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{{Short description|Order of extinct placental mammals}}
{{automatic Taxobox
{{Automatic taxobox
| name = Embrithopoda
| fossil_range= [[Eocene]] - [[Oligocene]], {{fossilrange|56|28|ref=<ref>Emmanuel Gheerbrant; Arnaud Schmitt; László Kocsis (2018). "Early African fossils elucidate the origin of embrithopod mammals". Current Biology. Online edition. doi:10.1016/j.cub.2018.05.032.</ref>}}
| fossil_range = [[Paleocene]] - [[Oligocene]], {{fossilrange|56|28|ref=<ref>Emmanuel Gheerbrant; Arnaud Schmitt; László Kocsis (2018). "Early African fossils elucidate the origin of embrithopod mammals". Current Biology. Online edition. doi:10.1016/j.cub.2018.05.032.</ref>}}
| image = Arsinoitherium zitteli.jpg
| image = Arsinoitherium zitteli.jpg
| image_caption = ''[[Arsinoitherium zitteli]]''<ref name="Rose-242">{{Harvnb|Rose|2006| pp=242–3}}</ref>
| image_caption = ''[[Arsinoitherium zitteli]]''<ref name="Rose-242">{{Harvnb|Rose|2006| pp=242–3}}</ref>
| display_parents = 3
| taxon = Embrithopoda
| authority = {{Harvnb|Andrews|1906}}
| authority = {{Harvnb|Andrews|1906}}
| subdivision_ranks = [[Family (biology)|Families]]
| subdivision_ranks = Families
| subdivision =
| subdivision = * †''[[Stylolophus]]''
* †[[Arsinoitheriidae]]
* †[[Arsinoitheriidae]]
* †[[Palaeoamasiidae]]
* †[[Palaeoamasiidae]]
}}
}}


'''Embrithopoda''' ("heavy-footed") is an [[Order (biology)|order]] of extinct [[mammal]]s known from Asia, Africa and eastern Europe. Most of the embrithopod genera are known exclusively from jaws and teeth dated from the late [[Paleocene]] to the late [[Eocene]], but the order is best known from its terminal member, the elephantine ''[[Arsinoitherium]]''.<ref name="Rose-2006">{{Harvnb|Rose|2006| p=265}}</ref>
'''Embrithopoda''' ("heavy-footed") is an [[Order (biology)|order]] of extinct [[mammal]]s known from Asia, Africa and [[Eastern Europe]]. Most of the embrithopod genera are known exclusively from jaws and teeth dated from the late [[Paleocene]] to the late [[Eocene]]; however, the order is best known from its terminal member, the elephantine ''[[Arsinoitherium]]''.<ref name="Rose-2006">{{Harvnb|Rose|2006| p=265}}</ref>


==Description==
==Description==
While embrithopods bore a superficial resemblance to [[rhinoceros]]es, their horns had bony cores covered in [[keratin]]ized skin and were not made of [[hair]]. Not all embrithopods possessed horns, either. Despite their appearance, they have been regarded as related to [[elephant]]s, not [[perissodactyl]]s.<ref name="UCMP">{{Cite web
While embrithopods bore a superficial resemblance to [[rhinoceros]]es, their horns had bony cores covered in [[keratin]]ized skin. Not all embrithopods possessed horns, either. Despite their appearance, they have been regarded as related to [[elephant]]s, not [[perissodactyl]]s.<ref name="UCMP">{{Cite web
| title = Introduction to the Embrithopoda
| title = Introduction to the Embrithopoda
| publisher = University of California Museum of Paleontology
| publisher = University of California Museum of Paleontology
| url = http://www.ucmp.berkeley.edu/mammal/mesaxonia/embrithopoda.html
| url = http://www.ucmp.berkeley.edu/mammal/mesaxonia/embrithopoda.html
| accessdate = March 2013}}</ref>
| access-date = 7 March 2013}}</ref>


As [[Tethytheria|tethytheres]],<ref>{{Paleodb|43309|Embrithopoda}}. Retrieved March 2013.</ref> the Embrithopoda have been believed to be part of the clade [[Afrotheria]]. However, a study of the basal arsinoitheriid, ''Palaeoamasia'', suggests that embrithopods are not [[Tethytheria|tethytheres]] or even [[Paenungulata|paenungulates]], and that they need to be better sampled in an analysis of [[eutheria]]n relationships to clarify if they are even [[afrotheria]]ns.<ref name="Erdal2016">{{cite journal|last1= Erdal|first1= O.|last2= Antoine|first2= P.-O.|last3= Sen|first3= S.|last4= Smith|first4= A.|title= New material of ''Palaeoamasia kansui'' (Embrithopoda, Mammalia) from the Eocene of Turkey and a phylogenetic analysis of Embrithopoda at the species level|journal= Palaeontology|volume= 59|issue= 5|year= 2016|pages= 631–655|doi= 10.1111/pala.12247}}[Erdal et al.'s inclusion of Embrithopoda in Tabuce et al. (2007) finds them outside of (Ungulata + Afrotheria). Since this clade is not supported by molecular data, it suggests the need to explore the relationships of embrithopods, as they could potentially have affinities with laurasiatheres and "true" ungulates.]</ref> It is also not clear if embrithopods originated in Africa or Eurasia.<ref name="Erdal2016" /> However, recent findings demonstrate an african origin for embrithopods and furthermore a relationship with other paenungulates, albeit having diverged earlier than previously thought.<ref>E. Gheerbrant, A. Schmitt, and L. Kocsis. 2018. Early African Fossils Elucidate the Origin of Embrithopod Mammals. Current Biology 28:1-7</ref>
As [[Tethytheria|tethytheres]],<ref>{{Paleodb|43309|Embrithopoda}}. Retrieved March 2013.</ref> the Embrithopoda have been believed to be part of the clade [[Afrotheria]]. However, a study of the basal arsinoitheriid, ''Palaeoamasia'', suggests that embrithopods are not [[Tethytheria|tethytheres]] or even [[Paenungulata|paenungulates]], and that they need to be better sampled in an analysis of [[eutheria]]n relationships to clarify if they are even [[afrotheria]]ns.<ref name="Erdal2016">{{cite journal|last1= Erdal|first1= O.|last2= Antoine|first2= P.-O.|last3= Sen|first3= S.|last4= Smith|first4= A.|title= New material of ''Palaeoamasia kansui'' (Embrithopoda, Mammalia) from the Eocene of Turkey and a phylogenetic analysis of Embrithopoda at the species level|journal= Palaeontology|volume= 59|issue= 5|year= 2016|pages= 631–655|doi= 10.1111/pala.12247|url= https://hal.sorbonne-universite.fr/hal-01346066/document}}[Erdal et al.'s inclusion of Embrithopoda in Tabuce et al. (2007) finds them outside of (Ungulata + Afrotheria). Since this clade is not supported by molecular data, it suggests the need to explore the relationships of embrithopods, as they could potentially have affinities with laurasiatheres and "true" ungulates.]</ref> It is also not clear if embrithopods originated in Africa or Eurasia.<ref name="Erdal2016" /> However, recent findings demonstrate an African origin for embrithopods and furthermore a relationship with other paenungulates, albeit having diverged earlier than previously thought.<ref name="E. Gheerbrant, A. Schmitt 2018">{{cite journal |last1= Gheerbrant |first1= E. |last2= Schmitt |first2= A. |last3= Kocsis |first3= L. |date= July 2018 |title= Early African fossils elucidate the origin of embrithopod mammals |journal= Current Biology |volume= 28 |issue= 13 |pages= 2167-2173.e2 |url= https://www.cell.com/current-biology/fulltext/S0960-9822(18)30668-7 |access-date= 28 September 2024}}</ref>


Fossils of embrithopods, such as ''[[Arsinoitherium]]'', have been found in [[Egypt]], [[Mongolia]], [[Turkey]], [[Romania]], [[Namibia]]<ref name=Namatherium>{{Cite journal |author1=M. Pickford |author2=B. Senut |author3=J. Morales |author4=P. Mein |author5=I. M. Sanchez | year = 2008 | journal = Memoir of the Geological Survey of Namibia | volume = 20 | pages = 465–514 | title = Mammalia from the Lutetian of Namibia }}</ref> and [[Tunisia]].<ref>{{Cite journal |author1=Nicolas Vialle |author2=Gilles Merzeraud |author3=Cyrille Delmer |author4=Monique Feist |author5=Suzanne Jiquel |author6=Laurent Marivaux |author7=Anusha Ramdarshan |author8=Monique Vianey-Liaud |author9=El Mabrouk Essid |author10=Wissem Marzougui |author11=Hayet Khayati Ammar |author12=Rodolphe Tabuce | year = 2013 | journal = Journal of African Earth Sciences | volume = 87 | pages = 86–92 | title = Discovery of an embrithopod mammal (''Arsinoitherium''?) in the late Eocene of Tunisia | doi=10.1016/j.jafrearsci.2013.07.010 }}</ref> Until the 1970s, only ''Arsinoitherium'' itself was known, appearing isolated in the fossil record.<ref name="UCMP" />
Fossils of embrithopods, such as ''[[Arsinoitherium]]'', have been found in [[Egypt]], [[Ethiopia]], [[Kenya]], [[Morocco]], [[Mongolia]], [[Turkey]], [[Romania]], [[Namibia]],<ref name=Namatherium>{{Cite journal |author1=M. Pickford |author2=B. Senut |author3=J. Morales |author4=P. Mein |author5=I. M. Sanchez | year = 2008 | journal = Memoir of the Geological Survey of Namibia | volume = 20 | pages = 465–514 | title = Mammalia from the Lutetian of Namibia }}</ref> [[Tunisia]]<ref>{{Cite journal |author1=Nicolas Vialle |author2=Gilles Merzeraud |author3=Cyrille Delmer |author4=Monique Feist |author5=Suzanne Jiquel |author6=Laurent Marivaux |author7=Anusha Ramdarshan |author8=Monique Vianey-Liaud |author9=El Mabrouk Essid |author10=Wissem Marzougui |author11=Hayet Khayati Ammar |author12=Rodolphe Tabuce | year = 2013 | journal = Journal of African Earth Sciences | volume = 87 | pages = 86–92 | title = Discovery of an embrithopod mammal (''Arsinoitherium''?) in the late Eocene of Tunisia | doi=10.1016/j.jafrearsci.2013.07.010 }}</ref> and [[Croatia]].<ref>[https://www.rabdanas.com/index.php/vijesti/item/7241-dan-georaznolikosti-na-rabu-obiljezen-medunarodnim-simpozijem-geopark-and-sciences/ Rab danas]</ref> Until the 1970s, only ''Arsinoitherium'' itself was known, appearing isolated in the fossil record.<ref name="UCMP" />


==Classification==
==Classification==
{{Harvnb|McKenna|Manning|1977}} and {{Harvnb|McKenna|Bell|1997}} considered ''[[Phenacolophus]]'' from Mongolia a primitive embrithopod, although this attribution was challenged by several other authors.<ref>{{Cite journal|last=Koenigswald|first=W. v.|date=2012|title=Unique differentiation of radial enamel in Arsinoitherium (Embrithopoda, Tethytheria)|url=http://s3.amazonaws.com/academia.edu.documents/32095413/205_-_Koenigswald__2012_Arsinoitherium_enamel.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1496545134&Signature=lfX%2FPhwuQs%2B4oz%2F5mtUgH4NPb8k%3D&response-content-disposition=inline%3B%20filename%3DUnique_differentiation_of_radial_enamel.pdf|journal=Historical Biology|volume=|pages=|via=|doi=10.1080/08912963.2012.714658}}</ref> A 2016 cladistic study found ''Phenacolophus'' as a stem-perissodactyl and the embrithopods at the base of [[Altungulata]].<ref>{{Harvnb|Radulesco|Sudre|1985}}; {{Harvnb|Maas|Thewissen|Kappelman|1998|p=291}}; {{Harvnb|Rose|Archibald|2005|pp=97–98}}</ref><ref name="Erdal2016" />
{{Harvnb|McKenna|Manning|1977}} and {{Harvnb|McKenna|Bell|1997}} considered ''Phenacolophus'' from Mongolia a primitive embrithopod, although this attribution was challenged by several other authors.<ref>{{Cite journal|last=Koenigswald|first=W. v.|date=2012|title=Unique differentiation of radial enamel in Arsinoitherium (Embrithopoda, Tethytheria)|journal=Historical Biology|volume=25|issue=2|pages=183–192|doi=10.1080/08912963.2012.714658}}</ref> A 2016 cladistic study found ''Phenacolophus'' as a stem-perissodactyl and the embrithopods at the base of [[Altungulata]].<ref>{{Harvnb|Radulesco|Sudre|1985}}; {{Harvnb|Maas|Thewissen|Kappelman|1998|p=291}}; {{Harvnb|Rose|Archibald|2005|pp=97–98}}</ref><ref name="Erdal2016" /> More recently, an afrothere identity has been vindicated, albeit more basal than previously assumed.<ref name="E. Gheerbrant, A. Schmitt 2018"/>


Order '''Embrithopoda''' <small>{{Harvnb|Andrews|1906}} sensu Prothero & Schoch 1989 </small> (=Barypoda <small>Andrews 1904</small>)<ref name="mikko">[[Mikko's Phylogeny Archive]] [http://www.helsinki.fi/~mhaaramo/] {{cite web|last=Haaramo|first=Mikko|year=2007|title=''†Embrithopoda - arsinoitheres'' |url=http://www.helsinki.fi/~mhaaramo/metazoa/deuterostoma/chordata/synapsida/eutheria/basal_ungulata/embrithopoda.html |accessdate= 30 December 2015}}</ref>
Order '''Embrithopoda''' <small>{{Harvnb|Andrews|1906}} sensu Prothero & Schoch 1989 </small> (=Barypoda <small>Andrews 1904</small>)<ref name="mikko">[[Mikko's Phylogeny Archive]] [http://www.helsinki.fi/~mhaaramo/] {{cite web |last=Haaramo |first=Mikko |year=2007 |title=''†Embrithopoda - arsinoitheres'' |url=http://www.helsinki.fi/~mhaaramo/metazoa/deuterostoma/chordata/synapsida/eutheria/basal_ungulata/embrithopoda.html |access-date=30 December 2015}}</ref>
* Genus †''{{ill|Stylolophus|de}}'' <small>Gheerbrant ''et al'', 2018</small>
* Family †[[Arsinoitheriidae]] <small>{{Harvnb|Andrews|1904}}</small>
* Family †[[Arsinoitheriidae]] <small>{{Harvnb|Andrews|1904}}</small>
** Genus ''[[Namatherium]]'' <small>Pickford ''et al.'', 2008</small><ref name=Namatherium />
** Genus ''{{ill|Namatherium|de}}'' <small>Pickford ''et al.'', 2008</small><ref name=Namatherium />
** Genus †''[[Arsinoitherium]]'' <small>{{Harvnb|Beadnell|1902}}</small>
** Genus †''[[Arsinoitherium]]'' <small>{{Harvnb|Beadnell|1902}}</small>
* Family †[[Palaeoamasiidae]] <small>{{Harvnb|Şen|Heintz|1979}}</small>
* Family †[[Palaeoamasiidae]] <small>{{Harvnb|Şen|Heintz|1979}}</small>
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| title = Further notes on the mammals of the Eocene of Egypt
| title = Further notes on the mammals of the Eocene of Egypt
| year = 1904 | journal = Geological Magazine | volume = 1 | issue = 4
| year = 1904 | journal = Geological Magazine | volume = 1 | issue = 4
| doi = 10.1017/S0016756800119491 | oclc = 4668923377 | ref = harv}}<!-- {{Harvnb|Andrews|1904}} -->
| pages = 157–162 | doi = 10.1017/S0016756800119491 | bibcode = 1904GeoM....1..157A | oclc = 4668923377 | url = https://zenodo.org/record/1687665 }}<!-- {{Harvnb|Andrews|1904}} -->
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| last = Andrews | first = C.W.
| title = A descriptive catalogue of the Tertiary Vertebrata of the Fayûm, Egypt
| title = A descriptive catalogue of the Tertiary Vertebrata of the Fayûm, Egypt
| year = 1906 | location = London | publisher = British Museum
| year = 1906 | location = London | publisher = British Museum
| oclc = 3675777 | ref = harv}}<!-- {{Harvnb|Andrews|1906}} -->
| oclc = 3675777 }}<!-- {{Harvnb|Andrews|1906}} -->
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| last = Beadnell | first = H.J.C.
| last = Beadnell | first = H.J.C.
| title = A preliminary note on Arsinoitherium zitteli Beadnell, from the Upper Eocene strata of Egypt
| title = A preliminary note on Arsinoitherium zitteli Beadnell, from the Upper Eocene strata of Egypt
| year = 1902 | publisher = Egyptian Survey Department, Public Works Ministry | location = Cairo
| year = 1902 | publisher = Egyptian Survey Department, Public Works Ministry | location = Cairo
| oclc = 20609512 | ref = harv}}<!-- {{Harvnb|Beadnell|1902}} --->
| oclc = 20609512 }}<!-- {{Harvnb|Beadnell|1902}} --->
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| last = Court | first = N.
| last = Court | first = N.
| year = 1990 | journal = Journal of Vertebrate Paleontology | volume = 10 | issue = 2 | pages = 170–82
| year = 1990 | journal = Journal of Vertebrate Paleontology | volume = 10 | issue = 2 | pages = 170–82
| title = Periotic anatomy of Arsinoitherium (Mammalia, Embrithopoda) and its phylogenetic implications
| title = Periotic anatomy of Arsinoitherium (Mammalia, Embrithopoda) and its phylogenetic implications
| oclc = 4899524631 | ref = harv | doi=10.1080/02724634.1990.10011806}}<!-- {{Harvnb|Court|1990}} -->
| oclc = 4899524631 | doi=10.1080/02724634.1990.10011806}}<!-- {{Harvnb|Court|1990}} -->
* {{Cite book | last1 = Maas | first1 = M.C. | author2-link = Hans Thewissen | last2 = Thewissen | first2 = J.G.M. | last3 = Kappelman | first3 = J. | chapter = ''Hypsamasia seni'' (Mammalia: Embrithopoda) and other mammals from the Eocene Kartal Formation of Turkey | pages = 286–297 | editor1-last = Beard | editor1-first = K.C. | editor2-last = Dawson | editor2-first = M.R. | title = Dawn of the Age of Mammals in Asia | year = 1998 | series = Bulletin of Carnegie Museum of Natural History | volume = 34 | chapter-url = http://www.neomed.edu/DEPTS/ANAT/Thewissen/pdf/1998MaasEtAlHypsamasia.pdf | access-date = 9 May 2013 | oclc = 493312921 }} {{Dead link|date=April 2019 |bot=InternetArchiveBot |fix-attempted=yes }}<!-- {{Harvnb|Maas|Thewissen|Kappelman|1998}} -->
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| last1 = Maas | first1 = M.C.
| last2 = Thewissen | first2 = J.G.M.
| last3 = Kappelman | first3 = J.
| chapter = ''Hypsamasia seni'' (Mammalia: Embrithopoda) and other mammals from the Eocene Kartal Formation of Turkey | pages = 286–297
| editor1-last = Beard | editor1-first = K.C.
| editor2-last = Dawson | editor2-first = M.R.
| title = Dawn of the Age of Mammals in Asia
| year = 1998 | series = Bulletin of Carnegie Museum of Natural History | volume = 34
| url = http://www.neomed.edu/DEPTS/ANAT/Thewissen/pdf/1998MaasEtAlHypsamasia.pdf | accessdate = May 2013
| oclc = 493312921 | ref = harv}}<!-- {{Harvnb|Maas|Thewissen|Kappelman|1998}} -->
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| last1 = McKenna | first1 = Malcolm C.
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| title = Classification of Mammals Above the Species Level
| title = Classification of Mammals Above the Species Level
| year = 1997 | publisher = Columbia University Press | location = New York
| year = 1997 | publisher = Columbia University Press | location = New York
| isbn = 0231110138 | oclc = 37345734 | ref = harv}}<!-- {{Harvnb|McKenna|Bell|1997}} -->
| isbn = 0231110138 | oclc = 37345734 }}<!-- {{Harvnb|McKenna|Bell|1997}} -->
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| last1 = McKenna | first1 = M.C.
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| title = Affinities and palaeobiogeographic significance of the Mongolian Paleogene genus Phenacolophus
| title = Affinities and palaeobiogeographic significance of the Mongolian Paleogene genus Phenacolophus
| year = 1977 | journal = Geobios, Memoire special | volume = 1 | pages = 61–85
| year = 1977 | journal = Geobios| volume = 10 |issue= Suppl 1| pages = 61–85
| doi = 10.1016/S0016-6995(77)80008-9 | oclc = 4656767437}}<!-- {{Harvnb|McKenna|Manning|1977}} -->
| doi = 10.1016/S0016-6995(77)80008-9 | oclc = 4656767437}}<!-- {{Harvnb|McKenna|Manning|1977}} -->
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| year = 1966 | publisher = Ankara University Press
| year = 1966 | publisher = Ankara University Press
| series = A.Ü. D.T.C.F. (University of Ankara, Faculty of Languages, History and Geography Publications) | volume = 172 | pages = 1–104
| series = A.Ü. D.T.C.F. (University of Ankara, Faculty of Languages, History and Geography Publications) | volume = 172 | pages = 1–104
| oclc = 16763756 | ref = harv}}<!-- {{Harvnb|Ozansoy|1966}} -->
| oclc = 16763756 }}<!-- {{Harvnb|Ozansoy|1966}} -->
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| last3 = Iliesco | first3 = M.
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| title = Decouverte d'un Embrithopode nouveau (Mammalia) dans la Paléogène de la dépression de Hateg (Roumanie) et considération générales sur la géologie de la région
| title = Decouverte d'un Embrithopode nouveau (Mammalia) dans la Paléogène de la dépression de Hateg (Roumanie) et considération générales sur la géologie de la région
| year = 1976 | journal = Neues Jahrbuch für Geologie und Paläontologie Monatshefte | volume = 1 | issue = 11 | pages = 690–698
| year = 1976 | journal = Neues Jahrbuch für Geologie und Paläontologie - Monatshefte | volume = 1 | issue = 11 | pages = 690–698
| ref = harv}}<!-- {{Harvnb|Radulesco|Iliesco|Iliesco|1976}} -->
}}<!-- {{Harvnb|Radulesco|Iliesco|Iliesco|1976}} -->
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| title = ''Crivadiatherium iliescui'' n. sp., nouvel Embrithopode (Mammalia) dans le Paléogène ancien de la depression de Hateg (Roumanie)
| title = ''Crivadiatherium iliescui'' n. sp., nouvel Embrithopode (Mammalia) dans le Paléogène ancien de la depression de Hateg (Roumanie)
| year = 1985 | journal = Palaeovertebrata | volume = 15 | issue = 3 | pages = 139–57
| year = 1985 | journal = Palaeovertebrata | volume = 15 | issue = 3 | pages = 139–57
| ref = harv}}<!-- {{Harvnb|Radulesco|Sudre|1985}} -->
}}<!-- {{Harvnb|Radulesco|Sudre|1985}} -->
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| year = 2006 | publisher = JHU Press | location = Baltimore
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| title = The Rise of Placental Mammals: Origins and Relationships of the Major Extant Clades
| title = The Rise of Placental Mammals: Origins and Relationships of the Major Extant Clades
| year = 2005 | publisher = JHU Press
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| isbn = 9780801880223 | oclc = 55801049 | ref = harv}}<!-- {{Harvnb|Rose|Archibald|2005}} -->
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| title = ''Palaeoamasia kansui'' Ozansoy 1966, embrithopode (Mammalia) de l'Eocene de Anatolie
| title = ''Palaeoamasia kansui'' Ozansoy 1966, embrithopode (Mammalia) de l'Eocene de Anatolie
| year = 1979 | journal = Annales de paléontologie (Vértébres) | volume = 65 | issue = 1 | pages = 73–91
| year = 1979 | journal = Annales de paléontologie (Vértébres) | volume = 65 | issue = 1 | pages = 73–91
| ref = harv}}<!-- {{Harvnb|Şen|Heintz|1979}} -->
}}<!-- {{Harvnb|Şen|Heintz|1979}} -->
{{Refend}}
{{Refend}}


{{Paenungulata}}
{{Taxonbar|from=Q131666}}
{{Taxonbar|from=Q131666}}


[[Category:Embrithopods|Embrithopods]]
[[Category:Embrithopods| ]]
[[Category:Eocene mammals]]
[[Category:Eocene mammals]]
[[Category:Oligocene mammals]]
[[Category:Oligocene mammals]]

Latest revision as of 15:45, 28 September 2024

Embrithopoda
Temporal range: Paleocene - Oligocene, 56–28 Ma[1]
Arsinoitherium zitteli[2]
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Clade: Paenungulatomorpha
Grandorder: Paenungulata
Mirorder: Tethytheria
Order: Embrithopoda
Andrews 1906
Families

Embrithopoda ("heavy-footed") is an order of extinct mammals known from Asia, Africa and Eastern Europe. Most of the embrithopod genera are known exclusively from jaws and teeth dated from the late Paleocene to the late Eocene; however, the order is best known from its terminal member, the elephantine Arsinoitherium.[3]

Description

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While embrithopods bore a superficial resemblance to rhinoceroses, their horns had bony cores covered in keratinized skin. Not all embrithopods possessed horns, either. Despite their appearance, they have been regarded as related to elephants, not perissodactyls.[4]

As tethytheres,[5] the Embrithopoda have been believed to be part of the clade Afrotheria. However, a study of the basal arsinoitheriid, Palaeoamasia, suggests that embrithopods are not tethytheres or even paenungulates, and that they need to be better sampled in an analysis of eutherian relationships to clarify if they are even afrotherians.[6] It is also not clear if embrithopods originated in Africa or Eurasia.[6] However, recent findings demonstrate an African origin for embrithopods and furthermore a relationship with other paenungulates, albeit having diverged earlier than previously thought.[7]

Fossils of embrithopods, such as Arsinoitherium, have been found in Egypt, Ethiopia, Kenya, Morocco, Mongolia, Turkey, Romania, Namibia,[8] Tunisia[9] and Croatia.[10] Until the 1970s, only Arsinoitherium itself was known, appearing isolated in the fossil record.[4]

Classification

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McKenna & Manning 1977 and McKenna & Bell 1997 considered Phenacolophus from Mongolia a primitive embrithopod, although this attribution was challenged by several other authors.[11] A 2016 cladistic study found Phenacolophus as a stem-perissodactyl and the embrithopods at the base of Altungulata.[12][6] More recently, an afrothere identity has been vindicated, albeit more basal than previously assumed.[7]

Order Embrithopoda Andrews 1906 sensu Prothero & Schoch 1989 (=Barypoda Andrews 1904)[13]

Notes

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  1. ^ Emmanuel Gheerbrant; Arnaud Schmitt; László Kocsis (2018). "Early African fossils elucidate the origin of embrithopod mammals". Current Biology. Online edition. doi:10.1016/j.cub.2018.05.032.
  2. ^ Rose 2006, pp. 242–3
  3. ^ Rose 2006, p. 265
  4. ^ a b "Introduction to the Embrithopoda". University of California Museum of Paleontology. Retrieved 7 March 2013.
  5. ^ Embrithopoda in the Paleobiology Database. Retrieved March 2013.
  6. ^ a b c Erdal, O.; Antoine, P.-O.; Sen, S.; Smith, A. (2016). "New material of Palaeoamasia kansui (Embrithopoda, Mammalia) from the Eocene of Turkey and a phylogenetic analysis of Embrithopoda at the species level". Palaeontology. 59 (5): 631–655. doi:10.1111/pala.12247.[Erdal et al.'s inclusion of Embrithopoda in Tabuce et al. (2007) finds them outside of (Ungulata + Afrotheria). Since this clade is not supported by molecular data, it suggests the need to explore the relationships of embrithopods, as they could potentially have affinities with laurasiatheres and "true" ungulates.]
  7. ^ a b Gheerbrant, E.; Schmitt, A.; Kocsis, L. (July 2018). "Early African fossils elucidate the origin of embrithopod mammals". Current Biology. 28 (13): 2167-2173.e2. Retrieved 28 September 2024.
  8. ^ a b M. Pickford; B. Senut; J. Morales; P. Mein; I. M. Sanchez (2008). "Mammalia from the Lutetian of Namibia". Memoir of the Geological Survey of Namibia. 20: 465–514.
  9. ^ Nicolas Vialle; Gilles Merzeraud; Cyrille Delmer; Monique Feist; Suzanne Jiquel; Laurent Marivaux; Anusha Ramdarshan; Monique Vianey-Liaud; El Mabrouk Essid; Wissem Marzougui; Hayet Khayati Ammar; Rodolphe Tabuce (2013). "Discovery of an embrithopod mammal (Arsinoitherium?) in the late Eocene of Tunisia". Journal of African Earth Sciences. 87: 86–92. doi:10.1016/j.jafrearsci.2013.07.010.
  10. ^ Rab danas
  11. ^ Koenigswald, W. v. (2012). "Unique differentiation of radial enamel in Arsinoitherium (Embrithopoda, Tethytheria)". Historical Biology. 25 (2): 183–192. doi:10.1080/08912963.2012.714658.
  12. ^ Radulesco & Sudre 1985; Maas, Thewissen & Kappelman 1998, p. 291; Rose & Archibald 2005, pp. 97–98
  13. ^ Mikko's Phylogeny Archive [1] Haaramo, Mikko (2007). "†Embrithopoda - arsinoitheres". Retrieved 30 December 2015.

References

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