Plotopteridae

{{Short description|Extinct family of sea birds}}

{{Automatic taxobox

| name = Plotopterids

| fossil_range = Eocene – Miocene
~{{fossil range|Priabonian|Burdigalian}}

| image = Fossil of sternum of Copepteryx from Kitahata Karatsu Japan.JPG

| image_caption = Copepteryx

| taxon = Plotopteridae

| authority = Howard, 1969

| subdivision_ranks = Genera

| subdivision =

• {{extinct}}Empeirodytes
• {{extinct}}Phocavis
• {{extinct}}Plotopterum
• {{extinct}}Stemec
• {{extinct}}Stenornis
• {{extinct}}Tonsalinae Mayr & Goedert, 2018
• {{extinct}}Copepteryx
• {{extinct}}Hokkaidornis
• {{extinct}}Klallamornis
• {{extinct}}Olympidytes
• {{extinct}}Tonsala

}}

Plotopteridae{{cite journal|author= Howard, H. |year=1969|title= A new avian fossil from Kern County, California|journal=Condor|volume=71|issue=1|pages= 68–69|url=http://sora.unm.edu/sites/default/files/journals/condor/v071n01/p0068-p0069.pdf|doi= 10.2307/1366050|jstor=1366050}} is an extinct family of flightless seabirds with uncertain placement, generally considered as member of order Suliformes.{{Cite journal |last1=Mori |first1=Hirotsugu |last2=Miyata |first2=Kazunori |date=2021 |title=Early Plotopteridae Specimens (Aves) from the Itanoura and Kakinoura Formations (Latest Eocene to Early Oligocene), Saikai, Nagasaki Prefecture, Western Japan |url=https://bioone.org/journals/paleontological-research/volume-25/issue-2/2020PR018/Early-Plotopteridae-Specimens-Aves-from-the-Itanoura-and-Kakinoura-Formations/10.2517/2020PR018.full |journal=Paleontological Research |volume=25 |issue=2 |pages=145–159 |doi=10.2517/2020PR018 |s2cid=233029559 |issn=1342-8144}} They exhibited remarkable convergent evolution with the penguins, particularly with the now extinct giant penguins.{{cite journal|author= Olson, Storrs L.|author2= Hasegawa, Yoshikazu |year=1979|title= Fossil Counterparts of Giant Penguins from the North Pacific|journal=Science|volume=206|issue=4419|pages= 688–689|bibcode=1979Sci...206..688O|doi= 10.1126/science.206.4419.688|pmid=17796934|s2cid= 12404154 }}{{cite journal|author1=Olson, Storrs L. |author2=Hasegawa, Yoshikazu |name-list-style=amp |year=1996|title= A new genus and two new species of gigantic Plotopteridae from Japan (Aves: Pelecaniformes)|journal=J. Vertebr. Paleontol.|volume=16|issue=4|pages= 742–751|doi= 10.1080/02724634.1996.10011362|bibcode=1996JVPal..16..742O }} That they lived in the North Pacific, the other side of the world from the penguins, has led to them being described at times as the Northern Hemisphere's penguins, though they were not closely related. More recent studies have shown, however, that the shoulder-girdle, forelimb and sternum of plotopterids differ significantly from those of penguins, so comparisons in terms of function may not be entirely accurate.Tatsuro et al., New Skeletal Remains of Plotopterids from Japan, SVP 2015 Plotopterids are regarded as closely related to Anhingidae (darters) and Phalacrocoracidae (cormorants). On the other hand, there is a theory that this group may have a common ancestor with penguins due to the similarity of forelimb and brain morphology.{{Cite journal |last1=Kawabe |first1=Soichiro |last2=Ando |first2=Tatsuro |last3=Endo |first3=Hideki |year=2014 |title=Enigmatic affinity in the brain morphology between plotopterids and penguins, with a comprehensive comparison among water birds|url=https://onlinelibrary.wiley.com/doi/abs/10.1111/zoj.12072|journal=Zoological Journal of the Linnean Society |language=en |volume=170|issue=3|pages=467–493|doi=10.1111/zoj.12072|issn=1096-3642}}{{Cite journal|last1=Mayr|first1=Gerald|last2=Goedert|first2=James L.|last3=Vogel|first3=Olaf|date=2015-07-04|title=Oligocene plotopterid skulls from western North America and their bearing on the phylogenetic affinities of these penguin-like seabirds|url=https://doi.org/10.1080/02724634.2014.943764|journal=Journal of Vertebrate Paleontology|volume=35|issue=4|pages=e943764|doi=10.1080/02724634.2014.943764|bibcode=2015JVPal..35E3764M |s2cid=83729696 |issn=0272-4634}} However, the endocast morphology of stem group Sphenisciformes differs from both Plotopteridae and modern penguins.{{Cite journal|last1=Mayr|first1=Gerald|last2=Goedert|first2=James L.|last3=Pietri|first3=Vanesa L. De|last4=Scofield|first4=R. Paul|date=2021|title=Comparative osteology of the penguin-like mid-Cenozoic Plotopteridae and the earliest true fossil penguins, with comments on the origins of wing-propelled diving|journal=Journal of Zoological Systematics and Evolutionary Research|language=en|volume=59|issue=1|pages=264–276|doi=10.1111/jzs.12400|issn=1439-0469|doi-access=free}}

Their fossils have been found in California, Oregon,{{cite journal|author= Goedert, James L. |year=1988| title= A new late Eocene species of Plototpteridae (Aves: Pelecaniformes) from northwestern Oregon |journal= Proceedings of the California Academy of Sciences |volume= 45 |pages= 97–102}} Washington,{{Cite journal|author1=Gerald Mayr |author2=James L. Goedert |name-list-style=amp |year=2016 |title=New late Eocene and Oligocene remains of the flightless, penguin-like plotopterids (Aves, Plotopteridae) from western Washington State, U.S.A. |journal=Journal of Vertebrate Paleontology |volume=36 |issue=4 |pages=e1163573 |doi=10.1080/02724634.2016.1163573 |bibcode=2016JVPal..36E3573M |s2cid=88129671 }}{{cite journal| author1= Mayr, Gerald| author2= Goedert, James L.| year= 2018| title= First record of a tarsometatarsus of Tonsala hildegardae (Plotopteridae) and other avian remains from the late Eocene/early Oligocene of Washington State (USA)| journal= Geobios| volume= 51| issue= 1| pages= 51–59| doi= 10.1016/j.geobios.2017.12.006 | bibcode= 2018Geobi..51...51M}} British Columbia,{{Cite journal|author1=Gary Kaiser|author2=Junya Watanabe |author3=Marji Johns |name-list-style=amp |year=2015 |title=A new member of the family Plotopteridae (Aves) from the late Oligocene of British Columbia, Canada |journal=Palaeontologia Electronica |volume=18 |issue=3 |pages=Article number 18.3.52A |url=http://palaeo-electronica.org/content/2015/1359-plotopterid-in-canada }} Hokkaido, Tōhoku, Chūbu, Kyushu.{{Cite press release|url=http://www2.pref.fukui.jp/press/atfiles/pafd15972185673e.pdf |publisher=西海市教育委員会、蒲郡市教育委員会、福井県立恐竜博物館 |title=西海市から発見された世界最古級のペンギンモドキの化石について |accessdate=2022-02-26 |date=2020-08-18}} They seem to have evolved on arctic islands during the mid-Eocene, spreading southwards with the formation of kelp forests {{cite journal |last1=Mayr |first1=G.|last2=Goedert |first2=J. |year=2021 |title=New late Eocene and Oligocene plotopterid fossils from Washington State (USA), with a revision of "Tonsala" buchanani (Aves, Plotopteridae) |journal= Journal of Paleontology |volume=(e1163573) online preview |pages=224–236|doi=10.1017/jpa.2021.81 |s2cid=240582610 }} They ranged in size from that of a large cormorant (such as a Brandt's cormorant), to very large size, with femur length two times longer than emperor penguin. They had shortened wings optimised for underwater wing-propelled pursuit diving (like penguins or the now extinct great auk), and a body skeleton similar to that of the darter.

File:Tonsala hildegardae.png

The second species to be named from rocks along the eastern Pacific Ocean was Tonsala hildegardae{{Cite journal| author= Olson, Storrs L.| year= 1980| title= A new genus of penguin-like pelecaniform bird from the Oligocene of Washington (Pelecaniformes: Plotopteridae)| journal= Contributions in Science| volume= 330| pages= 51–57 | doi= 10.5962/p.208144| s2cid= 4803730| doi-access= free}} from the late Oligocene lower part of the Pysht Formation in Washington State. More fossils of T. hildegardae have since been described {{Cite journal| author1 = Goedert, James L.| author2= Cornish, John| year= 2002| title= A preliminary report on the diversity and stratigraphic distribution of the Plotopteridae (Pelecaniformes) in Paleogene rocks of Washington State, USA| journal= Proceedings of the 5th Symposium of the Society of Avian Paleontology and Evolution, Beijing| pages= 63–76}}{{cite journal| author1= Mayr, Gerald| author2= Goedert, James L.| year= 2017| title= First record of a tarsometatarsus of Tonsala hildegardae (Plotopteridae) and other avian remains from the late Eocene/early Oligocene of Washington State (USA)| journal= Geobios| volume= 51| issue= 1| pages= 51–59| doi= 10.1016/j.geobios.2017.12.006| bibcode= 2018Geobi..51...51M}} and included some of the first known examples of borings made by the marine bone-eating worm Osedax in bird bones.{{Cite journal| author1= Kiel, Steffen| author2= Kahl, W.-A.| author3= Goedert, James L.| year= 2011| title= Osedax borings in fossil marine bird bones| journal= Naturwissenschaften| volume= 98| issue= 1| pages= 51–55| doi= 10.1007/s00114-010-0740-5| pmid= 21103978| pmc= 3018246| bibcode= 2011NW.....98...51K}}

File:Copepteryx hexeris.jpg.]]

The earliest known member of the family, Phocavis maritimus lived in the late Eocene, but most of the known species lived during Oligocene time, becoming extinct in the early to mid-Miocene. That they became extinct at the same time as the giant penguins of the Southern Hemisphere, which also coincided with the radiation of the seals and dolphins, has led to speculation that the expansion of marine mammals was responsible for the extinction of the Plotopteridae, though this has not been formally tested.