Reconstruction of Palaeeudyptes klekowskii, copyright A. C. Tatarinov.

Belongs within: Aequorlitornithes.
Contains: Spheniscinae.

The Spheniscidae, penguins, are a group of flightless, wing-propelled diving birds found primarily in cold waters of the Southern Hemisphere. As well as features related to their mode of swimming, members of the clade have marked supraorbital fossae on the skull for the salt glands (Mayr 2009).

Spheniscidae [Dypsicleidae, Sphenisciformes]
    |--Waimanu Jones, Ando & Fordyce in Slack et al. 2006 M09, CC10
    |    |--*W. manneringi Jones, Ando & Fordyce in Slack et al. 2006 CC10
    |    `--W. tuatahi Jones, Ando & Fordyce in Slack et al. 2006 CC10
    `--+--Delphinornis Wiman 1905 M09, S46
       |    |--*D. larsenii Wiman 1905 [=D. larseni] S46
       |    |--D. arctowskii Myrcha et al. 2002 M09
       |    `--D. gracilis Myrcha et al. 2002 M09
       `--+--Mesetaornis polaris Myrcha et al. 2002 M09
          |--Marambiornis exilis Myrcha et al. 2002 M09
          `--+--+--Eretiscus tonnii (Simpson 1981) M09
             |  `--Spheniscinae M09
             |--Palaeeudyptes Huxley 1859 M09, CC10 (see below for synonymy)
             |    |--*P. antarcticus Huxley 1859 S46
             |    |--P. gunnari (Wiman 1905) S72, A05 [=*Eosphaeniscus gunnari A05]
             |    |--P. klekowskii Myrcha et al. 1990 M09
             |    `--P. marplesi Brodkorb 1963 CC10
             `--Anthropornithinae S46
                  |--Pachydyptes Oliver 1930 S46
                  |    |--*P. ponderosus Oliver 1930 (see below for synonymy) CC10
                  |    `--P. simpsoni Jenkins 1974 U93
                  |--Arthrodytes Ameghino 1905 S46,
                  |    |--*A. grandis (Ameghino 1901) [=Paraptenodytes grandis] S72
                  |    `--A. andrewsi (Ameghino 1901) M09 [=Paraptenodytes andrewsi S72]
                  `--Anthropornis Wiman 1905 M09, S46
                       |--*A. nordenskjoeldi Wiman 1905 [=A. nordenskjoeldii] S46
                       `--A. grandis (Wiman 1905) M09

Spheniscidae incertae sedis:
  Duntroonornis Marples 1952 CC10
    `--*D. parvus Marples 1952 CC10
  Korora Marples 1952 CC10
    `--*K. oliveri Marples 1952 CC10
  Marplesornis Simpson 1972 CC10
    `--*M. novaezealandiae (Marples 1960) [=Palaeospheniscus novaezealandiae] CC10
  Tereingaornis Scarlett 1984 CC10
    `--*T. moisleyi Scarlett 1984 CC10
  Archaeospheniscus Marples 1952 [incl. Notodyptes Marples 1953] CC10
    |--*A. lowei Marples 1952 CC10
    |--A. lopdelli Marples 1952 CC10
    `--A. wimani (Marples 1953) [=*Notodyptes wimani] CC10
  Platydyptes Marples 1952 CC10
    |--*P. novaezealandiae (Oliver 1930) [=Pachydyptes novaezealandiae, Pa. novaezelandiae] CC10
    |--P. amiesi Marples 1952 CC10
    `--P. marplesi Simpson 1971 [=P. marplesigi] CC10
  Chubutodyptes Simpson 1970 S72
    `--*C. biloculata Simpson 1970 S72
  Neculus Ameghino 1905 (n. d.) S72
    `--*N. rothi Ameghino 1905 (n. d.) S72
  Palaeoapterodytes Ameghino 1905 (n. d.) [=Apterodytes Ameghino 1901 non Hermann 1783] S72
    `--*P. ictus (Ameghino 1901) (n. d.) [=*Apterodytes ictus] AH10
  Wimanornis seymourensis S72
  Anthropodyptes gilli S72
  Orthopteryx Wiman 1905 S46
    `--*O. gigas Wiman 1905 S46
  Pachypteryx Wiman 1905 S46
    `--*P. grandis Wiman 1905 S46
  Ichthyopteryx Wiman 1905 (n. d.) [=Ichtyopteryx] S46
    `--*I. gracilis Wiman 1905 [=*Ichtyopteryx gracilis] S46
  Catarractes Brisson 1760 (n. d.) CC10
  Penguinus Brünnich 1771 (n. d.) CC10
  Madrynornis mirandus Acosta Hospitaleche, Tambussi et al. 2007 AH10
  Paraptenodytes Ameghino 1891 S72 (see below for synonymy)
    |--*P. antarcticus (Moreno & Mercerat 1891) (see below for synonymy) S72
    |--P. brodkorbi Simpson 1972 S72
    `--P. robustus (Ameghino 1895) M09 (see below for synonymy)
  Palaeospheniscus Moreno & Mercerat 1891 S72 (see below for synonymy)
    |--*P. patagonicus Moreno & Mercerat 1891 (see below for synonymy) S72
    |--P. bergi Moreno & Mercerat 1891 (see below for synonymy) S72
    |--P. biloculata (Simpson 1970) AH10
    |--P. gracilis Ameghino 1899 (see below for synonymy) S72
    `--P. wimani (Ameghino 1905) [=*Perispheniscus wimani] S72
  Crossvallia unienwillia Tambussi et al. 2005 M09
  Tonniornis M09
    |--T. mesetaensis Tambussi et al. 2006 M09
    `--T. minimum Tambussi et al. 2006 M09
  Perudyptes devriesi Clarke et al. 2007 M09
  Icadyptes salsi Clarke et al. 2007 M09

*Pachydyptes ponderosus Oliver 1930 [=Anthropornis ponderosa, A. ponderosus, Pachydyptes ponderosa] CC10

Palaeeudyptes Huxley 1859 M09, CC10 [incl. Eosphaeniscus Wiman 1905 CC10, Eospheniscus S46; Palaeeudyptinae S46]

Palaeospheniscus Moreno & Mercerat 1891 S72 [incl. Paraspheniscus Ameghino 1905 S72, Perispheniscus Ameghino 1905 S72, Pseudospheniscus Ameghino 1905 S72; Palaeospheniscinae S46]

Palaeospheniscus bergi Moreno & Mercerat 1891 [=Pal. bergii, *Paraspheniscus bergi; incl. Pseudospheniscus concavus Ameghino 1905, Ps. convexus Ameghino 1905, *Ps. interplanus Ameghino 1905, Ps. planus Ameghino 1905, Palaeospheniscus planus Ameghino 1905, Pal. rothi Ameghino 1905] S72

Palaeospheniscus gracilis Ameghino 1899 [incl. Pal. medianus Ameghino 1905, Pal. nereius Ameghino 1901, Paraspheniscus nereius] S72

*Palaeospheniscus patagonicus Moreno & Mercerat 1891 [incl. P. affinis Ameghino 1905, P. intermedius Ameghino 1905, P. interruptus Ameghino 1905, P. menzbieri Moreno & Mercerat 1891] S72

Paraptenodytes Ameghino 1891 S72 [incl. Isotremornis Ameghino 1905 S72, Metancylornis Ameghino 1905 S72, Treleudytes Ameghino 1905 S72; Paraptenodytinae S46]

*Paraptenodytes antarcticus (Moreno & Mercerat 1891) [=Palaeospheniscus antarcticus; incl. *Isotremornis nordenskjoeldi Ameghino 1905] S72

Paraptenodytes robustus (Ameghino 1895) M09 [=Palaeospheniscus robustus S72, Perispheniscus robustus S72; incl. *Treleudytes crassa Ameghino 1905 S72, T. crassus S72, Paraptenodytes curtus Ameghino 1901 S72, *Metancylornis curtus S72]

*Type species of generic name indicated


[AH10] Acosta Hospitaleche, C. 2010. Taxonomic status of Apterodytes ictus Ameghino, 1901 (Aves; Sphenisciformes) from the Early Miocene of Patagonia, Argentina. Neues Jahrbuch für Geologie und Paläontologie—Abhandlungen 255 (3): 371–375.

[A05] Ameghino, F. 1905. Enumeración de los Impennes fósiles de Patagonia y de la Isla Seymour. Anales del Museo Nacional de Buenos Aires, serie 3, 6: 97–167, pls 1–8.

[CC10] Checklist Committee (OSNZ). 2010. Checklist of the Birds of New Zealand, Norfolk and Macquarie Islands, and the Ross Dependency, Antarctica, 4th ed. Ornithological Society of New Zealand and Te Papa Press: Wellington.

[M09] Mayr, G. 2009. Paleogene Fossil Birds. Springer.

[S46] Simpson, G. G. 1946. Fossil penguins. Bulletin of the American Museum of Natural History 87 (1): 1–99.

[S72] Simpson, G. G. 1972. Conspectus of Patagonian fossil penguins. American Museum Novitates 2488: 1–37.

[U93] Unwin, D. M. 1993. Aves. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 717–737. Chapman & Hall: London.

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