a caseid of modest size with proportions comparable to ''[[Ennatosaurus]]''.<ref name="Matamales-Andreu&al.2021">{{cite journal\|last1=Matamales-Andreu \|first1=R. \|author2=Mujal, E. \|author3=Galobart, À. \|author4=Fortuny, J. \|title=Insight on the evolution of synapsid locomotion based on tetrapod tracks from the lower Permian of Mallorca (Balearic Islands, western Mediterranean) \|journal=Palaeogeography, Palaeoclimatology, Palaeoecology \|year=2021 \|volume=579 \|issue= \|pages=110589 \|doi=10.1016/j.palaeo.2021.110589\|bibcode=2021PPP...579k0589M }}</ref>

	==Paleobiology==		==Paleobiology==

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	[[File:Arisierpeton simplex, premaxillae.png\|thumb\|right\|Premaxillae of ''[[Arisierpeton\|Arisierpeton simplex]]''. (A) Specimen GAA 00242 in anterior, labial, posterior, ventral, and dorsal views; (B) Specimen GAA 00239 in labial (showing a complete tooth with tricuspid distal end), partial labial, posterior and ventral views.]][[File:Cotylorhynchus romeria from Norman, Oklahoma.jpg\|thumb\|left\|Skeleton of ''Cotylorhynchus romeri'' on display at the [[Sam Noble Oklahoma Museum of Natural History]].]]		[[File:Arisierpeton simplex, premaxillae.png\|thumb\|right\|Premaxillae of ''[[Arisierpeton\|Arisierpeton simplex]]''. (A) Specimen GAA 00242 in anterior, labial, posterior, ventral, and dorsal views; (B) Specimen GAA 00239 in labial (showing a complete tooth with tricuspid distal end), partial labial, posterior and ventral views.]][[File:Cotylorhynchus romeria from Norman, Oklahoma.jpg\|thumb\|left\|Skeleton of ''Cotylorhynchus romeri'' on display at the [[Sam Noble Oklahoma Museum of Natural History]].]]

	The forelimbs of caseids are often more robust than the hindlimbs. It has indeed been observed that the bones of the forelimbs gained in robustness from the beginning of the evolution of the group, before the appearance of large species, while the hindlimbs remained slenderer. These characteristics suggest that the initial strengthening of the forelimbs was probably related to a particular function such as digging, and that this trait was later [[Exaptation\|exapted]] by more derived and larger caseids to support their weights of up to several hundred kilograms.<ref name="Romano2017"/> During their evolutionary history, caseids also show a reduction in their phalangeal formula.{{refn\|group=nb\|The phalangeal formula corresponds to the number of phalanges constituting each digits of the manus and pes of [[Tetrapoda\|tetrapods]]. It is listed starting from digit I (corresponding in men to the thumb and big toe) to digit V (the equivalent of the little finger and little toe).}} The most basal caseids like ''Eoasea'', ''[[Callibrachion]]'', and ''Martensius'' possessed the [[Plesiomorphy and symplesiomorphy\|plesiomorphic]] condition of early [[amniote]]s with a [[Phalanx bone#Development\|phalangeal formula]] of 2-3-4-5-3 for [[Manus (anatomy)\|manus]] and 2-3-4-5-4 for [[Pes (anatomy)\|pes]].<ref name="Reisz&Fröbisch2014"/><ref name="Spindler&al2015">{{cite journal\|last1=Spindler \|first1=F. \|author2=Falconnet, J. \|author3=Fröbisch, J. \|title=''Callibrachion'' and ''Datheosaurus'', two historical and previously mistaken basal caseasaurian synapsids from Europe \|journal=Acta Palaeontologica Polonica \|year=2015 \|volume=361 \|issue=3 \|pages=597–616 \|doi=10.4202/app.00221.2015 \|url=http://app.pan.pl/article/item/app002212015.html \|doi-access=free}}</ref><ref name="Berman&al.2020"/> In ''Euromycter'' the manus has a formula of 2-3-4-4-3 (the pes is unknown).<ref name="sigogneau-russell1974"/> In the more derived forms like ''Cotylorhynchus'' the manus and pes show a phalangeal formula of 2-2-3-3-2.{{refn\|group=nb\|In 1962 Olson and Barghusen had reported in the species ''Cotylorhynchus bransoni'' a foot with the phalangeal formula of 2-2-2-3-2, i.e. a smaller formula than in the type species ''C. romeri''. In 2015 Romano and Nicosia however demonstrated that the third digit of the foot of this species indeed had three phalanges and not two. Consequently the phalangeal formula of the foot of ''C. bransoni'' was 2-2-3-3-2 like ''C. romeri''.}}<ref name="Stovall&al.1966">{{cite journal\|last1=Stovall \|first1=J.W. \|author2=Price, L.I. \|author3=Romer, A.S. \|year=1966 \|title=The Postcranial Skeleton of the Giant Permian Pelycosaur ''Cotylorhynchus romeri'' \|journal=Bulletin of the Museum of Comparative Zoology \|volume=135 \|issue=1 \|pages=1–30 \|doi=}}</ref><ref name="Olson1962">{{cite journal\|last1=Olson \|first1=E.C. \|year=1962 \|title=Late Permian terrestrial vertebrates, U.S.A and U.S.S.R. \|journal=Transactions of the American Philosophical Society \|series=New Series \|volume=52 \|issue= 2\|pages=1–224 \|doi=10.2307/1005904\|jstor=1005904 }}</ref><ref name="Olson1968"/><ref name="Romano&Nicosia2015">{{cite journal\|last1=Romano \|first1=M. \|author2=Nicosia, U. \|year=2015 \|title=Cladistic analysis of Caseidae (Caseasauria, Synapsida): using the gap-weighting method to include taxa based on incomplete specimens \|journal=Palaeontology \|volume=58 \|issue=6 \|pages=1109–1130 \|doi=10.1111/pala.12197\|s2cid=86489484 }}</ref> Along with this reduction in the number of phalanges, the proportions of the autopods also change in derived caseids with [[Metacarpal bones\|metacarpals]], [[Metatarsal bones\|metatarsals]], and [[Phalanx bone\|phalanges]] becoming shorter and broader. At the extreme of this specialization the genus ''Angelosaurus'' has short, broad, and smooth [[ungual]] phalanges which resemble [[Hoof\|hooves]] rather than [[claw]]s.<ref name="Olson1968"/><ref name="Ronchi&al.2011">{{cite journal\|last1=Ronchi \|first1=A. \|author2=Sacchi, E. \|author3=Romano, M. \|author4=Nicosia, U. \|year=2011 \|title=A huge caseid pelycosaur from north-western Sardinia and its bearing on European Permian stratigraphy and palaeobiogeography \|journal=Acta Palaeontologica Polonica \|volume=56 \|issue=4 \|pages=723–738 \|doi=10.4202/app.2010.0087\|s2cid=55085495 \|doi-access=free }}</ref>		The forelimbs of caseids are often more robust than the hindlimbs. It has indeed been observed that the bones of the forelimbs gained in robustness from the beginning of the evolution of the group, before the appearance of large species, while the hindlimbs remained slenderer. These characteristics suggest that the initial strengthening of the forelimbs was probably related to a particular function such as digging, and that this trait was later [[Exaptation\|exapted]] by more derived and larger caseids to support their weights of up to several hundred kilograms.<ref name="Romano2017"/> During their evolutionary history, caseids also show a reduction in their phalangeal formula.{{refn\|group=nb\|The phalangeal formula corresponds to the number of phalanges constituting each digits of the manus and pes of [[Tetrapoda\|tetrapods]]. It is listed starting from digit I (corresponding in men to the thumb and big toe) to digit V (the equivalent of the little finger and little toe).}} The most basal caseids like ''Eoasea'', ''[[Callibrachion]]'', and ''Martensius'' possessed the [[Plesiomorphy and symplesiomorphy\|plesiomorphic]] condition of early [[amniote]]s with a [[Phalanx bone#Development\|phalangeal formula]] of 2-3-4-5-3 for [[Manus (anatomy)\|manus]] and 2-3-4-5-4 for [[Pes (anatomy)\|pes]].<ref name="Reisz&Fröbisch2014"/><ref name="Spindler&al2015">{{cite journal\|last1=Spindler \|first1=F. \|author2=Falconnet, J. \|author3=Fröbisch, J. \|title=''Callibrachion'' and ''Datheosaurus'', two historical and previously mistaken basal caseasaurian synapsids from Europe \|journal=Acta Palaeontologica Polonica \|year=2015 \|volume=361 \|issue=3 \|pages=597–616 \|doi=10.4202/app.00221.2015 \|url=http://app.pan.pl/article/item/app002212015.html \|doi-access=free}}</ref><ref name="Berman&al.2020"/> In ''Euromycter'' the manus has a formula of 2-3-4-4-3 (the pes is unknown).<ref name="sigogneau-russell1974"/> In the more derived forms like ''Cotylorhynchus'' the manus and pes show a phalangeal formula of 2-2-3-3-2.{{refn\|group=nb\|In 1962 Olson and Barghusen had reported in the species ''Cotylorhynchus bransoni'' a foot with the phalangeal formula of 2-2-2-3-2, i.e. a smaller formula than in the type species ''C. romeri''. In 2015 Romano and Nicosia however demonstrated that the third digit of the foot of this species indeed had three phalanges and not two. Consequently the phalangeal formula of the foot of ''C. bransoni'' was 2-2-3-3-2 like ''C. romeri''.}}<ref name="Stovall&al.1966">{{cite journal\|last1=Stovall \|first1=J.W. \|author2=Price, L.I. \|author3=Romer, A.S. \|year=1966 \|title=The Postcranial Skeleton of the Giant Permian Pelycosaur ''Cotylorhynchus romeri'' \|journal=Bulletin of the Museum of Comparative Zoology \|volume=135 \|issue=1 \|pages=1–30 \|doi=}}</ref><ref name="Olson1962">{{cite journal\|last1=Olson \|first1=E.C. \|year=1962 \|title=Late Permian terrestrial vertebrates, U.S.A and U.S.S.R. \|journal=Transactions of the American Philosophical Society \|series=New Series \|volume=52 \|issue= 2\|pages=1–224 \|doi=10.2307/1005904\|jstor=1005904 }}</ref><ref name="Olson1968"/><ref name="Romano&Nicosia2015">{{cite journal\|last1=Romano \|first1=M. \|author2=Nicosia, U. \|year=2015 \|title=Cladistic analysis of Caseidae (Caseasauria, Synapsida): using the gap-weighting method to include taxa based on incomplete specimens \|journal=Palaeontology \|volume=58 \|issue=6 \|pages=1109–1130 \|doi=10.1111/pala.12197\|bibcode=2015Palgy..58.1109R \|s2cid=86489484 }}</ref> Along with this reduction in the number of phalanges, the proportions of the autopods also change in derived caseids with [[Metacarpal bones\|metacarpals]], [[Metatarsal bones\|metatarsals]], and [[Phalanx bone\|phalanges]] becoming shorter and broader. At the extreme of this specialization the genus ''Angelosaurus'' has short, broad, and smooth [[ungual]] phalanges which resemble [[Hoof\|hooves]] rather than [[claw]]s.<ref name="Olson1968"/><ref name="Ronchi&al.2011">{{cite journal\|last1=Ronchi \|first1=A. \|author2=Sacchi, E. \|author3=Romano, M. \|author4=Nicosia, U. \|year=2011 \|title=A huge caseid pelycosaur from north-western Sardinia and its bearing on European Permian stratigraphy and palaeobiogeography \|journal=Acta Palaeontologica Polonica \|volume=56 \|issue=4 \|pages=723–738 \|doi=10.4202/app.2010.0087\|s2cid=55085495 \|doi-access=free }}</ref>

	==Footprints==		==Footprints==
Line 79:		Line 79:
	\|footer= Left: paleogeographic map of Earth at the end of the [[Paleozoic]] showing the known distribution of caseid synapsids. Right: close-up of the paleogeographic location of the caseid sites. '''1''' and '''2''' ''[[Ennatosaurus\|Ennatosaurus tecton]]'', Arkhangelsk Oblast, Russia, late Roadian – early Wordian; '''3''' ''[[Phreatophasma\|Phreatophasma aenigmaticum]]'', Bashkortostan, Russia, early Roadian; '''4''' ''[[Datheosaurus\|Datheosaurus macrourus]]'' Lower Silesian Voivodeship, Poland, Gzhelian; '''5''' ''[[Martensius\|Martensius bromackerensis]]'', Thuringia, Germany, Sakmarian; '''6''' ''[[Callibrachion\|Callibrachion gaudryi]]'', Saône-et-Loire, France, Asselian; '''7''' ''[[Euromycter\|Euromycter rutenus]]'' and ''[[Ruthenosaurus\|Ruthenosaurus russellorum]]'', Aveyron, France, late Artinskian; '''8''' ''[[Lalieudorhynchus\|Lalieudorhynchus gandi]]'', Hérault, France, Wordian – early Capitanian; '''9''' ''[[Alierasaurus\|Alierasaurus ronchii]]'', Nurra, Sardinia, Italy, Roadian; '''10''' ''[[Eocasea\|Eocasea martini]]'', Greenwood County, Kansas, late Pennsylvanian; '''11''' ''[[Angelosaurus\|Angelosaurus romeri]]'' and ''[[Cotylorhynchus\|Cotylorhynchus bransoni]]'', Kingfisher County, Oklahoma, early Roadian; '''12''' ''[[Cotylorhynchus\|Cotylorhynchus bransoni]]'', Blaine County, Oklahoma, early Roadian; '''13''' ''[[Cotylorhynchus\|Cotylorhynchus romeri]]'', Logan County, Oklahoma, mid-late Kungurian; '''14''' ''[[Cotylorhynchus\|Cotylorhynchus romeri]]'', Cleveland County, Oklahoma, mid-late Kungurian; '''15''' ''[[Oromycter\|Oromycter dolesorum]]'' and ''[[Arisierpeton\|Arisierpeton simplex]]'', Comanche County, Oklahoma, early Artinskian; '''16''' ''[[Cotylorhynchus\|Cotylorhynchus hancocki]]'', Hardeman County, Texas, late Kungurian – early Roadian; '''17''' ''[[Cotylorhynchus\|Cotylorhynchus hancocki]]'', ''[[Angelosaurus\|Angelosaurus dolani]]'', ''[[Angelosaurus\|A. greeni]]'', ''[[Caseoides\|Caseoides sanangeloensis]]'', and ''[[Caseopsis\|Caseopsis agilis]]'', Knox County, Texas, late Kungurian – early Roadian; '''18''' ''[[Casea\|Casea broilii]]'', Baylor County, Texas, mid-late Kungurian.}}		\|footer= Left: paleogeographic map of Earth at the end of the [[Paleozoic]] showing the known distribution of caseid synapsids. Right: close-up of the paleogeographic location of the caseid sites. '''1''' and '''2''' ''[[Ennatosaurus\|Ennatosaurus tecton]]'', Arkhangelsk Oblast, Russia, late Roadian – early Wordian; '''3''' ''[[Phreatophasma\|Phreatophasma aenigmaticum]]'', Bashkortostan, Russia, early Roadian; '''4''' ''[[Datheosaurus\|Datheosaurus macrourus]]'' Lower Silesian Voivodeship, Poland, Gzhelian; '''5''' ''[[Martensius\|Martensius bromackerensis]]'', Thuringia, Germany, Sakmarian; '''6''' ''[[Callibrachion\|Callibrachion gaudryi]]'', Saône-et-Loire, France, Asselian; '''7''' ''[[Euromycter\|Euromycter rutenus]]'' and ''[[Ruthenosaurus\|Ruthenosaurus russellorum]]'', Aveyron, France, late Artinskian; '''8''' ''[[Lalieudorhynchus\|Lalieudorhynchus gandi]]'', Hérault, France, Wordian – early Capitanian; '''9''' ''[[Alierasaurus\|Alierasaurus ronchii]]'', Nurra, Sardinia, Italy, Roadian; '''10''' ''[[Eocasea\|Eocasea martini]]'', Greenwood County, Kansas, late Pennsylvanian; '''11''' ''[[Angelosaurus\|Angelosaurus romeri]]'' and ''[[Cotylorhynchus\|Cotylorhynchus bransoni]]'', Kingfisher County, Oklahoma, early Roadian; '''12''' ''[[Cotylorhynchus\|Cotylorhynchus bransoni]]'', Blaine County, Oklahoma, early Roadian; '''13''' ''[[Cotylorhynchus\|Cotylorhynchus romeri]]'', Logan County, Oklahoma, mid-late Kungurian; '''14''' ''[[Cotylorhynchus\|Cotylorhynchus romeri]]'', Cleveland County, Oklahoma, mid-late Kungurian; '''15''' ''[[Oromycter\|Oromycter dolesorum]]'' and ''[[Arisierpeton\|Arisierpeton simplex]]'', Comanche County, Oklahoma, early Artinskian; '''16''' ''[[Cotylorhynchus\|Cotylorhynchus hancocki]]'', Hardeman County, Texas, late Kungurian – early Roadian; '''17''' ''[[Cotylorhynchus\|Cotylorhynchus hancocki]]'', ''[[Angelosaurus\|Angelosaurus dolani]]'', ''[[Angelosaurus\|A. greeni]]'', ''[[Caseoides\|Caseoides sanangeloensis]]'', and ''[[Caseopsis\|Caseopsis agilis]]'', Knox County, Texas, late Kungurian – early Roadian; '''18''' ''[[Casea\|Casea broilii]]'', Baylor County, Texas, mid-late Kungurian.}}

	Caseid [[fossil]]s have been found in the southern and central [[United States]] ([[Texas]], [[Oklahoma]], [[Kansas]]), [[France]], [[Germany]], [[Poland]], [[Sardinia]], and [[European Russia]]. This geographical distribution corresponds to the paleoequatorial belt of [[Pangaea]] during the Carboniferous and the Permian, with the exception of the Russian localities which were located at the level of the [[30th parallel north]] where a more temperate climate prevailed. The absence in Russia of terrestrial vertebrate localities prior to the [[Kungurian]] does not allow us to know the precise moment of the arrival of caseids in this region of the world.<ref name="Brocklehurst&Fröbisch2017"/> According to Werneburg and colleagues, the presence of caseids at this [[Latitude\|paleolatitude]] suggests their possible existence in the temperate regions of [[Gondwana]].<ref name="Werneburg&al.2022"/> A possible Gondwanan occurrence was proposed as early as the 1990s by Michael S.Y. Lee and in 2021 by Asher J. Lichtig and Spencer G. Lucas, who reinterpreted the middle Permian taxon ''[[Eunotosaurus]]'' from [[South Africa]] as a small burrowing caseid.<ref name="Lee1995">{{Cite thesis \|degree=Ph.D. \|title= Evolutionary Morphology of Pareiasaurs \|url= https://www.academia.edu/9770306 \|last=Lee \|first=M.S.Y. \|year=1995 \|publisher= Dept of Zoology, University of Cambridge \|location=Cambridge \|pages=151–155}}</ref><ref name="Lichtig&Lucas2021">{{cite journal\|last1=Lichtig \|first1=A.J. \|author2=Lucas, S.G. \|title=''Chinlechelys'' from the Upper Triassic of New Mexico, USA, and the origin of turtles \|journal=Palaeontologia Electronica \|year=2021 \|volume=24\|issue=(1):a13 \|pages=1–49 \|doi=10.26879/886\|s2cid=233454789 \|doi-access=free }}</ref> Between these two dates, other researchers classified ''Eunotosaurus'' as a [[Parareptilia\|parareptile]]<ref name="Modesto2000">{{cite journal \|last1=Modesto \|first1=S.P. \|year=2000 \|title=''Eunotosaurus africanus'' and the Gondwanan ancestry of anapsid reptiles \|journal=Palaeontologia Africana \|volume=36 \|issue= \|pages=15–20 \|doi= \|url=https://wiredspace.wits.ac.za/handle/10539/16375}}</ref> or a [[pantestudines]].<ref name="Bever&al.2015">{{cite journal \|last1=Bever \|first1=G.S. \|author2=Lyson, T.R. \|author3=Field, D.J. \|author4=Bhullar, B-A.S. \|year=2015 \|title=Evolutionary origin of the turtle skull \|journal=Nature \|volume=525 \|issue=7568 \|pages=239–242 \|doi=10.1038/nature14900\|pmid=26331544 \|bibcode=2015Natur.525..239B \|s2cid=4401555 }}</ref><ref name="Lyson&al.2016">{{cite journal \|last1=Lyson \|first1=T.R. \|author2=Rubidge, B.S. \|author3=Scheyer, T.M. \|author4=de Queiroz, K. \|author5=Schachner, E.R. \|author6=Smith, R.M.H. \|author7=Botha-Brink, J. \|author8=Bever, G.S. \|year=2016 \|title=Fossorial origin of the turtle shell \|journal=Current Biology \|volume=26 \|issue=14 \|pages=1887–1894 \|doi=10.1016/j.cub.2016.05.020\|pmid=27426515 \|s2cid=3935231 \|doi-access=free }}</ref> If the first three authors are correct, ''Eunotosaurus'' would be the first caseid found in the [[Southern Hemisphere]] and the last known representative of the family with a latest [[Capitanian]] age for the most recent specimens.<ref name="Day&Smith2020">{{cite journal\|last1=Day \|first1=M.O. \|author2=Smith, R.M.H. \|year=2020 \|title=Biostratigraphy of the ''Endothiodon'' Assemblage Zone (Beaufort Group, Karoo Supergroup), South Africa \|journal=South African Journal of Geology \|volume=123 \|issue=2 \|pages=165–180 \|doi=10.25131/sajg.123.0011\|s2cid=225834576 }}</ref> Elsewhere in Gondwana, the presence of the [[ichnotaxon]] ''[[Dimetropus]]'' in the Permian of [[Morocco]] suggests the future discovery of basal synapsids (including caseids) in the Permian basins of [[North Africa]], which were also part of the paleoequatorial belt and constituted one of the main [[Wildlife corridor\|migratory routes]] between the Northern and Southern Hemispheres.<ref name="Voigt&al.2011a">{{cite journal\|last1=Voigt \|first1=S. \|author2=Lagnaoui, A. \|author3=Hminna, A. \|author4=Saber, H. \|author5=Schneider, J.W. \|year=2011 \|title=Revisional notes on the Permian tetrapod ichnofauna from the Tiddas Basin, central Morocco \|journal=Palaeogeography, Palaeoclimatology, Palaeoecology \|volume=302 \|issue=3–4 \|pages=474–483 \|doi=10.1016/j.palaeo.2011.02.010\|bibcode=2011PPP...302..474V }}</ref><ref name="Voigt&al.2011b">{{cite journal\|last1=Voigt \|first1=S. \|author2=Saber, H. \|author3=Schneider, J.W. \|author4=Hmich, D. \|author5=Hminna, A. \|year=2011 \|title=Late Carboniferous-Early Permian Tetrapod Ichnofauna from the Khenifra Basin, Central Morocco \|journal=Geobios \|volume=44 \|issue=4 \|pages=399–407 \|doi=10.1016/j.geobios.2010.11.008}}</ref>		Caseid [[fossil]]s have been found in the southern and central [[United States]] ([[Texas]], [[Oklahoma]], [[Kansas]]), [[France]], [[Germany]], [[Poland]], [[Sardinia]], and [[European Russia]]. This geographical distribution corresponds to the paleoequatorial belt of [[Pangaea]] during the Carboniferous and the Permian, with the exception of the Russian localities which were located at the level of the [[30th parallel north]] where a more temperate climate prevailed. The absence in Russia of terrestrial vertebrate localities prior to the [[Kungurian]] does not allow us to know the precise moment of the arrival of caseids in this region of the world.<ref name="Brocklehurst&Fröbisch2017"/> According to Werneburg and colleagues, the presence of caseids at this [[Latitude\|paleolatitude]] suggests their possible existence in the temperate regions of [[Gondwana]].<ref name="Werneburg&al.2022"/> A possible Gondwanan occurrence was proposed as early as the 1990s by Michael S.Y. Lee and in 2021 by Asher J. Lichtig and Spencer G. Lucas, who reinterpreted the middle Permian taxon ''[[Eunotosaurus]]'' from [[South Africa]] as a small burrowing caseid.<ref name="Lee1995">{{Cite thesis \|degree=Ph.D. \|title= Evolutionary Morphology of Pareiasaurs \|url= https://www.academia.edu/9770306 \|last=Lee \|first=M.S.Y. \|year=1995 \|publisher= Dept of Zoology, University of Cambridge \|location=Cambridge \|pages=151–155}}</ref><ref name="Lichtig&Lucas2021">{{cite journal\|last1=Lichtig \|first1=A.J. \|author2=Lucas, S.G. \|title=''Chinlechelys'' from the Upper Triassic of New Mexico, USA, and the origin of turtles \|journal=Palaeontologia Electronica \|year=2021 \|volume=24\|issue=(1):a13 \|pages=1–49 \|doi=10.26879/886\|s2cid=233454789 \|doi-access=free }}</ref> Between these two dates, other researchers classified ''Eunotosaurus'' as a [[Parareptilia\|parareptile]]<ref name="Modesto2000">{{cite journal \|last1=Modesto \|first1=S.P. \|year=2000 \|title=''Eunotosaurus africanus'' and the Gondwanan ancestry of anapsid reptiles \|journal=Palaeontologia Africana \|volume=36 \|issue= \|pages=15–20 \|doi= \|url=https://wiredspace.wits.ac.za/handle/10539/16375}}</ref> or a [[pantestudines]].<ref name="Bever&al.2015">{{cite journal \|last1=Bever \|first1=G.S. \|author2=Lyson, T.R. \|author3=Field, D.J. \|author4=Bhullar, B-A.S. \|year=2015 \|title=Evolutionary origin of the turtle skull \|journal=Nature \|volume=525 \|issue=7568 \|pages=239–242 \|doi=10.1038/nature14900\|pmid=26331544 \|bibcode=2015Natur.525..239B \|s2cid=4401555 }}</ref><ref name="Lyson&al.2016">{{cite journal \|last1=Lyson \|first1=T.R. \|author2=Rubidge, B.S. \|author3=Scheyer, T.M. \|author4=de Queiroz, K. \|author5=Schachner, E.R. \|author6=Smith, R.M.H. \|author7=Botha-Brink, J. \|author8=Bever, G.S. \|year=2016 \|title=Fossorial origin of the turtle shell \|journal=Current Biology \|volume=26 \|issue=14 \|pages=1887–1894 \|doi=10.1016/j.cub.2016.05.020\|pmid=27426515 \|s2cid=3935231 \|doi-access=free }}</ref> If the first three authors are correct, ''Eunotosaurus'' would be the first caseid found in the [[Southern Hemisphere]] and the last known representative of the family with a latest [[Capitanian]] age for the most recent specimens.<ref name="Day&Smith2020">{{cite journal\|last1=Day \|first1=M.O. \|author2=Smith, R.M.H. \|year=2020 \|title=Biostratigraphy of the ''Endothiodon'' Assemblage Zone (Beaufort Group, Karoo Supergroup), South Africa \|journal=South African Journal of Geology \|volume=123 \|issue=2 \|pages=165–180 \|doi=10.25131/sajg.123.0011\|bibcode=2020SAJG..123..165D \|s2cid=225834576 }}</ref> Elsewhere in Gondwana, the presence of the [[ichnotaxon]] ''[[Dimetropus]]'' in the Permian of [[Morocco]] suggests the future discovery of basal synapsids (including caseids) in the Permian basins of [[North Africa]], which were also part of the paleoequatorial belt and constituted one of the main [[Wildlife corridor\|migratory routes]] between the Northern and Southern Hemispheres.<ref name="Voigt&al.2011a">{{cite journal\|last1=Voigt \|first1=S. \|author2=Lagnaoui, A. \|author3=Hminna, A. \|author4=Saber, H. \|author5=Schneider, J.W. \|year=2011 \|title=Revisional notes on the Permian tetrapod ichnofauna from the Tiddas Basin, central Morocco \|journal=Palaeogeography, Palaeoclimatology, Palaeoecology \|volume=302 \|issue=3–4 \|pages=474–483 \|doi=10.1016/j.palaeo.2011.02.010\|bibcode=2011PPP...302..474V }}</ref><ref name="Voigt&al.2011b">{{cite journal\|last1=Voigt \|first1=S. \|author2=Saber, H. \|author3=Schneider, J.W. \|author4=Hmich, D. \|author5=Hminna, A. \|year=2011 \|title=Late Carboniferous-Early Permian Tetrapod Ichnofauna from the Khenifra Basin, Central Morocco \|journal=Geobios \|volume=44 \|issue=4 \|pages=399–407 \|doi=10.1016/j.geobios.2010.11.008\|bibcode=2011Geobi..44..399V }}</ref>

	==Phylogeny==		==Phylogeny==

← Previous revision		Revision as of 15:11, 22 August 2024
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	\|footer=''Cotylorhynchus romeri''. Top : skull reconstruction in left lateral view, and medial and lateral views of mandible; bottom : skull reconstruction in dorsal, ventral, posterior and anterior views.}}[[File:Euromycter rutenus 6778 retouched.jpg\|thumb\|right\|Skull of ''[[Euromycter\|Euromycter rutenus]]'' in ventral view allowing to see the numerous small teeth adorning the bones of the palate (the [[parasphenoid]], triangular in shape in the center, and the very elongated [[Pterygoid bone\|pterygoids]] on each side). The hyoid apparatus is not present here because it was removed during the preparation of the palate.]]		\|footer=''Cotylorhynchus romeri''. Top : skull reconstruction in left lateral view, and medial and lateral views of mandible; bottom : skull reconstruction in dorsal, ventral, posterior and anterior views.}}[[File:Euromycter rutenus 6778 retouched.jpg\|thumb\|right\|Skull of ''[[Euromycter\|Euromycter rutenus]]'' in ventral view allowing to see the numerous small teeth adorning the bones of the palate (the [[parasphenoid]], triangular in shape in the center, and the very elongated [[Pterygoid bone\|pterygoids]] on each side). The hyoid apparatus is not present here because it was removed during the preparation of the palate.]]

	Caseids measured from less than {{convert\|1\|m}} to {{convert\|7\|m}} in length.<ref name="Romano2017">{{cite journal\|last1=Romano \|first1=M. \|year=2017 \|title=Long bone scaling of caseid synapsids: a combined morphometric and cladistic approach \|journal=Lethaia \|volume=50 \|issue=4 \|pages=511–526 \|doi=10.1111/let.12207}}</ref><ref name="Angielczyk&Kammerer2018">{{cite book\|last=Angielczyk\|first=K.D. \|author2=Kammerer, C.F. \|year=2018 \|chapter=Non-Mammalian synapsids : the deep roots of the mammalian family tree \|pages=117–198 \|title=Handbook of Zoology : Mammalian Evolution, Diversity and Systematics \|editor1-last=Zachos\|editor1-first=F.E. \|editor2=Asher, R.J. \|publisher=de Gruyter \|location=Berlin \|isbn=978-3-11-027590-2}}</ref><ref name="Romano&al.2018">{{cite journal\|last1=Romano\|first1=M. \|author2=Citton, P. \|author3=Maganuco, S. \|author4=Sacchi, E. \|author5=Caratelli, M. \|author6=Ronchi, A. \|author7=Nicosia, U. \|year=2018 \|title=New basal synapsid discovery at the Permian outcrop of Torre del Porticciolo (Alghero, Italy) \|journal=Geological Journal \|volume=54 \|issue=3 \|pages=1554–1566 \|doi=10.1002/gj.3250\|s2cid=133755506 }}</ref> They had a small head wider than high and with a forward-inclined snout, a very short neck, a long tail, robust forelimbs, and a body of variable proportions depending on their diet. Small insectivorous species like ''[[Eocasea]]'' had an unexpanded trunk.<ref name="Reisz&Fröbisch2014">{{cite journal\|last1=Reisz\|first1=R.R. \|author2=Fröbisch, J. \|year=2014 \|title=Oldest caseid synapsid from the late Pennsylvanian of Kansas, and the evolution of herbivory in terrestrial vertebrates \|journal=PLOS ONE \|volume=9 \|issue=4 \|pages=e94518 \|doi=10.1371/journal.pone.0094518 \|pmid=24739998 \|pmc=3989228\|bibcode=2014PLoSO...994518R \|doi-access=free }}</ref> Others with an omnivorous diet like ''[[Martensius]]'' had a barely enlarged rib cage, a more elongated skull, smaller [[nostril]]s, and a snout less inclined forward than in herbivorous caseids.<ref name="Berman&al.2020">{{cite journal\|last1=Berman \|first1=D.S. \|author2=Maddin, H.C. \|author3=Henrici, A.C. \|author4=Sumida, S.S. \|author5=Scott, D. \|author6=Reisz, R.R. \|title=New primitive Caseid (Synapsida, Caseasauria) from the Early Permian of Germany \|journal=Annals of Carnegie Museum \|year=2020 \|volume=86 \|issue=1 \|pages=43–75 \|doi=10.2992/007.086.0103\|s2cid=216027787 }}</ref> The latter were characterized by their disproportionately small [[skull]] compared to the size of the body. The postcranial skeleton indeed shows a spectacular increase in the volume of the [[rib cage]], which becomes very wide and barrel-shaped, probably to accommodate a particularly developed [[Gastrointestinal tract#Lower gastrointestinal tract\|intestine]], necessary for the digestion of high-fiber rich plants. In these forms, the skull has very large external nostrils and a very short facial region with a strong forward inclination of the end of the snout which clearly overhangs the dental row. The [[Skull#Fenestrae\|temporal fenestrae]] are also relatively large (especially in ''[[Ennatosaurus]]''), the [[Supratemporal bone\|supratemporals]] are large in size, and, on the [[Occipital bone\|occipital]] surface, the paroccipital [[Process (anatomy)\|processes]] are massively developed, establishing strong supporting contacts with the [[Squamosal bone\|squamosals]].<ref name="Kemp1982">{{cite book\|last=Kemp \|first=T.S. \|year=1982 \|chapter=Pelycosaurs \|pages=39 \|title=Mammal-like reptiles and the origin of Mammals \|editor1-last=Kemp\|editor1-first=T.S. \|publisher=Academic Press \|location=London \|isbn=978-0124041202}}</ref> The dorsal surface of the skull is covered with numerous small pits. These suggest the presence of large [[Reptile scale\|scales]] on the head of these animals.<ref name="Romano&al.2017a">{{cite journal\|last1=Romano \|first1=M. \|author2=Brocklehurst, N. \|author3=Fröbisch, J. \|year=2017 \|title=The postcranial skeleton of ''Ennatosaurus tecton'' (Synapsida, Caseidae) \|journal=Journal of Systematic Palaeontology \|volume=16 \|issue= 13\|pages=1097–1122 \|doi=10.1080/14772019.2017.1367729\|s2cid=89922565 }}</ref> Numerous [[Lip\|labial]] [[Foramen\|foramina]] running parallel to the ventral edge of the [[premaxilla]] and [[maxilla]], as well as along the dorsal edge of the [[Mandible#Other vertebrates\|dentary]], suggest the presence of scaly "lips" which must have concealed the dentition when the jaws were closed.<ref name="Reisz&al.2022">{{cite journal\|last1=Reisz \|first1=R.R. \|author2=Scott, D. \|author3=Modesto, S.P. \|year=2022 \|title=Cranial Anatomy of the Caseid Synapsid ''Cotylorhynchus romeri'', a Large Terrestrial Herbivore From the Lower Permian of Oklahoma, U.S.A \|journal=Frontiers in Earth Science \|volume=10 \|issue= \|pages=1–19 \|doi=10.3389/feart.2022.847560\|doi-access=free }}</ref><ref name="Romano&al.2017a"/> The [[Tooth\|teeth]], simply conical and pointed in insectivorous species, adopt in herbivorous species a leafy or spatulate morphology and are provided with more or less numerous cuspules. Numerous small teeth also adorned several bones of the palate. Herbivorous species do not show a simple evolutionary trend towards increasing tooth complexity.<ref name="Maddin&al.2008">{{cite journal \|last1=Maddin \|first1=H.C. \|author2=Sidor, C.A. \|author3=Reisz, R.R. \|year=2008 \|title=Cranial anatomy of ''Ennatosaurus tecton'' (Synapsida: Caseidae) from the Middle Permian of Russia and the evolutionary relationships of Caseidae \|journal=Journal of Vertebrate Paleontology \|volume=28 \|issue=1 \|pages=160–180 \|doi=10.1671/0272-4634(2008)28[160:CAOETS]2.0.CO;2\|s2cid=44064927 }}</ref> Thus, the teeth of the basal taxa ''[[Casea]]'' and ''[[Arisierpeton]]'' have three cuspules<ref name="Olson1968">{{cite journal\|last1=Olson \|first1=E.C. \|year=1968 \|title=The family Caseidae \|journal=Fieldiana: Geology \|volume=17 \|issue= \|pages=225–349\| doi= }}</ref><ref name="Reisz2019">{{cite journal \|last1=Reisz \|first1=R. \|year=2019 \|title=A small caseid synapsid, ''Arisierpeton simplex'' gen. et sp. nov., from the early Permian of Oklahoma, with a discussion of synapsid diversity at the classic Richards Spur locality \|journal=PeerJ \|volume=7e6615 \|issue= \|pages=1–20 \|doi=10.7717/peerj.6615 \|pmid=30997285 \|pmc=6462398 \|doi-access=free }}</ref> just like in the more derived forms ''[[Cotylorhynchus]]'' and ''[[Caseopsis]]''.<ref name="Olson1968"/> ''Ennatosaurus'' and ''[[Euromycter]]'', which occupy an intermediate [[Phylogenetics\|phylogenetic]] position, have teeth bearing 5 to 7 cuspules and 5 to 8 cuspules respectively.<ref name="Maddin&al.2008"/><ref name="sigogneau-russell1974">{{cite journal \|last1=Sigoneau-Russell \|first1=D. \|author2=Russel, D.E. \|year=1974 \|title=Étude du premier caséidé (Reptilia, Pelycosauria) d'Europe occidentale \|journal=Bulletin du Muséum National d'Histoire Naturelle \|series=Série 3 \|volume=38 \|issue=230 \|pages=145–215 \|doi= \|url=http://bibliotheques.mnhn.fr/EXPLOITATION/infodoc/digitalCollections/viewerpopup.aspx?seid=BMSCT_S003_1974_T230_N038}}</ref> ''[[Angelosaurus]]'', one of the most derived caseids, has teeth with 5 cuspules.<ref name="Olson1968"/> In ''Angelosaurus'' the teeth have a bulbous morphology with very short and wide crowns. Their sturdiness and the significant wear they show, indicate that ''Angelosaurus'' must have fed on tougher plants than those on which most other herbivorous caseids fed.<ref name="Olson1968"/> Herbivorous caseids also show very different dietary adaptations from those seen in another group of basal synapsids, the [[Edaphosauridae]]. The latter had, in addition to the marginal dentition, a dental battery made up of numerous teeth located both on the [[palate]] and on the inner surface of the lower jaws. In herbivorous caseids, the palatal teeth are smaller, and the inner surface of the lower jaws bears no teeth. Instead of a dental battery, they had a massive [[tongue]] (as indicated by the presence of a highly developed [[hyoid apparatus]] found in ''Ennatosaurus'' and ''Euromycter'') perhaps rough, with which they had to compress food against the palatal teeth.<ref name="Olson1968"/><ref name="Reisz&al.2022"/>		Caseids measured from less than {{convert\|1\|m}} to {{convert\|7\|m}} in length.<ref name="Romano2017">{{cite journal\|last1=Romano \|first1=M. \|year=2017 \|title=Long bone scaling of caseid synapsids: a combined morphometric and cladistic approach \|journal=Lethaia \|volume=50 \|issue=4 \|pages=511–526 \|doi=10.1111/let.12207}}</ref><ref name="Angielczyk&Kammerer2018">{{cite book\|last=Angielczyk\|first=K.D. \|author2=Kammerer, C.F. \|year=2018 \|chapter=Non-Mammalian synapsids : the deep roots of the mammalian family tree \|pages=117–198 \|title=Handbook of Zoology : Mammalian Evolution, Diversity and Systematics \|editor1-last=Zachos\|editor1-first=F.E. \|editor2=Asher, R.J. \|publisher=de Gruyter \|location=Berlin \|isbn=978-3-11-027590-2}}</ref><ref name="Romano&al.2018">{{cite journal\|last1=Romano\|first1=M. \|author2=Citton, P. \|author3=Maganuco, S. \|author4=Sacchi, E. \|author5=Caratelli, M. \|author6=Ronchi, A. \|author7=Nicosia, U. \|year=2018 \|title=New basal synapsid discovery at the Permian outcrop of Torre del Porticciolo (Alghero, Italy) \|journal=Geological Journal \|volume=54 \|issue=3 \|pages=1554–1566 \|doi=10.1002/gj.3250\|s2cid=133755506 }}</ref> They had a small head wider than high and with a forward-inclined snout, a very short neck, a long tail, robust forelimbs, and a body of variable proportions depending on their diet. Small insectivorous species like ''[[Eocasea]]'' had an unexpanded trunk.<ref name="Reisz&Fröbisch2014">{{cite journal\|last1=Reisz\|first1=R.R. \|author2=Fröbisch, J. \|year=2014 \|title=Oldest caseid synapsid from the late Pennsylvanian of Kansas, and the evolution of herbivory in terrestrial vertebrates \|journal=PLOS ONE \|volume=9 \|issue=4 \|pages=e94518 \|doi=10.1371/journal.pone.0094518 \|pmid=24739998 \|pmc=3989228\|bibcode=2014PLoSO...994518R \|doi-access=free }}</ref> Others with an omnivorous diet like ''[[Martensius]]'' had a barely enlarged rib cage, a more elongated skull, smaller [[nostril]]s, and a snout less inclined forward than in herbivorous caseids.<ref name="Berman&al.2020">{{cite journal\|last1=Berman \|first1=D.S. \|author2=Maddin, H.C. \|author3=Henrici, A.C. \|author4=Sumida, S.S. \|author5=Scott, D. \|author6=Reisz, R.R. \|title=New primitive Caseid (Synapsida, Caseasauria) from the Early Permian of Germany \|journal=Annals of Carnegie Museum \|year=2020 \|volume=86 \|issue=1 \|pages=43–75 \|doi=10.2992/007.086.0103\|s2cid=216027787 }}</ref> The latter were characterized by their disproportionately small [[skull]] compared to the size of the body. The postcranial skeleton indeed shows a spectacular increase in the volume of the [[rib cage]], which becomes very wide and barrel-shaped, probably to accommodate a particularly developed [[Gastrointestinal tract#Lower gastrointestinal tract\|intestine]], necessary for the digestion of high-fiber rich plants. In these forms, the skull has very large external nostrils and a very short facial region with a strong forward inclination of the end of the snout which clearly overhangs the dental row. The [[Skull#Fenestrae\|temporal fenestrae]] are also relatively large (especially in ''[[Ennatosaurus]]''), the [[Supratemporal bone\|supratemporals]] are large in size, and, on the [[Occipital bone\|occipital]] surface, the paroccipital [[Process (anatomy)\|processes]] are massively developed, establishing strong supporting contacts with the [[Squamosal bone\|squamosals]].<ref name="Kemp1982">{{cite book\|last=Kemp \|first=T.S. \|year=1982 \|chapter=Pelycosaurs \|pages=39 \|title=Mammal-like reptiles and the origin of Mammals \|editor1-last=Kemp\|editor1-first=T.S. \|publisher=Academic Press \|location=London \|isbn=978-0124041202}}</ref> The dorsal surface of the skull is covered with numerous small pits. These suggest the presence of large [[Reptile scale\|scales]] on the head of these animals.<ref name="Romano&al.2017a">{{cite journal\|last1=Romano \|first1=M. \|author2=Brocklehurst, N. \|author3=Fröbisch, J. \|year=2017 \|title=The postcranial skeleton of ''Ennatosaurus tecton'' (Synapsida, Caseidae) \|journal=Journal of Systematic Palaeontology \|volume=16 \|issue= 13\|pages=1097–1122 \|doi=10.1080/14772019.2017.1367729\|s2cid=89922565 }}</ref> Numerous [[Lip\|labial]] [[Foramen\|foramina]] running parallel to the ventral edge of the [[premaxilla]] and [[maxilla]], as well as along the dorsal edge of the [[Mandible#Other vertebrates\|dentary]], suggest the presence of scaly "lips" which must have concealed the dentition when the jaws were closed.<ref name="Reisz&al.2022">{{cite journal\|last1=Reisz \|first1=R.R. \|author2=Scott, D. \|author3=Modesto, S.P. \|year=2022 \|title=Cranial Anatomy of the Caseid Synapsid ''Cotylorhynchus romeri'', a Large Terrestrial Herbivore From the Lower Permian of Oklahoma, U.S.A \|journal=Frontiers in Earth Science \|volume=10 \|issue= \|pages=1–19 \|doi=10.3389/feart.2022.847560\|doi-access=free }}</ref><ref name="Romano&al.2017a"/> The [[Tooth\|teeth]], simply conical and pointed in insectivorous species, adopt in herbivorous species a leafy or spatulate morphology and are provided with more or less numerous cuspules. Numerous small teeth also adorned several bones of the palate. Herbivorous species do not show a simple evolutionary trend towards increasing tooth complexity.<ref name="Maddin&al.2008">{{cite journal \|last1=Maddin \|first1=H.C. \|author2=Sidor, C.A. \|author3=Reisz, R.R. \|year=2008 \|title=Cranial anatomy of ''Ennatosaurus tecton'' (Synapsida: Caseidae) from the Middle Permian of Russia and the evolutionary relationships of Caseidae \|journal=Journal of Vertebrate Paleontology \|volume=28 \|issue=1 \|pages=160–180 \|doi=10.1671/0272-4634(2008)28[160:CAOETS]2.0.CO;2\|s2cid=44064927 }}</ref> Thus, the teeth of the basal taxa ''[[Casea]]'' and ''[[Arisierpeton]]'' have three cuspules<ref name="Olson1968">{{cite journal\|last1=Olson \|first1=E.C. \|year=1968 \|title=The family Caseidae \|journal=Fieldiana: Geology \|volume=17 \|issue= \|pages=225–349\| doi= }}</ref><ref name="Reisz2019">{{cite journal \|last1=Reisz \|first1=R. \|year=2019 \|title=A small caseid synapsid, ''Arisierpeton simplex'' gen. et sp. nov., from the early Permian of Oklahoma, with a discussion of synapsid diversity at the classic Richards Spur locality \|journal=PeerJ \|volume=7e6615 \|issue= \|pages=1–20 \|doi=10.7717/peerj.6615 \|pmid=30997285 \|pmc=6462398 \|doi-access=free }}</ref> just like in the more derived forms ''[[Cotylorhynchus]]'' and ''[[Caseopsis]]''.<ref name="Olson1968"/> ''Ennatosaurus'' and ''[[Euromycter]]'', which occupy an intermediate [[Phylogenetics\|phylogenetic]] position, have teeth bearing 5 to 7 cuspules and 5 to 8 cuspules respectively.<ref name="Maddin&al.2008"/><ref name="sigogneau-russell1974">{{cite journal \|last1=Sigoneau-Russell \|first1=D. \|author2=Russel, D.E. \|year=1974 \|title=Étude du premier caséidé (Reptilia, Pelycosauria) d'Europe occidentale \|journal=Bulletin du Muséum National d'Histoire Naturelle \|series=Série 3 \|volume=38 \|issue=230 \|pages=145–215 \|doi= \|url=http://bibliotheques.mnhn.fr/EXPLOITATION/infodoc/digitalCollections/viewerpopup.aspx?seid=BMSCT_S003_1974_T230_N038}}</ref> ''[[Angelosaurus]]'', one of the most derived caseids, has teeth with 5 cuspules.<ref name="Olson1968"/> In ''Angelosaurus'' the teeth have a bulbous morphology with very short and wide crowns. Their sturdiness and the significant wear they show indicate that ''Angelosaurus'' must have fed on tougher plants than those on which most other herbivorous caseids fed.<ref name="Olson1968"/> Herbivorous caseids also show very different dietary adaptations from those seen in another group of basal synapsids, the [[Edaphosauridae]]. The latter had, in addition to the marginal dentition, a dental battery made up of numerous teeth located both on the [[palate]] and on the inner surface of the lower jaws. In herbivorous caseids, the palatal teeth are smaller, and the inner surface of the lower jaws bears no teeth. Instead of a dental battery, they had a massive [[tongue]] (as indicated by the presence of a highly developed [[hyoid apparatus]] found in ''Ennatosaurus'' and ''Euromycter'') perhaps rough, with which they had to compress food against the palatal teeth.<ref name="Olson1968"/><ref name="Reisz&al.2022"/>

	[[File:Arisierpeton simplex, premaxillae.png\|thumb\|right\|Premaxillae of ''[[Arisierpeton\|Arisierpeton simplex]]''. (A) Specimen GAA 00242 in anterior, labial, posterior, ventral, and dorsal views; (B) Specimen GAA 00239 in labial (showing a complete tooth with tricuspid distal end), partial labial, posterior and ventral views.]][[File:Cotylorhynchus romeria from Norman, Oklahoma.jpg\|thumb\|left\|Skeleton of ''Cotylorhynchus romeri'' on display at the [[Sam Noble Oklahoma Museum of Natural History]].]]		[[File:Arisierpeton simplex, premaxillae.png\|thumb\|right\|Premaxillae of ''[[Arisierpeton\|Arisierpeton simplex]]''. (A) Specimen GAA 00242 in anterior, labial, posterior, ventral, and dorsal views; (B) Specimen GAA 00239 in labial (showing a complete tooth with tricuspid distal end), partial labial, posterior and ventral views.]][[File:Cotylorhynchus romeria from Norman, Oklahoma.jpg\|thumb\|left\|Skeleton of ''Cotylorhynchus romeri'' on display at the [[Sam Noble Oklahoma Museum of Natural History]].]]

← Previous revision		Revision as of 22:10, 3 December 2023
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	\|footer=''Cotylorhynchus romeri''. Top : skull reconstruction in left lateral view, and medial and lateral views of mandible; bottom : skull reconstruction in dorsal, ventral, posterior and anterior views.}}[[File:Euromycter rutenus 6778 retouched.jpg\|thumb\|right\|Skull of ''[[Euromycter\|Euromycter rutenus]]'' in ventral view allowing to see the numerous small teeth adorning the bones of the palate (the [[parasphenoid]], triangular in shape in the center, and the very elongated [[Pterygoid bone\|pterygoids]] on each side). The hyoid apparatus is not present here because it was removed during the preparation of the palate.]]		\|footer=''Cotylorhynchus romeri''. Top : skull reconstruction in left lateral view, and medial and lateral views of mandible; bottom : skull reconstruction in dorsal, ventral, posterior and anterior views.}}[[File:Euromycter rutenus 6778 retouched.jpg\|thumb\|right\|Skull of ''[[Euromycter\|Euromycter rutenus]]'' in ventral view allowing to see the numerous small teeth adorning the bones of the palate (the [[parasphenoid]], triangular in shape in the center, and the very elongated [[Pterygoid bone\|pterygoids]] on each side). The hyoid apparatus is not present here because it was removed during the preparation of the palate.]]

	Caseids measured from less than {{convert\|1\|m}} to {{convert\|7\|m}} in length.<ref name="Romano2017">{{cite journal\|last1=Romano \|first1=M. \|year=2017 \|title=Long bone scaling of caseid synapsids: a combined morphometric and cladistic approach \|journal=Lethaia \|volume=50 \|issue=4 \|pages=511–526 \|doi=10.1111/let.12207}}</ref><ref name="Angielczyk&Kammerer2018">{{cite book\|last=Angielczyk\|first=K.D. \|author2=Kammerer, C.F. \|year=2018 \|chapter=Non-Mammalian synapsids : the deep roots of the mammalian family tree \|pages=117–198 \|title=Handbook of Zoology : Mammalian Evolution, Diversity and Systematics \|editor1-last=Zachos\|editor1-first=F.E. \|editor2=Asher, R.J. \|publisher=de Gruyter \|location=Berlin \|isbn=978-3-11-027590-2}}</ref><ref name="Romano&al.2018">{{cite journal\|last1=Romano\|first1=M. \|author2=Citton, P. \|author3=Maganuco, S. \|author4=Sacchi, E. \|author5=Caratelli, M. \|author6=Ronchi, A. \|author7=Nicosia, U. \|year=2018 \|title=New basal synapsid discovery at the Permian outcrop of Torre del Porticciolo (Alghero, Italy) \|journal=Geological Journal \|volume=54 \|issue=3 \|pages=1554–1566 \|doi=10.1002/gj.3250\|s2cid=133755506 }}</ref> They had a small head wider than high and with a forward-inclined snout, a very short neck, a long tail, robust forelimbs, and a body of variable proportions depending on their diet. Small insectivorous species like ''[[Eocasea]]'' had an unexpanded trunk.<ref name="Reisz&Fröbisch2014">{{cite journal\|last1=Reisz\|first1=R.R. \|author2=Fröbisch, J. \|year=2014 \|title=Oldest caseid synapsid from the late Pennsylvanian of Kansas, and the evolution of herbivory in terrestrial vertebrates \|journal=PLOS ONE \|volume=9 \|issue=4 \|pages=e94518 \|doi=10.1371/journal.pone.0094518 \|pmid=24739998 \|pmc=3989228\|bibcode=2014PLoSO...994518R \|doi-access=free }}</ref> Others with an omnivorous diet like ''[[Martensius]]'' had a barely enlarged rib cage, a more elongated skull, smaller [[nostril]]s, and a snout less inclined forward than in herbivorous caseids.<ref name="Berman&al.2020">{{cite journal\|last1=Berman \|first1=D.S. \|author2=Maddin, H.C. \|author3=Henrici, A.C. \|author4=Sumida, S.S. \|author5=Scott, D. \|author6=Reisz, R.R. \|title=New primitive Caseid (Synapsida, Caseasauria) from the Early Permian of Germany \|journal=Annals of Carnegie Museum \|year=2020 \|volume=86 \|issue=1 \|pages=43–75 \|doi=10.2992/007.086.0103\|s2cid=216027787 }}</ref> The latter were characterized by their disproportionately small [[skull]] compared to the size of the body. The postcranial skeleton indeed shows a spectacular increase in the volume of the [[rib cage]], which becomes very wide and barrel-shaped, probably to accommodate a particularly developed [[Gastrointestinal tract#Lower gastrointestinal tract\|intestine]], necessary for the digestion of high-fiber rich plants. In these forms, the skull has very large external nostrils and a very short facial region with a strong forward inclination of the end of the snout which clearly overhangs the dental row. The [[Skull#Fenestrae\|temporal fenestrae]] are also relatively large (especially in ''[[Ennatosaurus]]''), the [[Supratemporal bone\|supratemporals]] are large in size, and, on the [[Occipital bone\|occipital]] surface, the paroccipital [[Process (anatomy)\|processes]] are massively developed, establishing strong supporting contacts with the [[Squamosal bone\|squamosals]].<ref name="Kemp1982">{{cite book\|last=Kemp \|first=T.S. \|year=1982 \|chapter=Pelycosaurs \|pages=39 \|title=Mammal-like reptiles and the origin of Mammals \|editor1-last=Kemp\|editor1-first=T.S. \|publisher=Academic Press \|location=London \|isbn=978-0124041202}}</ref> The dorsal surface of the skull is covered with numerous small pits. These suggest the presence of large [[Reptile scale\|scales]] on the head of these animals.<ref name="Romano&al.2017a">{{cite journal\|last1=Romano \|first1=M. \|author2=Brocklehurst, N. \|author3=Fröbisch, J. \|year=2017 \|title=The postcranial skeleton of ''Ennatosaurus tecton'' (Synapsida, Caseidae) \|journal=Journal of Systematic Palaeontology \|volume=16 \|issue= 13\|pages=1097–1122 \|doi=10.1080/14772019.2017.1367729\|s2cid=89922565 }}</ref> Numerous [[Lip\|labial]] [[Foramen\|foramina]] running parallel to the ventral edge of the [[premaxilla]] and [[maxilla]], as well as along the dorsal edge of the [[Mandible#Other vertebrates\|dentary]], suggest the presence of scaly "lips" which must have concealed the dentition when the jaws were closed.<ref name="Reisz&al.2022">{{cite journal\|last1=Reisz \|first1=R.R. \|author2=Scott, D. \|author3=Modesto, S.P. \|year=2022 \|title=Cranial Anatomy of the Caseid Synapsid ''Cotylorhynchus romeri'', a Large Terrestrial Herbivore From the Lower Permian of Oklahoma, U.S.A \|journal=Frontiers in Earth Science \|volume=10 \|issue= \|pages=1–19 \|doi=10.3389/feart.2022.847560\|doi-access=free }}</ref><ref name="Romano&al.2017a"/> The [[Tooth\|teeth]], simply conical and pointed in insectivorous species, adopt in herbivorous species a leafy or spatulate morphology and are provided with more or less numerous cuspules. Numerous small teeth also adorned several bones of the palate. Herbivorous species do not show a simple evolutionary trend towards increasing tooth complexity.<ref name="Maddin&al.2008">{{cite journal \|last1=Maddin \|first1=H.C. \|author2=Sidor, C.A. \|author3=Reisz, R.R. \|year=2008 \|title=Cranial anatomy of ''Ennatosaurus tecton'' (Synapsida: Caseidae) from the Middle Permian of Russia and the evolutionary relationships of Caseidae \|journal=Journal of Vertebrate Paleontology \|volume=28 \|issue=1 \|pages=160–180 \|doi=10.1671/0272-4634(2008)28[160:CAOETS]2.0.CO;2\|s2cid=44064927 }}</ref> Thus, the teeth of the basal taxa ''[[Casea]]'' and ''[[Arisierpeton]]'' have three cuspules<ref name="Olson1968">{{cite journal\|last1=Olson \|~~first~~=E.C. \|year=1968 \|title=The family Caseidae \|journal=Fieldiana: Geology \|volume=17 \|issue= \|pages=225–349\| doi= }}</ref><ref name="Reisz2019">{{cite journal \|last1=Reisz \|first1=R. \|year=2019 \|title=A small caseid synapsid, ''Arisierpeton simplex'' gen. et sp. nov., from the early Permian of Oklahoma, with a discussion of synapsid diversity at the classic Richards Spur locality \|journal=PeerJ \|volume=7e6615 \|issue= \|pages=1–20 \|doi=10.7717/peerj.6615 \|pmid=30997285 \|pmc=6462398 \|~~url~~=~~http://doi.org/10.7717/peerj.6615~~}}</ref> just like in the more derived forms ''[[Cotylorhynchus]]'' and ''[[Caseopsis]]''.<ref name="Olson1968"/> ''Ennatosaurus'' and ''[[Euromycter]]'', which occupy an intermediate [[Phylogenetics\|phylogenetic]] position, have teeth bearing 5 to 7 cuspules and 5 to 8 cuspules respectively.<ref name="Maddin&al.2008"/><ref name="sigogneau-russell1974">{{cite journal \|last1=Sigoneau-Russell \|first1=D. \|author2=Russel, D.E. \|year=1974 \|title=Étude du premier caséidé (Reptilia, Pelycosauria) d'Europe occidentale \|journal=Bulletin du Muséum National d'Histoire Naturelle \|series=Série 3 \|volume=38 \|issue=230 \|pages=145–215 \|doi= \|url=http://bibliotheques.mnhn.fr/EXPLOITATION/infodoc/digitalCollections/viewerpopup.aspx?seid=BMSCT_S003_1974_T230_N038}}</ref> ''[[Angelosaurus]]'', one of the most derived caseids, has teeth with 5 cuspules.<ref name="Olson1968"/> In ''Angelosaurus'' the teeth have a bulbous morphology with very short and wide crowns. Their sturdiness and the significant wear they show, indicate that ''Angelosaurus'' must have fed on tougher plants than those on which most other herbivorous caseids fed.<ref name="Olson1968"/> Herbivorous caseids also show very different dietary adaptations from those seen in another group of basal synapsids, the [[Edaphosauridae]]. The latter had, in addition to the marginal dentition, a dental battery made up of numerous teeth located both on the [[palate]] and on the inner surface of the lower jaws. In herbivorous caseids, the palatal teeth are smaller, and the inner surface of the lower jaws bears no teeth. Instead of a dental battery, they had a massive [[tongue]] (as indicated by the presence of a highly developed [[hyoid apparatus]] found in ''Ennatosaurus'' and ''Euromycter'') perhaps rough, with which they had to compress food against the palatal teeth.<ref name="Olson1968"/><ref name="Reisz&al.2022"/>		Caseids measured from less than {{convert\|1\|m}} to {{convert\|7\|m}} in length.<ref name="Romano2017">{{cite journal\|last1=Romano \|first1=M. \|year=2017 \|title=Long bone scaling of caseid synapsids: a combined morphometric and cladistic approach \|journal=Lethaia \|volume=50 \|issue=4 \|pages=511–526 \|doi=10.1111/let.12207}}</ref><ref name="Angielczyk&Kammerer2018">{{cite book\|last=Angielczyk\|first=K.D. \|author2=Kammerer, C.F. \|year=2018 \|chapter=Non-Mammalian synapsids : the deep roots of the mammalian family tree \|pages=117–198 \|title=Handbook of Zoology : Mammalian Evolution, Diversity and Systematics \|editor1-last=Zachos\|editor1-first=F.E. \|editor2=Asher, R.J. \|publisher=de Gruyter \|location=Berlin \|isbn=978-3-11-027590-2}}</ref><ref name="Romano&al.2018">{{cite journal\|last1=Romano\|first1=M. \|author2=Citton, P. \|author3=Maganuco, S. \|author4=Sacchi, E. \|author5=Caratelli, M. \|author6=Ronchi, A. \|author7=Nicosia, U. \|year=2018 \|title=New basal synapsid discovery at the Permian outcrop of Torre del Porticciolo (Alghero, Italy) \|journal=Geological Journal \|volume=54 \|issue=3 \|pages=1554–1566 \|doi=10.1002/gj.3250\|s2cid=133755506 }}</ref> They had a small head wider than high and with a forward-inclined snout, a very short neck, a long tail, robust forelimbs, and a body of variable proportions depending on their diet. Small insectivorous species like ''[[Eocasea]]'' had an unexpanded trunk.<ref name="Reisz&Fröbisch2014">{{cite journal\|last1=Reisz\|first1=R.R. \|author2=Fröbisch, J. \|year=2014 \|title=Oldest caseid synapsid from the late Pennsylvanian of Kansas, and the evolution of herbivory in terrestrial vertebrates \|journal=PLOS ONE \|volume=9 \|issue=4 \|pages=e94518 \|doi=10.1371/journal.pone.0094518 \|pmid=24739998 \|pmc=3989228\|bibcode=2014PLoSO...994518R \|doi-access=free }}</ref> Others with an omnivorous diet like ''[[Martensius]]'' had a barely enlarged rib cage, a more elongated skull, smaller [[nostril]]s, and a snout less inclined forward than in herbivorous caseids.<ref name="Berman&al.2020">{{cite journal\|last1=Berman \|first1=D.S. \|author2=Maddin, H.C. \|author3=Henrici, A.C. \|author4=Sumida, S.S. \|author5=Scott, D. \|author6=Reisz, R.R. \|title=New primitive Caseid (Synapsida, Caseasauria) from the Early Permian of Germany \|journal=Annals of Carnegie Museum \|year=2020 \|volume=86 \|issue=1 \|pages=43–75 \|doi=10.2992/007.086.0103\|s2cid=216027787 }}</ref> The latter were characterized by their disproportionately small [[skull]] compared to the size of the body. The postcranial skeleton indeed shows a spectacular increase in the volume of the [[rib cage]], which becomes very wide and barrel-shaped, probably to accommodate a particularly developed [[Gastrointestinal tract#Lower gastrointestinal tract\|intestine]], necessary for the digestion of high-fiber rich plants. In these forms, the skull has very large external nostrils and a very short facial region with a strong forward inclination of the end of the snout which clearly overhangs the dental row. The [[Skull#Fenestrae\|temporal fenestrae]] are also relatively large (especially in ''[[Ennatosaurus]]''), the [[Supratemporal bone\|supratemporals]] are large in size, and, on the [[Occipital bone\|occipital]] surface, the paroccipital [[Process (anatomy)\|processes]] are massively developed, establishing strong supporting contacts with the [[Squamosal bone\|squamosals]].<ref name="Kemp1982">{{cite book\|last=Kemp \|first=T.S. \|year=1982 \|chapter=Pelycosaurs \|pages=39 \|title=Mammal-like reptiles and the origin of Mammals \|editor1-last=Kemp\|editor1-first=T.S. \|publisher=Academic Press \|location=London \|isbn=978-0124041202}}</ref> The dorsal surface of the skull is covered with numerous small pits. These suggest the presence of large [[Reptile scale\|scales]] on the head of these animals.<ref name="Romano&al.2017a">{{cite journal\|last1=Romano \|first1=M. \|author2=Brocklehurst, N. \|author3=Fröbisch, J. \|year=2017 \|title=The postcranial skeleton of ''Ennatosaurus tecton'' (Synapsida, Caseidae) \|journal=Journal of Systematic Palaeontology \|volume=16 \|issue= 13\|pages=1097–1122 \|doi=10.1080/14772019.2017.1367729\|s2cid=89922565 }}</ref> Numerous [[Lip\|labial]] [[Foramen\|foramina]] running parallel to the ventral edge of the [[premaxilla]] and [[maxilla]], as well as along the dorsal edge of the [[Mandible#Other vertebrates\|dentary]], suggest the presence of scaly "lips" which must have concealed the dentition when the jaws were closed.<ref name="Reisz&al.2022">{{cite journal\|last1=Reisz \|first1=R.R. \|author2=Scott, D. \|author3=Modesto, S.P. \|year=2022 \|title=Cranial Anatomy of the Caseid Synapsid ''Cotylorhynchus romeri'', a Large Terrestrial Herbivore From the Lower Permian of Oklahoma, U.S.A \|journal=Frontiers in Earth Science \|volume=10 \|issue= \|pages=1–19 \|doi=10.3389/feart.2022.847560\|doi-access=free }}</ref><ref name="Romano&al.2017a"/> The [[Tooth\|teeth]], simply conical and pointed in insectivorous species, adopt in herbivorous species a leafy or spatulate morphology and are provided with more or less numerous cuspules. Numerous small teeth also adorned several bones of the palate. Herbivorous species do not show a simple evolutionary trend towards increasing tooth complexity.<ref name="Maddin&al.2008">{{cite journal \|last1=Maddin \|first1=H.C. \|author2=Sidor, C.A. \|author3=Reisz, R.R. \|year=2008 \|title=Cranial anatomy of ''Ennatosaurus tecton'' (Synapsida: Caseidae) from the Middle Permian of Russia and the evolutionary relationships of Caseidae \|journal=Journal of Vertebrate Paleontology \|volume=28 \|issue=1 \|pages=160–180 \|doi=10.1671/0272-4634(2008)28[160:CAOETS]2.0.CO;2\|s2cid=44064927 }}</ref> Thus, the teeth of the basal taxa ''[[Casea]]'' and ''[[Arisierpeton]]'' have three cuspules<ref name="Olson1968">{{cite journal\|last1=Olson \|first1=E.C. \|year=1968 \|title=The family Caseidae \|journal=Fieldiana: Geology \|volume=17 \|issue= \|pages=225–349\| doi= }}</ref><ref name="Reisz2019">{{cite journal \|last1=Reisz \|first1=R. \|year=2019 \|title=A small caseid synapsid, ''Arisierpeton simplex'' gen. et sp. nov., from the early Permian of Oklahoma, with a discussion of synapsid diversity at the classic Richards Spur locality \|journal=PeerJ \|volume=7e6615 \|issue= \|pages=1–20 \|doi=10.7717/peerj.6615 \|pmid=30997285 \|pmc=6462398 \|doi-access=free }}</ref> just like in the more derived forms ''[[Cotylorhynchus]]'' and ''[[Caseopsis]]''.<ref name="Olson1968"/> ''Ennatosaurus'' and ''[[Euromycter]]'', which occupy an intermediate [[Phylogenetics\|phylogenetic]] position, have teeth bearing 5 to 7 cuspules and 5 to 8 cuspules respectively.<ref name="Maddin&al.2008"/><ref name="sigogneau-russell1974">{{cite journal \|last1=Sigoneau-Russell \|first1=D. \|author2=Russel, D.E. \|year=1974 \|title=Étude du premier caséidé (Reptilia, Pelycosauria) d'Europe occidentale \|journal=Bulletin du Muséum National d'Histoire Naturelle \|series=Série 3 \|volume=38 \|issue=230 \|pages=145–215 \|doi= \|url=http://bibliotheques.mnhn.fr/EXPLOITATION/infodoc/digitalCollections/viewerpopup.aspx?seid=BMSCT_S003_1974_T230_N038}}</ref> ''[[Angelosaurus]]'', one of the most derived caseids, has teeth with 5 cuspules.<ref name="Olson1968"/> In ''Angelosaurus'' the teeth have a bulbous morphology with very short and wide crowns. Their sturdiness and the significant wear they show, indicate that ''Angelosaurus'' must have fed on tougher plants than those on which most other herbivorous caseids fed.<ref name="Olson1968"/> Herbivorous caseids also show very different dietary adaptations from those seen in another group of basal synapsids, the [[Edaphosauridae]]. The latter had, in addition to the marginal dentition, a dental battery made up of numerous teeth located both on the [[palate]] and on the inner surface of the lower jaws. In herbivorous caseids, the palatal teeth are smaller, and the inner surface of the lower jaws bears no teeth. Instead of a dental battery, they had a massive [[tongue]] (as indicated by the presence of a highly developed [[hyoid apparatus]] found in ''Ennatosaurus'' and ''Euromycter'') perhaps rough, with which they had to compress food against the palatal teeth.<ref name="Olson1968"/><ref name="Reisz&al.2022"/>

	[[File:Arisierpeton simplex, premaxillae.png\|thumb\|right\|Premaxillae of ''[[Arisierpeton\|Arisierpeton simplex]]''. (A) Specimen GAA 00242 in anterior, labial, posterior, ventral, and dorsal views; (B) Specimen GAA 00239 in labial (showing a complete tooth with tricuspid distal end), partial labial, posterior and ventral views.]][[File:Cotylorhynchus romeria from Norman, Oklahoma.jpg\|thumb\|left\|Skeleton of ''Cotylorhynchus romeri'' on display at the [[Sam Noble Oklahoma Museum of Natural History]].]]		[[File:Arisierpeton simplex, premaxillae.png\|thumb\|right\|Premaxillae of ''[[Arisierpeton\|Arisierpeton simplex]]''. (A) Specimen GAA 00242 in anterior, labial, posterior, ventral, and dorsal views; (B) Specimen GAA 00239 in labial (showing a complete tooth with tricuspid distal end), partial labial, posterior and ventral views.]][[File:Cotylorhynchus romeria from Norman, Oklahoma.jpg\|thumb\|left\|Skeleton of ''Cotylorhynchus romeri'' on display at the [[Sam Noble Oklahoma Museum of Natural History]].]]
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	This hypothesis is however disputed by Kenneth Angielczyk and Christian Kammerer as well as by Robert Reisz and colleagues based on [[Paleontology\|paleontological]] and [[Taphonomy\|taphonomic]] data combined with the absence in these large caseids of morphological adaptations to an aquatic lifestyle. According to Angielczyk and Kammerer, the low bone density of caseids identified by Lambertz et al. does not resemble that of semiaquatic animals, which tend to have a more strongly ossified skeleton to provide passive [[buoyancy]] control and increased stability against current and wave action. ''Cotylorhynchus'' bone microstructure is more similar to what is seen in animals living in the [[Pelagic zone\|open ocean]], such as [[cetacea]]ns and [[pinniped]]s, which emphasize high maneuverability, rapid acceleration and hydrodynamic control of buoyancy. However, the caseid morphology was totally incompatible with a [[Pelagic zone\|pelagic]] lifestyle. Thus, due to these unusual data, Angielczyk and Kammerer consider that the available evidence is still insufficient to question the more widely assumed terrestrial lifestyle of caseids.<ref name="Angielczyk&Kammerer2018"/> Robert Reisz and colleagues also dispute the supposed semiaquatic lifestyle of the caseids on the fact that the latter possess no morphological adaptations to an aquatic lifestyle and, in the case of the species ''Cotylorhynchus romeri'', on the interpretation that this animal lived in a dry environment for part of the year as indicated by the presence of numerous skeletons of the amphibian ''[[Brachydectes]]'' preserved in [[aestivation]] and of the [[lungfish]] ''[[Gnathorhiza]]'', another well-known aestivator.<ref name="Reisz&al.2022"/>		This hypothesis is however disputed by Kenneth Angielczyk and Christian Kammerer as well as by Robert Reisz and colleagues based on [[Paleontology\|paleontological]] and [[Taphonomy\|taphonomic]] data combined with the absence in these large caseids of morphological adaptations to an aquatic lifestyle. According to Angielczyk and Kammerer, the low bone density of caseids identified by Lambertz et al. does not resemble that of semiaquatic animals, which tend to have a more strongly ossified skeleton to provide passive [[buoyancy]] control and increased stability against current and wave action. ''Cotylorhynchus'' bone microstructure is more similar to what is seen in animals living in the [[Pelagic zone\|open ocean]], such as [[cetacea]]ns and [[pinniped]]s, which emphasize high maneuverability, rapid acceleration and hydrodynamic control of buoyancy. However, the caseid morphology was totally incompatible with a [[Pelagic zone\|pelagic]] lifestyle. Thus, due to these unusual data, Angielczyk and Kammerer consider that the available evidence is still insufficient to question the more widely assumed terrestrial lifestyle of caseids.<ref name="Angielczyk&Kammerer2018"/> Robert Reisz and colleagues also dispute the supposed semiaquatic lifestyle of the caseids on the fact that the latter possess no morphological adaptations to an aquatic lifestyle and, in the case of the species ''Cotylorhynchus romeri'', on the interpretation that this animal lived in a dry environment for part of the year as indicated by the presence of numerous skeletons of the amphibian ''[[Brachydectes]]'' preserved in [[aestivation]] and of the [[lungfish]] ''[[Gnathorhiza]]'', another well-known aestivator.<ref name="Reisz&al.2022"/>

	In 2022, Werneburg and colleagues proposed a somewhat different semiaquatic lifestyle, in which large caseids like ''[[Lalieudorhynchus]]'' (whose bone texture is even more osteoporotic than in ''Cotylorhynchus'') would be ecological equivalents of modern [[Hippopotamus\|hippos]], passing part of their time in the water (being underwater walkers rather than swimming animals) but coming on dry land for food.<ref name="Werneburg&al.2022">{{cite journal\|last1=Werneburg \|~~first~~=R. \|author2=Spindler, F. \|author3=Falconnet, J. \|author4=Steyer, J.-S. \|author5=Vianey-Liaud, M. \|author6=Schneider, J.W. \|title=A new caseid synapsid from the Permian (Guadalupian) of the Lodève basin (Occitanie, France) \|journal=Palaeovertebrata \|year=2022 \|volume=45(2)-e2 \|issue= 2\|pages=e2 \|doi=10.18563/pv.45.2.e2\|s2cid=253542331 \|url=https://hal.archives-ouvertes.fr/hal-03851911/file/Werneburg%20et%20al%202022%20A%20new%20caseid%20synapsid%20from%20the%20Permian%20of%20the%20Lod%C3%A8ve%20Basin.pdf }}</ref>		In 2022, Werneburg and colleagues proposed a somewhat different semiaquatic lifestyle, in which large caseids like ''[[Lalieudorhynchus]]'' (whose bone texture is even more osteoporotic than in ''Cotylorhynchus'') would be ecological equivalents of modern [[Hippopotamus\|hippos]], passing part of their time in the water (being underwater walkers rather than swimming animals) but coming on dry land for food.<ref name="Werneburg&al.2022">{{cite journal\|last1=Werneburg \|first1=R. \|author2=Spindler, F. \|author3=Falconnet, J. \|author4=Steyer, J.-S. \|author5=Vianey-Liaud, M. \|author6=Schneider, J.W. \|title=A new caseid synapsid from the Permian (Guadalupian) of the Lodève basin (Occitanie, France) \|journal=Palaeovertebrata \|year=2022 \|volume=45(2)-e2 \|issue= 2\|pages=e2 \|doi=10.18563/pv.45.2.e2\|s2cid=253542331 \|url=https://hal.archives-ouvertes.fr/hal-03851911/file/Werneburg%20et%20al%202022%20A%20new%20caseid%20synapsid%20from%20the%20Permian%20of%20the%20Lod%C3%A8ve%20Basin.pdf }}</ref>

	==Evolution==		==Evolution==