PBDB Taxon

Basic info	Taxonomic history	Classification	Included Taxa
Morphology	Ecology and taphonomy	External Literature Search	Age range and collections

Sphenisciformes (penguin)

Reptilia - Theropoda

Taxonomy

Sphenisciformes was named by Sharpe (1891). It is extant.

It was reranked as the unranked clade Sphenisciformes by Clarke et al. (2007), Ksepka and Clarke (2010) and Sangster and Mayr (2021).

It was assigned to Carinatae by Gadow (1898); to Neognathae by Carroll (1988); to Pansphenisciformes by Clarke et al. (2003); to Neoaves by Slack et al. (2006); to Neognathae by Clarke et al. (2007), Ksepka and Clarke (2010) and Fordyce and Thomas (2011); to Aves by Baird (1992) and Park and Fitzgerald (2012); to Aves by Myrcha et al. (2002), Sepkoski (2002), Jadwiszczak (2006), Jadwiszczak (2006), Jadwiszczak (2012) and del Hoyo et al. (2014); to Procellariimorphae by Livezey and Zusi (2007) and Sangster and Mayr (2021); and to Aequornithes by Mayr (2022).

Synonymy list

Year	Name and author
1891	Sphenisciformes Sharpe
1898	Sphenisciformes Gadow p. 32
1988	Sphenisciformes Carroll
1992	Sphenisciformes Baird p. 24
2002	Sphenisciformes Myrcha et al. p. 17
2002	Sphenisciformes Sepkoski
2003	Sphenisciformes Clarke et al. p. 3
2006	Sphenisciformes Jadwiszczak p. 11
2006	Sphenisciformes Slack et al. p. 1150
2007	Sphenisciformes Clarke et al. p. 11545
2007	Sphenisciformes Livezey and Zusi p. 90
2010	Sphenisciformes Ksepka and Clarke p. 7
2011	Sphenisciformes Fordyce and Thomas p. 45
2012	Sphenisciformes Jadwiszczak p. 262
2012	Sphenisciformes Park and Fitzgerald p. 317
2014	Sphenisciformes del Hoyo et al.
2021	Sphenisciformes Sangster and Mayr p. 50
2022	Sphenisciformes Mayr p. 119 figs. Fig. 7.1

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Rank	Name	Author
kingdom	Animalia	()
—	Bilateria
—	Eubilateria	Ax 1987
—	Deuterostomia
phylum	Chordata	Haeckel 1874
subphylum	Vertebrata
superclass	Gnathostomata
—	Osteichthyes	()
subclass	Sarcopterygii	()
subclass	Dipnotetrapodomorpha	(Nelson 2006)
subclass	Tetrapodomorpha	()
—	Tetrapoda
—	Reptiliomorpha
—	Anthracosauria
subclass	Amphibiosauria	Kuhn 1967
—	Cotylosauria	()
—	Amniota
—	Sauropsida
class	Reptilia
subclass	Eureptilia	()
—	Romeriida
—	Diapsida	()
—	Archosauromorpha	(Huene 1946)

Rank	Name	Author
—	Crocopoda
—	Archosauriformes	Gauthier 1986
—	Eucrocopoda
—	Archosauria	()
informal	Avemetatarsalia
—	Ornithodira
—	Dinosauromorpha
—	Dinosauriformes
—	Dinosauria	()
order	Theropoda
—	Neotheropoda
—	Averostra	Paul 2002
—	Tetanurae
—	Coelurosauria	()
—	Maniraptora
—	Paraves
class	Aves
subclass	Neornithes	Gadow 1893
—	Aequornithes
—	Feraequornithes
superorder	Procellariimorphae
—	Sphenisciformes	(Sharpe 1891)
—	Sphenisciformes	(Sharpe 1891)

If no rank is listed, the taxon is considered an unranked clade in modern classifications. Ranks may be repeated or presented in the wrong order because authors working on different parts of the classification may disagree about how to rank taxa.

Unr. Sphenisciformes Sharpe 1891 [penguin]

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G. †Anthropornis Wiman 1905

†Anthropornis grandis Wiman 1905

†Anthropornis nordenskjoeldi Wiman 1905

Invalid names: Pachypteryx Wiman 1905 [synonym]

G. †Aprosdokitos Acosta Hospitaleche et al. 2017

†Aprosdokitos mikrotero Acosta Hospitaleche et al. 2017

G. †Archaeospheniscus Marples 1952

†Archaeospheniscus lopdelli Marples 1952

†Archaeospheniscus lowei Marples 1952

†Archaeospheniscus wimani Marples 1953

G. †Crossvallia Tambussi et al. 2005

†Crossvallia unienwillia Tambussi et al. 2005

†Crossvallia waiparensis Mayr et al. 2020

G. †Delphinornis Wiman 1905

†Delphinornis arctowskii Myrcha et al. 2002

†Delphinornis larseni Wiman 1905

Invalid names: Notodyptes Marples 1953 [synonym]

G. †Duntroonornis Marples 1952

†Duntroonornis parvus Marples 1952

G. †Icadyptes Clarke et al. 2007

†Icadyptes salasi Clarke et al. 2007

G. †Inkayacu Clarke et al. 2010

†Inkayacu paracasensis Clarke et al. 2010

G. †Kaiika Fordyce and Thomas 2011

†Kaiika maxwelli Fordyce and Thomas 2011

G. †Kairuku Ksepka et al. 2012

†Kairuku grebneffi Ksepka et al. 2012

†Kairuku waewaeroa Giovanardi et al. 2021

†Kairuku waitaki Ksepka et al. 2012

G. †Korora Marples 1952

†Korora oliveri Marples 1952

G. †Kumimanu Mayr et al. 2017

†Kumimanu biceae Mayr et al. 2017

†Kumimanu fordycei Ksepka et al. 2023

G. †Kupoupou Blokland et al. 2019

†Kupoupou stilwelli Blokland et al. 2019

G. †Marambiornis Myrcha et al. 2002

†Marambiornis exilis Myrcha et al. 2002

G. †Mesetaornis Myrcha et al. 2002

†Mesetaornis polaris Myrcha et al. 2002

G. †Muriwaimanu Mayr et al. 2017

†Muriwaimanu tuatahi Slack et al. 2006

G. †Pachydyptes Oliver 1930

†Pachydyptes ponderosus Oliver 1930

†Pachydyptes simpsoni Jenkins 1974

G. †Pakudyptes Ando et al. 2024

†Pakudyptes hakataramea Ando et al. 2024

Subfm. †Palaeospheniscinae Simpson 1946

G. †Eretiscus Olson 1986

†Eretiscus tonnii Simpson 1981

Invalid names: Microdytes Simpson 1981 [replaced]

G. †Palaeospheniscus Moreno and Mercerat 1891

†Palaeospheniscus bergi Moreno and Mercerat 1891

Invalid names: Palaeospheniscus concavus Ameghino 1905 [synonym], Palaeospheniscus gracilis Ameghino 1899 [synonym], Palaeospheniscus interplanus Ameghino 1905 [synonym], Palaeospheniscus planus Ameghino 1905 [synonym], Palaeospheniscus rothi Ameghino 1905 [synonym], Pseudospheniscus convexus Ameghino 1905 [synonym]

†Palaeospheniscus biloculata Simpson 1970

Invalid names: Perispheniscus wimani Ameghino 1905 [synonym]

†Palaeospheniscus huxleyorum Simpson 1973

†Palaeospheniscus patagonicus Moreno and Mercerat 1891

Invalid names: Palaeospheniscus affinis Ameghino 1905 [synonym], Palaeospheniscus intermedius Ameghino 1905 [synonym], Palaeospheniscus interruptus Ameghino 1905 [synonym], Palaeospheniscus menzbieri Moreno and Mercerat 1891 [synonym]

Invalid names: Chubutodyptes Simpson 1970 [synonym], Paraspheniscus Ameghino 1905 [synonym], Perispheniscus Ameghino 1905 [synonym], Pseudospheniscus Ameghino 1905 [synonym], Treleudytes Ameghino 1905 [synonym]

G. †Perudyptes Clarke et al. 2007

†Perudyptes devriesi Clarke et al. 2007

G. †Petradyptes Ksepka et al. 2023

†Petradyptes stonehousei Ksepka et al. 2023

G. †Platydyptes Marples 1952

†Platydyptes amiesi Marples 1952

†Platydyptes marplesi Simpson 1971

†Platydyptes novaezealandiae Oliver 1930

G. †Sequiwaimanu Mayr et al. 2017

†Sequiwaimanu rosieae Mayr et al. 2017

Fm. Spheniscidae Bonaparte 1831

G. †Anthropodyptes Simpson 1959

†Anthropodyptes gilli Simpson 1959

G. Aptenodytes Miller 1778

Aptenodytes forsteri Gray 1844 [emperor penguin]

Aptenodytes patagonicus Miller 1778 [king penguin]

†Aptenodytes ridgeni Simpson 1972

G. †Arthrodytes Ameghino 1905

†Arthrodytes andrewsi Ameghino 1901

G. †Dege Simpson 1979

†Dege hendeyi Simpson 1979

G. Eudyptes Vieillot 1816

†Eudyptes atatu Thomas et al. 2020

†Eudyptes calauina Chávez Hoffmeister et al. 2014

Eudyptes chrysocome Forster 1781 [southern rockhopper penguin]

Eudyptes chrysocome filholi Hutton 1879

Eudyptes chrysolophus Brandt 1837 [Macaroni penguin]

Eudyptes moseleyi Matthews and Iredale 1921 [Northern rockhopper penguin]

Eudyptes pachyrhynchus Gray 1845 [Fiordland penguin]

Invalid names: Tasidyptes hunteri van Tets and O'Connor 1983 [synonym]

Eudyptes robustus Oliver 1953 [Snares penguin]

Eudyptes schlegeli Finsch 1876 [royal penguin]

Eudyptes sclateri Buller 1888 [Erect-crested penguin]

†Eudyptes warhami Cole et al. 2019

Invalid names: Tasidyptes van Tets and O'Connor 1983 [synonym]

G. Eudyptula Bonaparte 1856

Eudyptula minor Forster 1781 [little penguin]

Eudyptula minor novaehollandiae Shaw 1826

†Eudyptula wilsonae Thomas et al. 2023

G. †Inguza Simpson 1975

†Inguza predemersus Simpson 1971

G. †Madrynornis Acosta Hospitaleche et al. 2007

†Madrynornis mirandus Acosta Hospitaleche et al. 2007

G. †Marambiornopsis Jadwiszczak et al. 2021

†Marambiornopsis sobrali Jadwiszczak et al. 2021

G. †Marplesornis Simpson 1972

†Marplesornis novaezealandiae Marples 1960

G. Megadyptes Milne-Edwards 1880

Megadyptes antipodes Hombron and Jacquinot 1841 [yellow-eyed penguin]

†Megadyptes antipodes richdalei Cole et al. 2019

†Megadyptes antipodes waitaha Boessenkool et al. 2009 [Waitaha penguin]

G. †Nucleornis Simpson 1979

†Nucleornis insolitus Simpson 1979

G. †Palaeeudyptes Huxley 1859

†Palaeeudyptes antarcticus Huxley 1859

†Palaeeudyptes gunnari Wiman 1905

†Palaeeudyptes klekowskii Myrcha et al. 2002

†Palaeeudyptes marplesi Brodkorb 1963

Invalid names: Eosphaeniscus Wiman 1905 [synonym]

G. †Paraptenodytes Ameghino 1891

†Paraptenodytes antarcticus Moreno and Mercerat 1891

†Paraptenodytes brodkorbi Simpson 1972

†Paraptenodytes robustus Ameghino 1894

Invalid names: Arthrodytes grandis Ameghino 1901 [synonym], Isotremornis nordenskjoldi Ameghino 1905 [synonym], Paraptenodytes crassus Ameghino 1905 [synonym], Paraptenodytes curtus Ameghino 1901 [synonym]

Invalid names: Isotremornis Ameghino 1905 [synonym], Metancylornis Ameghino 1905 [synonym]

G. †Pseudaptenodytes Simpson 1970

†Pseudaptenodytes macraei Simpson 1970

Invalid names: Pseudaptenodytes minor Simpson 1970 [nomen dubium]

G. Pygoscelis Wagler 1832

Pygoscelis adeliae Hombon and Jacquinot 1841 [Adelie penguin]

Pygoscelis antarcticus Forster 1781 [chinstrap penguin]

†Pygoscelis calderensis Acosta Hospitaleche et al. 2006

†Pygoscelis grandis Walsh and Suarez 2006

Pygoscelis papua Forester 1781 [gentoo penguin]

†Pygoscelis tyreei Simpson 1972

Invalid names: Catarrhactes Hombron and Jacquinot 1841 [synonym]

G. Spheniscus Brisson 1760

†Spheniscus chilensis Emslie and Correa 2003

Spheniscus demersus Linnaeus 1758 [African penguin]

Spheniscus humboldti Meyen 1834 [Humboldt penguin]

Spheniscus magellanicus Forster 1781 [Magellanic penguin]

†Spheniscus megaramphus Stucchi et al. 2003

Spheniscus mendiculus Sundevall 1871 [Galapagos penguin]

†Spheniscus muizoni Gohlich 2007

†Spheniscus urbinai Stucchi 2002

Invalid names: Anthropornithinae Simpson 1946 [empty], Apterodytes ictus Ameghino 1901 [nomen dubium], Delphinornis gracilis Wiman 1905 [nomen dubium], Ichtyopteryx Wiman 1905 [nomen dubium], Neculus Ameghino 1905 [nomen dubium], Orthopteryx Wiman 1905 [nomen dubium], Orthopteryx gigas Wiman 1905 [nomen dubium], Palaeeudyptinae Simpson 1946 [empty], Palaeoapterodytes Ameghino 1905 [nomen dubium], Spheniscinae Simpson 1946 [empty], Tereingaornis Scarlett 1983 [nomen dubium]

G. †Tonniornis Tambussi et al. 2006

†Tonniornis mesetaensis Tambussi et al. 2006

†Tonniornis minimum Tambussi et al. 2006

G. †Waimanu Jones et al. 2006

†Waimanu manneringi Jones et al. 2006

G. †Wimanornis Simpson 1971

†Wimanornis seymourensis Simpson 1971

Invalid names: Paraptenodytinae Simpson 1946 [empty]

Diagnosis

No diagnoses are available

Measurements

No measurements are available

Composition:

hydroxyapatiteo

Entire body:

yeso

Adult length:

10 to < 100o

Adult width:

1.0 to < 10o

Adult height:

1.0 to < 10o

Architecture:

compact or denseo

Ontogeny:

accretion, modification of partso

Grouping:

solitaryo

Environment:

marineo

Locomotion:

actively mobileo

Life habit:

ground dwellingo

Diet:

carnivoresubo

Reproduction:

oviparouso

Dispersal:

direct/internalo

Dispersal 2:

mobileo

Created:

2017-04-17 10:40:01

Modified:

2017-04-17 10:40:01

Source: subo = suborder, o = order

References: Benton 1983, Marsh 1875, Bush and Bambach 2015

Age range

Maximum range based only on fossils: base of the Selandian to the top of the Holocene or 61.60000 to 0.00000 Ma
Minimum age of oldest fossil (stem group age): 58.7 Ma

Collections (177 total)

Time interval	Ma	Country or state	Original ID and collection number
Oldest occurrences
Early/Lower Paleocene - Middle Paleocene	66.0 - 58.7	New Zealand (Chatham Islands Territory)	Sphenisciformes indet., Kupoupou stilwelli (207006)
Selandian - Thanetian	61.6 - 56.0	New Zealand (South Island)	Crossvallia waiparensis (203390 203391 203392) Muriwaimanu tuatahi (175680) Sequiwaimanu rosieae (161199)
Selandian - Thanetian	61.6 - 56.0	New Zealand	Sphenisciformes indet., Waimanu manneringi (68822) Waimanu tuatahi (85839 85840 85841)
Late/Upper Paleocene	58.7 - 55.8	New Zealand (South Island)	Sphenisciformes indet., Kumimanu biceae, Kumimanu fordycei, Petradyptes stonehousei (190583)
Late/Upper Paleocene	58.7 - 55.8	Antarctica	Crossvallia unienwillia (74613)
Ypresian	56.0 - 47.8	Antarctica	Spheniscidae indet. (32360 52067 120576) Spheniscidae indet., Anthropornis grandis, Archaeospheniscus wimani (100414) Sphenisciformes indet. (100188 175618)
Ypresian	56.0 - 47.8	Antarctica (Seymour Island)	Anthropornis grandis (63550) Palaeeudyptes gunnari (63549)
Ypresian - Lutetian	56.0 - 41.3	Antarctica	Anthropornis grandis (38816)
Ypresian - Lutetian	56.0 - 41.3	Antarctica (Seymour Island)	Delphinornis sp., Palaeeudyptes gunnari, Archaeospheniscus wimani, Delphinornis larseni, Mesetaornis sp. (63551)
Eocene	56.0 - 33.9	Antarctica	Sphenisciformes indet. (207756)
Eocene	56.0 - 33.9	Chile	Spheniscidae indet. (99572)
Waipawan - Mangaorapan	55.8 - 49.5	New Zealand	Kaiika maxwelli (121125)
Lutetian	47.8 - 41.3	Antarctica	Palaeeudyptes gunnari (211926)
Lutetian	47.8 - 41.3	Peru (Ica)	Perudyptes devriesi (73817)
Lutetian	47.8 - 41.3	Antarctica (Seymour Island)	Spheniscidae indet. (176083)
Lutetian - Bartonian	47.8 - 38.0	Antarctica (Seymour Island)	Anthropornis grandis, Palaeeudyptes klekowskii, Palaeeudyptes gunnari, Mesetaornis polaris, Marambiornis exilis, Anthropornis nordenskjoeldi, Delphinornis wimani (53666)
Middle Eocene - Late/Upper Eocene	47.8 - 33.9	Chile	Spheniscidae indet., Palaeeudyptes sp. (99479)
Bartonian	41.3 - 38.0	Argentina (Tierra del Fuego)	Pansphenisciformes indet. (59977)
Bartonian	41.3 - 38.0	Antarctica (Seymour Island)	Anthropornis grandis (207777) Delphinornis sp. (63552)
Bartonian	41.3 - 38.0	Antarctica	Anthropornis nordenskjoldii, Pachypteryx grandis, Eosphaeniscus gunnari, Delphinornis larsenii, Ichtyopteryx gracilis (77358)