Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Archosauromorpha
Cladus: Crurotarsi
Divisio: Archosauria
Cladus: Avemetatarsalia
Cladus: Ornithodira
Subtaxon: Dinosauromorpha
Cladus: Dinosauriformes
Cladus: Dracohors
Cladus: Dinosauria
Cladus: Saurischia
Cladus: Eusaurischia
Subordo: Theropoda
Cladus: Neotheropoda
Cladus: Averostra
Cladus: Tetanurae
Cladus: Avetheropoda
Cladus: Coelurosauria
Cladus: Tyrannoraptora
Cladus: Maniraptoromorpha
Cladus: Maniraptoriformes
Cladus: Maniraptora
Cladus: Pennaraptora
Cladus: Paraves
Cladus: Eumaniraptora
Cladus: Avialae
Infraclassis: Aves
Cladus: Avebrevicauda
Cladus: Pygostylia
Cladus: Ornithothoraces
Cladus: Ornithuromorpha
Cladus: Carinatae
Parvclassis: Neornithes
Cohors: Neognathae
Cladus: Neoaves
Cladus: Aequornithes
Ordo: Pelecaniformes
Familia: †Plotopteridae
Genera: †Copepteryx – †Hokkaidornis – †Klallamornis – †Olympidytes – †Phocavis – †Plotopterum – †Stemec – †Tonsala
Name
Plotopteridae Howard, 1969: 69
References
Primary references
Howard, H. 1969. A new avian fossil from Kern County, California. The Condor 71(1): 68–69. DOI: 10.2307/1366050 PDF Reference page.
Links
Plotopteridae – Taxon details on Fossilworks.
Plotopteridae[1] is an extinct family of flightless seabirds with uncertain placement, generally considered as member of order Suliformes.[2] They exhibited remarkable convergent evolution with the penguins, particularly with the now extinct giant penguins.[3][4] 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.[5] Plotopterids are regarded as closely related to Anhingidae (darters) and Phalacrocoracidae (cormorants).[2] 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.[2][6][7] However, the endocast morphology of stem group Sphenisciformes differs from both Plotopteridae and modern penguins.[8]
Their fossils have been found in California, Oregon,[9] Washington,[10][11] British Columbia,[12] Hokkaido, Tōhoku, Chūbu, Kyushu.[13] They seem to have evolved on arctic islands during the mid-Eocene, spreading southwards with the formation of kelp forests [14] 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.[2] 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.
Tonsala hildegardae fossils
The second species to be named from rocks along the eastern Pacific Ocean was Tonsala hildegardae[15] from the late Oligocene lower part of the Pysht Formation in Washington State. More fossils of T. hildegardae have since been described [16][17] and included some of the first known examples of borings made by the marine bone-eating worm Osedax in bird bones.[18]
Reconstruction of Copepteryx.
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.
References
Howard, H. (1969). "A new avian fossil from Kern County, California" (PDF). Condor. 71 (1): 68–69. doi:10.2307/1366050. JSTOR 1366050.
Mori, Hirotsugu; Miyata, Kazunori (2021). "Early Plotopteridae Specimens (Aves) from the Itanoura and Kakinoura Formations (Latest Eocene to Early Oligocene), Saikai, Nagasaki Prefecture, Western Japan". Paleontological Research. 25 (2): 145–159. doi:10.2517/2020PR018. ISSN 1342-8144. S2CID 233029559.
Olson, Storrs L.; Hasegawa, Yoshikazu (1979). "Fossil Counterparts of Giant Penguins from the North Pacific". Science. 206 (4419): 688–689. Bibcode:1979Sci...206..688O. doi:10.1126/science.206.4419.688. PMID 17796934. S2CID 12404154.
Olson, Storrs L. & Hasegawa, Yoshikazu (1996). "A new genus and two new species of gigantic Plotopteridae from Japan (Aves: Pelecaniformes)". J. Vertebr. Paleontol. 16 (4): 742–751. Bibcode:1996JVPal..16..742O. doi:10.1080/02724634.1996.10011362.
Tatsuro et al., New Skeletal Remains of Plotopterids from Japan, SVP 2015
Kawabe, Soichiro; Ando, Tatsuro; Endo, Hideki (2014). "Enigmatic affinity in the brain morphology between plotopterids and penguins, with a comprehensive comparison among water birds". Zoological Journal of the Linnean Society. 170 (3): 467–493. doi:10.1111/zoj.12072. ISSN 1096-3642.
Mayr, Gerald; Goedert, James L.; Vogel, Olaf (2015-07-04). "Oligocene plotopterid skulls from western North America and their bearing on the phylogenetic affinities of these penguin-like seabirds". Journal of Vertebrate Paleontology. 35 (4): e943764. Bibcode:2015JVPal..35E3764M. doi:10.1080/02724634.2014.943764. ISSN 0272-4634. S2CID 83729696.
Mayr, Gerald; Goedert, James L.; Pietri, Vanesa L. De; Scofield, R. Paul (2021). "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 of Zoological Systematics and Evolutionary Research. 59 (1): 264–276. doi:10.1111/jzs.12400. ISSN 1439-0469.
Goedert, James L. (1988). "A new late Eocene species of Plototpteridae (Aves: Pelecaniformes) from northwestern Oregon". Proceedings of the California Academy of Sciences. 45: 97–102.
Gerald Mayr & James L. Goedert (2016). "New late Eocene and Oligocene remains of the flightless, penguin-like plotopterids (Aves, Plotopteridae) from western Washington State, U.S.A.". Journal of Vertebrate Paleontology. 36 (4): e1163573. Bibcode:2016JVPal..36E3573M. doi:10.1080/02724634.2016.1163573. S2CID 88129671.
Mayr, Gerald; Goedert, James L. (2018). "First record of a tarsometatarsus of Tonsala hildegardae (Plotopteridae) and other avian remains from the late Eocene/early Oligocene of Washington State (USA)". Geobios. 51 (1): 51–59. Bibcode:2018Geobi..51...51M. doi:10.1016/j.geobios.2017.12.006.
Gary Kaiser; Junya Watanabe & Marji Johns (2015). "A new member of the family Plotopteridae (Aves) from the late Oligocene of British Columbia, Canada". Palaeontologia Electronica. 18 (3): Article number 18.3.52A.
"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" (PDF) (Press release). 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. 2020-08-18. Retrieved 2022-02-26.
Mayr, G.; Goedert, J. (2021). "New late Eocene and Oligocene plotopterid fossils from Washington State (USA), with a revision of "Tonsala" buchanani (Aves, Plotopteridae)". Journal of Paleontology. (e1163573) online preview: 224–236. doi:10.1017/jpa.2021.81. S2CID 240582610.
Olson, Storrs L. (1980). "A new genus of penguin-like pelecaniform bird from the Oligocene of Washington (Pelecaniformes: Plotopteridae)". Contributions in Science. 330: 51–57. doi:10.5962/p.208144. S2CID 4803730.
Goedert, James L.; Cornish, John (2002). "A preliminary report on the diversity and stratigraphic distribution of the Plotopteridae (Pelecaniformes) in Paleogene rocks of Washington State, USA". Proceedings of the 5th Symposium of the Society of Avian Paleontology and Evolution, Beijing: 63–76.
Mayr, Gerald; Goedert, James L. (2017). "First record of a tarsometatarsus of Tonsala hildegardae (Plotopteridae) and other avian remains from the late Eocene/early Oligocene of Washington State (USA)". Geobios. 51 (1): 51–59. Bibcode:2018Geobi..51...51M. doi:10.1016/j.geobios.2017.12.006.
Kiel, Steffen; Kahl, W.-A.; Goedert, James L. (2011). "Osedax borings in fossil marine bird bones". Naturwissenschaften. 98 (1): 51–55. Bibcode:2011NW.....98...51K. doi:10.1007/s00114-010-0740-5. PMC 3018246. PMID 21103978.
Howard, H. (1969). "A new avian fossil from Kern County, California" (PDF). Condor. 71 (1): 68–69. doi:10.2307/1366050. JSTOR 1366050.
Olson, Storrs L.; Hasegawa, Yoshikazu (1979). "Fossil Counterparts of Giant Penguins from the North Pacific". Science. 206 (4419): 688–689. Bibcode:1979Sci...206..688O. doi:10.1126/science.206.4419.688. PMID 17796934. S2CID 12404154.
Olson, Storrs L. & Hasegawa, Yoshikazu (1996). "A new genus and two new species of gigantic Plotopteridae from Japan (Aves: Pelecaniformes)". J. Vertebr. Paleontol. 16 (4): 742–751. Bibcode:1996JVPal..16..742O. doi:10.1080/02724634.1996.10011362.
Schreiber, E.A. & Burger, J. (2001). Biology of Marine Birds. Taylor & Francis. ISBN 0-8493-9882-7.
Mayr, Gerald (2005). "Tertiary plotopterids (Aves, Plotopteridae) and a novel hypothesis on the phylogenetic relationships of penguins (Spheniscidae)" (PDF). Journal of Zoological Systematics and Evolutionary Research. 43 (1): 67–71. doi:10.1111/j.1439-0469.2004.00291.x.
Gary Kaiser; Junya Watanabe & Marji Johns (2015). "A new member of the family Plotopteridae (Aves) from the late Oligocene of British Columbia, Canada". Palaeontologia Electronica. 18 (3): Article number 18.3.52A.
Gerald Mayr & James L. Goedert (2016). "New late Eocene and Oligocene remains of the flightless, penguin-like plotopterids (Aves, Plotopteridae) from western Washington State, U.S.A.". Journal of Vertebrate Paleontology. 36 (4): e1163573. Bibcode:2016JVPal..36E3573M. doi:10.1080/02724634.2016.1163573. S2CID 88129671.
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