View source for Ophthalmosauridae

{{Short description|Extinct family of reptiles}}
{{Automatic taxobox
| name = Ophthalmosaurids
| fossil_range = [[Middle Jurassic]]-[[Late Cretaceous]], {{fossil_range|174.1|93.9}}
| image = OphthalmosaurusIcenius-NaturalHistoryMuseum-August23-08.jpg
| image_upright = 1.15
| image_caption =  Composite skeleton (NHMUK PV R3702, R3893, R4124) of ''Ophthalmosaurus icenicus'' at the [[Natural History Museum, London]] 
| taxon = Ophthalmosauridae
| authority = [[George H. Baur|Baur]], [[1887 in paleontology|1887]]
| subdivision_ranks = Subgroups
| subdivision = *†''[[Arthropterygius]]''
*†''[[Keilhauia]]''
*†''[[Macropterygius]]''
*†''[[Muiscasaurus]]''
*†''[[Myobradypterygius]]''
*†[[Ophthalmosaurinae]]
*†'''Panplatypterygiine'''
**†''[[Catutosaurus]]''
**†[[Platypterygiinae]]
| synonyms = *Ophthalmosauria
}}

'''Ophthalmosauridae''' is an extinct family of [[thunnosaur]] [[ichthyosaurs]] from the [[Middle Jurassic]] to the early [[Late Cretaceous]] ([[Bajocian]] - [[Cenomanian]]) worldwide.<ref name=Fetal11>{{cite journal |last=Fischer |first=V. |author2=Masure, E. |author3=Arkhangelsky, M.S. |author4= Godefroit, P.  |year=2011 |title=A new Barremian (Early Cretaceous) ichthyosaur from western Russia |journal=Journal of Vertebrate Paleontology |volume=31 |issue=5 |pages=1010–1025 |doi=10.1080/02724634.2011.595464 |bibcode=2011JVPal..31.1010F |hdl=2268/92828 |s2cid=86036325 |url=https://orbi.uliege.be/handle/2268/92828 }}</ref> Almost all ichthyosaurs from the Middle Jurassic onwards belong to the family, until the extinction of ichthyosaurs in the early Late Cretaceous. Ophthalmosaurids appeared worldwide during early Bajocian, subsequent to the disappearance of most other ichthyosaur lineages after the end of the [[Toarcian]].<ref name=":0">{{Cite journal|last1=Fischer|first1=Valentin|last2=Weis|first2=Robert|last3=Thuy|first3=Ben|date=2021-02-22|title=Refining the marine reptile turnover at the Early–Middle Jurassic transition|journal=PeerJ|volume=9|pages=e10647|doi=10.7717/peerj.10647|issn=2167-8359|pmc=7906043|pmid=33665003 |doi-access=free }}</ref> Currently, the oldest known ophthalmosaurids is ''[[Mollesaurus]]'' from the early Bajocian of Argentina,<ref name=Mollesaurus>{{cite journal |author=Marta S. Fernández |year=1999 |title=A new ichthyosaur from the Los Molles Formation (Early Bajocian), Neuquen Basin, Argentina |journal=Journal of Paleontology |volume=73 |issue=4 |pages=677–681 |jstor=1306766|doi=10.1017/S0022336000032492 |bibcode=1999JPal...73..677F |s2cid=130751356 }}</ref> as well as indeterminate remains of the same age from Luxembourg and Canada.<ref name=":0" /> Named by George H. Baur, in 1887, the family contains the basal taxa like ''[[Ophthalmosaurus]]''.<ref name=Fetal11/> Appleby (1956) named the taxon Ophthalmosauria which was followed by some authors,<ref name=Motani>{{cite journal |author=Ryosuke Motani |year=1999 |title=Phylogeny of the Ichthyopterygia |url=http://mygeologypage.ucdavis.edu/motani/pdf/Motani1999c.pdf |journal=Journal of Vertebrate Paleontology |volume=19 |issue=3 |pages=472–495 |doi=10.1080/02724634.1999.10011160 |bibcode=1999JVPal..19..473M |access-date=2011-10-19 |archive-url=https://web.archive.org/web/20160305104153/http://mygeologypage.ucdavis.edu/motani/pdf/Motani1999c.pdf |archive-date=2016-03-05  }}</ref><ref name=DM10>{{cite journal |author1=Patrick S. Druckenmiller |author2=Erin E. Maxwell |year=2010 |title=A new Lower Cretaceous (lower Albian) ichthyosaur genus from the Clearwater Formation, Alberta, Canada |journal=Canadian Journal of Earth Sciences |volume=47 |issue=8 |pages=1037–1053 |doi=10.1139/E10-028 |url=http://rparticle.web-p.cisti.nrc.ca/rparticle/AbstractTemplateServlet?calyLang=eng&journal=cjes&volume=47&year=0&issue=8&msno=e10-028 |bibcode=2010CaJES..47.1037D }}{{dead link|date=March 2018 |bot=InternetArchiveBot |fix-attempted=yes }}</ref> but these two names are often treated as synonyms; Ophthalmosauridae has the priority over Ophthalmosauria.<ref name=Fetal11/> However, some researchers argue that Ophthalmosauridae should be restricted to the group typically referred to as [[Ophthalmosaurinae]], with classic [[Platypterygiinae]] instead being referred to as Undorosauridae or Brachypterygiidae and  Ophthalmosauria being used to unite these two groups.<ref name="Zverkov2019">{{cite journal|last1=Zverkov|first1=N. G.|last2=Efimov|first2=V. M.|year=2019|title=Revision of ''Undorosaurus'', a mysterious Late Jurassic ichthyosaur of the Boreal Realm|journal=Journal of Systematic Palaeontology|volume=17|issue=14|pages=963–993|doi=10.1080/14772019.2018.1515793|bibcode=2019JSPal..17.1183Z |s2cid=91912834 |url=http://paleorxiv.org/s95xm/ }}</ref><ref name="Cortes2021">{{cite journal|first1=D|last1=Cortés|first2=E. E.|last2=Maxwell|first3=H. C. E.|last3=Larsson|year=2021|title=Re-appearance of hypercarnivore ichthyosaurs in the Cretaceous with differentiated dentition: Revision of ''"Platypterygius" sachicarum'' (Reptilia: Ichthyosauria, Ophthalmosauridae) from Colombia|journal=Journal of Systematic Palaeontology|volume=19|issue=14|pages=969–1002|doi=10.1080/14772019.2021.1989507|bibcode=2021JSPal..19..969C |s2cid=244512087 }}</ref>

== Phylogeny ==
[[Image:Ophthalmosaurus_icenicus_updated_reconstruction.png|thumb|right|[[Paleoart|Life restoration]] of ''Ophthalmosaurus'']]
Ophthalmosauridae is a [[node-based taxon]] defined by Ryosuke Motani (1999) as "the last common ancestor of ''[[Brachypterygius extremus]]'' and ''[[Ophthalmosaurus icenicus]]'' and all of its descendants". The definition he proposed for Ophthalmosauria was exactly the same.<ref name=Motani/> In this case both definitions are synonyms. The [[cladogram]] below follows the topology from a 2010 analysis by Patrick S. Druckenmiller and Erin E. Maxwell.<ref name=DM10/>

{{clade| style=font-size:85%;line-height:85%
|label1=[[Thunnosauria]]&nbsp;
|1={{clade
   |1=''[[Ichthyosaurus]]'' 
   |2={{clade
      |1=''[[Stenopterygius]]''
      |2={{clade
         |1=''"[[Ophthalmosaurus]]" natans'' (=''[[Baptanodon]]'')
         |label2=&nbsp;'''Ophthalmosauridae'''&nbsp;
         |2={{clade
            |1={{clade
               |1=''[[Aegirosaurus]]''
               |2=''[[Ophthalmosaurus]]'' <small>(type species)</small> }}
            |2={{clade
               |1=''[[Mollesaurus]]''
               |2={{clade
                  |1=''[[Athabascasaurus]]''
                  |2={{clade
                     |1=''[[Brachypterygius]]''
                     |2={{clade
                        |1=''[[Arthropterygius]]''
                        |2=''[[Caypullisaurus]]''
                        |3=''"[[Platypterygius]]" hercynicus''
                        |4=''"[[Platypterygius]]" australis'' (=''Longirostria'')<ref name=Platypterygius>Arkhangel'sky, M. S., 1998, On the Ichthyosaurian Genus Platypterygius: Palaeontological Journal, v. 32, n. 6, p. 611-615.</ref>
                        |5=''[[Platypterygius]]'' <small>(type species)</small>
                        |6={{clade
                           |1=''[[Maiaspondylus]]''
                           |2=''"[[Platypterygius]]" americanus'' (=''Tenuirostria'')<ref name=Platypterygius/> }} }} }} }} }} }} }} }} }} }}

Valentin Fischer, Edwige Masure, Maxim S. Arkhangelsky and [[Pascal Godefroit]] (2011) described a new genus of Ophthalmosauridae. They redefined it as "the last common ancestor of ''[[Arthropterygius chrisorum]]'' and ''Ophthalmosaurus icenicus'' and all of its descendants". In this case Ophthalmosauria isn't a [[junior synonym]] of Ophthalmosauridae. The cladogram below follows Fischer ''et al.'' 2011.<ref name=Fetal11/>

{{clade| style=font-size:85%;line-height:85%
|label1=[[Thunnosauria]]&nbsp;
|1={{clade
   |1=''[[Ichthyosaurus]]'' 
   |2={{clade
      |1=''[[Stenopterygius]]''
      |2={{clade
         |1=''[[Chacaicosaurus]]''
         |label2=&nbsp;'''Ophthalmosauridae'''&nbsp;
         |2={{clade
            |1=''[[Arthropterygius]]''
            |label2='''*'''
            |2={{clade
               |1=''[[Brachypterygius]]''
               |2=''[[Ophthalmosaurus]]'' <small>(type species)</small>
               |3={{clade
                  |1={{clade
                     |1=''[[Maiaspondylus]]''
                     |2={{clade
                        |1=''[[Mollesaurus]]''
                        |2={{clade
                           |1=''[[Aegirosaurus]]''
                           |2=''[[Sveltonectes]]'' }} }} }}
                  |2={{clade
                     |1=''"[[Platypterygius]]" hercynicus''
                     |2={{clade
                        |1=''[[Caypullisaurus]]''
                        |2={{clade
                           |1=''[[Athabascasaurus]]''
                           |2=''"[[Platypterygius]]" australis'' (=''Longirostria'') <ref name=Platypterygius/> }} }} }} }} }} }} }} }} }} }}

<small>*<u>Note</u>: Placement of Ophthalmosauria by definition.</small>

Fischer ''et al.'' (2012) described another new genus of Ophthalmosauridae. They defined for the first time two [[subfamily (biology)|subfamilies]] within the Ophthalmosauridae, [[Ophthalmosaurinae]] and [[Platypterygiinae]]. ''Ophthalmosaurus'' and ''Platypterygius'' were found to be non-[[monophyletic]]. Ophthalmosauridae was characterized by a reduced extracondylar area, a plate-like dorsal trochanter, a [[humerus]] with a facet for an anterior accessory element and the absence of notching on the paddle elements of the [[fin|forefin]]. The cladogram below follows Fischer ''et al.'' 2012.<ref name=Acamptonectes>{{cite journal |author1=Valentin Fischer |author2=Michael W. Maisch |author3=Darren Naish |author4=Ralf Kosma |author5=Jeff Liston |author6=Ulrich Joger |author7=Fritz J. Krüger |author8=Judith Pardo Pérez |author9=Jessica Tainsh |author10=Robert M. Appleby |year=2012 |title=New Ophthalmosaurid Ichthyosaurs from the European Lower Cretaceous Demonstrate Extensive Ichthyosaur Survival across the Jurassic–Cretaceous Boundary |journal=PLOS ONE |volume=7 |issue=1 |pages=e29234 |doi=10.1371/journal.pone.0029234 |pmid=22235274 |pmc=3250416|bibcode=2012PLoSO...729234F |doi-access=free }}</ref>

{{clade| style=font-size:85%;line-height:85%
|label1=[[Thunnosauria]]&nbsp;
|1={{clade
   |1=''[[Ichthyosaurus]]'' 
   |2={{clade
      |1=''[[Stenopterygius]]''
      |2={{clade
         |1=''[[Chacaicosaurus]]''
         |label2=&nbsp;'''Ophthalmosauridae'''&nbsp;
         |2={{clade
            |1=''[[Arthropterygius]]''
            |label2='''*'''
            |2={{clade
               |label1=&nbsp;[[Ophthalmosaurinae]]&nbsp;
               |1={{clade
                  |1=''[[Mollesaurus]]''
                  |2={{clade
                     |1=''[[Ophthalmosaurus]]'' <small>(type species)</small>
                     |2={{clade
                        |1=''[[Baptanodon]]'' <small>(''"O." natans'')</small>
                        |2=''[[Acamptonectes]]'' }} }} }}
               |label2=&nbsp;[[Platypterygiinae]]&nbsp;
               |2={{clade
                  |1={{clade
                     |1=''[[Brachypterygius]]''
                     |2={{clade
                        |1=''[[Maiaspondylus]]''
                        |2={{clade
                           |1=''[[Aegirosaurus]]''
                           |2=''[[Sveltonectes]]'' }} }} }}
                  |2={{clade
                     |1=''"[[Platypterygius]]" hercynicus''
                     |2={{clade
                        |1=''[[Caypullisaurus]]''
                        |2={{clade
                           |1=''[[Athabascasaurus]]''
                           |2=''"[[Platypterygius]]" australis'' (=''Longirostria'')<ref name=Platypterygius/> }} }} }} }} }} }} }} }} }} }}

=== Since 2019 ===
In 2019, two species of ''[[Cryopterygius]]'' were assigned to the genus ''[[Undorosaurus]]'' by Zverkov & Efimov (2019). The authors considered the type species of the former genus, ''C. kristiansenae'', to be synonymous with ''Undorosaurus gorodischensis''; second species of ''Cryopterygius'', ''C. kielanae'', was tentatively maintained by the authors as a distinct species within the genus ''Undorosaurus''.<ref>{{cite journal |author1=Nikolay G. Zverkov |author2=Vladimir M. Efimov |name-list-style=amp |year=2019 |title=Revision of ''Undorosaurus'', a mysterious Late Jurassic ichthyosaur of the Boreal Realm |journal=Journal of Systematic Palaeontology |volume=17 |issue=14 |pages=963–993 |doi=10.1080/14772019.2018.1515793 |bibcode=2019JSPal..17.1183Z |s2cid=91912834 |url=https://figshare.com/articles/Revision_of_i_Undorosaurus_i_a_mysterious_Late_Jurassic_ichthyosaur_of_the_Boreal_Realm/7624868 }}</ref>

The following cladogram shows a possible phylogenetic positions within Ophthalmosauridae according to the analysis performed by Zverkov and Jacobs (2020).<ref name="Zverkov2020">{{cite journal |author1=Nikolay G. Zverkov |author2=Megan L. Jacobs |name-list-style=amp |orig-date=2020 |title=Revision of ''Nannopterygius'' (Ichthyosauria: Ophthalmosauridae): reappraisal of the 'inaccessible' holotype resolves a taxonomic tangle and reveals an obscure ophthalmosaurid lineage with a wide distribution |journal=Zoological Journal of the Linnean Society |volume=191 |issue=1 |year=2021 |pages=228–275 |doi=10.1093/zoolinnean/zlaa028}}</ref>

{{clade
|label1='''Ophthalmosauria'''
|1={{clade
   |1={{clade
      |label1=[[Ophthalmosaurinae]]
      |1={{clade
         |1={{clade
            |1={{clade
               |1={{clade
                  |1=''[[Acamptonectes|Acamptonectes densus]]''
                  
                     |2={{clade
                        |1=''[[Mollesaurus|Mollesaurus periallus]]''
                        
                           |2={{clade
                              |1=''[[Ophthalmosaurus natans]]''
                              |2=''[[Ophthalmosaurus icenicus]]''
                              }}
                           }}
                        }} }} }}
         |2={{clade
            |1={{clade
               |1=''[[Gengasaurus nicosiai]]''
               
                  |2={{clade
                     |1={{clade
                        |1=''[[Nannopterygius|Nannopterygius yasykovi]]''
                           |2={{clade
                              |1=''[[Nannopterygius|Nannopterygius enthekiodon]]''
                              |2=''[[Nannopterygius|Nannopterygius saveljeviensis]]''
                              |3=''[[Nannopterygius|Nannopterygius borealis]]''
                              }}
                           }}
                     |2={{clade
                          |1={{clade
                             |1=''[[Arthropterygius|Arthropterygius volgensis]]''
                             |2={{clade
                                |1=''[[Arthropterygius|Arthropterygius lundi]]''
                                |2=''[[Arthropterygius|Arthropterygius thalassonotus]]''
                                |3={{clade
                                      |1=''[[Arthropterygius|Arthropterygius hoybergeti]]''
                                      |2=''[[Arthropterygius|Arthropterygius chrisorum]]''
                                    }}
                                 }}
                              }}
                           }}
                        }}
                     }}
                  }}
               }}
            
        
      |label2=[[Platypterygiinae]]
      |2={{clade
         |1={{clade
            |1=''[[Brachypterygius extremus]]''
            |2=''[[Aegirosaurus leptospondylus]]''
            
               |3={{clade
                  |1=''[[Muiscasaurus catheti]]''
                  
                        |2={{clade
                           |1=''[[Leninia stellans]]''
                           
                              |2={{clade
                                 |1=''[[Sveltonectes insolitus]]''
                                 
                                 |2={{clade
                                    |1=''[[Athabascasaurus bitumineus]]''
                                    
                                    |2={{clade
                                       |1=''[[Platypterygius americanus]]''
                                       
                                       |2={{clade
                                          |1={{clade
                                             |1=''[[Acuetzpalin|Acuetzpalin carranzai]]''
                                             |2=''[[Platypterygius sachicarum]]''
                                             |3=''[[Caypullisaurus bonapartei]]''
                                             }}
                                          |2={{clade
                                             |1={{clade
                                                |1={{clade
                                                   |1=''[[Grendelius mordax]]''
                                                   |2=''[[Grendelius alekseevi]]''
                                                   |3=''[[Grendelius|Grendelius pseudoscythicus]]''
                                                   |4=''[[Grendelius|Grendelius zhuravlevi]]''
                                                   }}
                                                |2={{clade
                                                   |1={{clade
                                                   |1={{clade
                                                      |1=''[[Undorosaurus|Undorosaurus kielanae]]''
                                                      |2={{clade
                                                         |1=''[[Undorosaurus|Undorosaurus nessovi]]''
                                                         |2=''[[Undorosaurus|Undorosaurus gorodischensis]]''
                                                         }}
                                                      }}
                                                   }}
                                                }}
                                             }}
                                          |3={{clade
                                             |1=''[[Platypterygius australis]]''
                                             |2={{clade
                                                |1=''[[Plutoniosaurus bedengensis]]''
                                                |2=''[[Simbirskiasaurus|Simbirskiasaurus birjukovi]]''
                                                |3=''[[Platypterygius hercynicus]]''
                                                |4=''[[Sisteronia seeleyi]]''
                                                |5=''[[Platypterygius|Platypterygius platydactylus]]''
                                                |6=''[[Maiaspondylus lindoei]]''
                                                }}
                                             }} 
                                          }}
                                       }}
                                    }} }} }} }} }} }} }} }} }} }}

==References==
{{Reflist}}
{{Portal|Paleontology}}
{{Ichthyosauria|Parvipelvia}}
{{Taxonbar|from=Q1936147}}

[[Category:Ophthalmosauridae| ]]
[[Category:Middle Jurassic ichthyosaurs]]
[[Category:Late Jurassic ichthyosaurs]]
[[Category:Early Cretaceous ichthyosaurs]]
[[Category:Aalenian first appearances]]
[[Category:Turonian extinctions]]
[[Category:Ichthyosaurs]]
[[Category:Prehistoric reptile families]]