Procellariidae

Grey petrel Procellaria cinerea, copyright JJ Harrison.


Belongs within: Procellariiformes.
Contains: Pterodroma, Puffinus.

The Procellariidae, petrels and related forms, are a group of medium-sized to huge seabirds found primarily in cooler waters of the world, particularly in the Southern Hemisphere. They have historically been divided between two subfamilies, the Fulmarinae and Procellariinae, based on the fused or unfused condition respectively of the lachrymal bone in the skull, but recent phylogenetic analyses have suggested that the 'fulmarines' are paraphyletic with regard to the 'procellariines'. A position within the Procellariidae has also been indicated for the diving petrels of the genus Pelecanoides, small, stocky birds with short wings and widely placed, powerful legs that have often been placed in their own family.

Within the family, the more mobile, 'procellariine'-type bill is found in members of the genera Procellaria and Puffinus. The petrels of the genus Procellaria have more strongly hooked bills than the shearwaters of the genus Puffinus. Other members of the Procellariidae include the prions of the genus Pachyptila, small grey petrels that primarily feed on zooplankton. The giant petrels of the genus Macronectes are the largest members of the family and are aggressive predators as well as scavengers on other vertebrates in the southern oceans. The fulmars of the genus Fulmarus are whitish-grey birds whose two species inhabit the northern (F. glacialis) and southern (F. glacialoides) polar regions.

Characters (from Cramp & Simmons 1977): Body more or less ovate. Wings long and narrow; eleven primaries: p10 longest, p11 minute; 20–29 secondaries, short, diastataxic. Tail short, usually twelve feathers (occasionally 14 or 16). Bill heavy, sometimes more slender or broad; hooked at tip; horny sheath divided into plates; nostrils in dorsal tubes. Legs set far back, bird crouching on tarsi when on land, gait shuffling except in Macronectes. Tarsus laterally flattened, round in Pterodroma; lower part of tibia bare; three anterior toes webbed, hind toe rudimentary, elevated; nails sharp and curved. Oil gland feathered. Aftershaft small. Peculiar musky odour. Sexes similar in plumage; mostly black or grey above and white below or all dark. One moult per year, primaries moulting descendantly. Young semi-altricial and nidicolous, development slow; two downs, both long and fluffy, juvenile feathers growing immediately under second down. Juvenile usually like adult.

<==Procellariidae [Fulmaridae, Fulmarinae, Procellariinae, Wagellidae]
    |  i. s.: Argyrodyptes Ameghino 1905 S72
    |           `--*A. microtarsus Ameghino 1905 S72
    |         Prion Lacépède 1799 (n. d.) [=Priamphus Rafinesque 1815] CC10
    |         Eopuffinus kazachstanensis Nessov 1986 U93
    |--+--Halobaena Bonaparte 1856 BKB15, CC10 [=Zaprium Coues 1875 CC10]
    |  |    `--*H. caerulea (Gmelin 1789) (see below for synonymy) CC10
    |  `--Pachyptila Illiger 1811 BKB15, CC10 (see below for synonymy)
    |       |  i. s.: P. belcheri (Mathews 1912) (see below for synonymy) CC10
    |       |         P. crassirostris (Mathews 1912) (see below for synonymy) CC10
    |       |           |--P. c. crassirostris (see below for synonymy) CC10
    |       |           |--P. c. flemingi Tennyson & Bartle 2005 CC10
    |       |           `--P. c. pyramidalis Fleming 1939 [=P. (Pseudoprion) eatoni pyramidalis] CC10
    |       |         P. salax Olson 1985 B93
    |       |--P. turtur (Kuhl 1820) JT12, CC10 (see below for synonymy)
    |       `--+--P. vittata (Forster 1777) JT12, CC10 (see below for synonymy)
    |          `--+--P. desolata (Gmelin 1789) JT12, CC10 (see below for synonymy)
    |             `--P. salvini (Mathews 1912) JT12, CC10 (see below for synonymy)
    |                  |--P. s. salvini (see below for synonymy) CC10
    |                  `--P. s. macgillivrayi (Mathews 1912) CC10
    `--+--+--Lugensa Mathews 1942 CC10 [incl. Aphrodroma Olson 2000 BKB15, CC10]
       |  |    `--L. brevirostris (Lesson 1833) (see below for synonymy) CC10
       |  `--+--+--Thalassoica Reichenbach 1853 BKB15, CC10 [=Aeipetes Forbes 1882 CC10, Thalassoeca Salvin 1896 CC10]
       |     |  |    `--*T. antarctica (Gmelin 1789) (see below for synonymy) CC10
       |     |  `--Pagodroma Bonaparte 1856 JT12, CC10 [Pagodromidae]
       |     |       `--*P. nivea (Forster 1777) (see below for synonymy) CC10
       |     |            |--P. n. nivea (see below for synonymy) CC10
       |     |            `--P. n. major (Schlegel 1863) (see below for synonymy) CC10
       |     `--+--Daption Stephens in Shaw 1826 JT12, CC10 (see below for synonymy)
       |        |    `--*D. capense (Linnaeus 1758) (see below for synonymy) CC10
       |        |         |--D. c. capense [incl. Procellaria naevia Bonaparte 1857, P. pardela Oken 1816] CC10
       |        |         `--D. c. australe Mathews 1913 [=D. capensis australis] CC10
       |        `--+--Macronectes Richmond 1905 JT12, CC10 (see below for synonymy)
       |           |    |--*M. giganteus (Gmelin 1789) (see below for synonymy) CC10
       |           |    `--M. halli Mathews 1912 [=M. giganteus halli] CC10
       |           `--Fulmarus Stephens in Shaw 1826 JT12, CC10 (see below for synonymy)
       |                |--*F. glacialis (Linnaeus 1761) CC10, CS77 (see below for synonymy)
       |                |    |--F. g. glacialis [incl. F. glacialis auduboni Bonaparte 1857] CS77
       |                |    `--F. g. rodgersii Cassin 1862 CS77
       |                `--F. glacialoides (Smith 1840) (see below for synonymy) CC10
       `--+--+--Pterodroma BKB15
          |  `--Pelecanoides Lacépède 1799 BKB15, CC10 (see below for synonymy)
          |       |  i. s.: P. cymatotrypetes Olson 1984 U93
          |       |         P. miokuaka Worthy, Tennyson et al. 2007 CC10
          |       |--P. garnotii JT12 [=*Puffinuria garnotii CC10]
          |       `--+--*P. urinatrix (Gmelin 1789) CC10, JT12, CC10 (see below for synonymy)
          |          |    |--P. u. urinatrix (see below for synonymy) CC10
          |          |    |--P. u. berard (Gaimard 1823) (see below for synonymy) CC10
          |          |    |--P. u. chathamensis Murphy & Harper 1916 [=P. urinator chathamensis] CC10
          |          |    |--P. u. coppingeri Mathews 1912 CC10
          |          |    |--P. u. dacunhae Nicoll 1906 CC10
          |          |    `--P. u. exsul Salvin 1896 [=P. urinator exsul] CC10
          |          `--+--P. georgicus Murphy & Harper 1916 JT12, CC10 (see below for synonymy)
          |             `--P. magellani JT12 [=Puffinuria garnotii magellani CC10, Pe. (*Porthmornis) magellani CC10]
          `--+--+--Puffinus BKB15
             |  `--Pseudobulweria Mathews 1936 JT12, CC10
             |       |--*P. macgillivrayi (Gray 1859) CC10, CS77 [=Thalassidroma (Bulweria) macgillivrayi CC10]
             |       |--P. aterrima JT12 [=Pterodroma aterrima BKB15]
             |       |--P. becki JT12 [=P. rostrata becki CC10]
             |       `--P. rostrata (Peale 1848) CC10 (see below for synonymy)
             |            |--P. r. rostrata CC10
             |            `--P. r. trouessarti CC10
             `--+--Bulweria Bonaparte 1843 BKB15, CC10 [Bulweriidae]
                |    |--*B. bulwerii (Jardine & Selby 1828) (see below for synonymy) CC10
                |    |--B. bifax HSS13
                |    `--B. fallax Jouanin 1955 CS77
                `--Procellaria Linnaeus 1758 BKB15, CC10 (see below for synonymy)
                     |  i. s.: P. atlantica Gould 1844 [=Fulmarus atlanticus] CC10
                     |         P. fregata Linnaeus 1766 (n. d.) [=Fregetta fregata] CC10
                     |         P. fuliginosa Gmelin 1789 (n. d.) [=Oestrelata fuliginosa] CC10
                     |         P. ‘grisea’ Kuhl 1820 (n. d.) non Gmelin 1789 (see below for synonymy) CC10
                     |         P. incerta Schlegel 1863 [=Oestrelata incerta] CC10
                     |         P. lugens Kuhl 1820 (n. d.) CC10
                     |         P. melanopus Gmelin 1789 (n. d.) CC10
                     |         P. munda Kuhl 1820 (nom. rej.) [=Nectris munda] CC10
                     |         P. nigra Pallas 1769 (n. d.) CC10
                     |         P. velox Kuhl 1820 (n. d.) CC10
                     |--P. westlandica Falla 1946 BKB15, CC10 [=P. parkinsoni westlandica CC10]
                     `--+--P. cinerea Gmelin 1789 BKB15, CC10 (see below for synonymy)
                        `--+--P. parkinsoni Gray 1862 BKB15, CC10 (see below for synonymy)
                           `--+--*P. aequinoctialis Linnaeus 1758 CC10, BKB15, CC10 (see below for synonymy)
                              `--P. conspicillata Gould 1844 JT12, CC10 [=P. aequinoctialis conspicillata CC10]

*Bulweria bulwerii (Jardine & Selby 1828) [=Procellaria bulwerii, Aestrelata bulweri, Thalassidroma bulweri; incl. Procellaria anjinho Heineken 1829, Bulweria bulweri pacifica Mathews & Iredale 1915] CC10

Daption Stephens in Shaw 1826 JT12, CC10 [=Calopetes Sundevall 1873 CC10, Petrella Mathews 1914 CC10; Daptionidae]

*Daption capense (Linnaeus 1758) [=Procellaria capensis, *Calopetes capensis, Daption capenses (l. c.), Fulmarus (Daption) capensis, *Petrella capensis; incl. Procellaria puncata Ellman 1861] CC10

Fulmarus Stephens in Shaw 1826 JT12, CC10 [=Halohippus Billberg 1828 CC10, Rhantistes Kaup 1829 CC10, Wagellus Gray 1840 CC10; incl. Priocella Hombron & Jacquinot 1844 CC10]

*Fulmarus glacialis (Linnaeys 1761) CC10, CS77 [=Procellaria glacialis CC10, *Halohippus glacialis CC10, *Rhantistes glacialis CC10, *Wagellus glacialis CC10]

Fulmarus glacialoides (Smith 1840) [=Procellaria glacialoides, F. (Priocella) glacialoides, Priocella antarctica glacialoides, Thalassoeca glacialoides, Thalassoica glacialoides, Procellaria smithi Schlegel 1863; incl. Priocella antarctica addenda Mathews 1915, Fulmarus antarcticus Stephens in Shaw 1826 non Pro. antarctica Gmelin 1789, Pri. antarctica, *Pri. garnotii Jacquinot & Pucheran in Dumont d’Urville 1844, Procellaria garnotii, Thalassoica glacialoides polaris Bonaparte 1857, Procellaria tenuirostris Audubon 1839 non Temminck 1835, Fulmarus tenuirostris, Priocella tenuirostris, Thalassoeca tenuirostris, Thalassoica glacialoides tenuirostris] CC10

*Halobaena caerulea (Gmelin 1789) [=Procellaria caerulea, Fulmarus (Halobaena) caeruleus, Halobaena coerulea (l. c.), Pachyptila caerulea, Prion caerulea, Procellaria coerulea (l. c.), *Zaprium caeruleum; incl. Procellaria forsteri Smith 1840 non Latham 1790, Halobaena murphyi Brooks 1917, H. caerulea murphyi, Procellaria similis Forster in Lichtenstein 1844, H. caerulea victoriae Mathews 1916] CC10

Lugensa brevirostris (Lesson 1833) [=Procellaria brevirostris, Aphrodroma brevirostris, Bulweria brevirostris, Oestrelata brevirostris, Pterodroma brevirostris; incl. O. kidderi Coues 1875, *Aphrodroma kidderi, Pterodroma kidderi, Pt. lugens Mathews 1936, Bulweria lugens, *Lugensa lugens, Pt. kidderi okahia Mathews 1935, Lugensa lugens okahia, Pt. lugens okahia, Fulmarus (Cookilaria) unicolor Gray 1871, Procellaria unicolor, Rhantistes unicolor Bonaparte 1856, Pterodrome (l. c.) whitlocki Mathews 1946] CC10

Macronectes Richmond 1905 JT12, CC10 [=Ossifraga Hombron & Jacquinot 1844 non Wood 1835 CC10; Macronectidae]

*Macronectes giganteus (Gmelin 1789) [=Procellaria gigantea, Fulmarus (*Ossifraga) giganteus, *Ossifraga gigantea, Procellaria gigas (l. c.); incl. O. alba Potts 1874, Macronectes giganteus albus, Procellaria brasilia Kuhl 1820, M. giganteus dovei Mathews 1916, M. giganteus forsteri Mathews 1912, Procellaria maxima fusca Mathews 1933, P. ossifraga Forster in Lichtenstein 1844, Macronectes giganteus solanderi Mathews 1912, M. giganteus wilsoni Mathews 1912] CC10

Pachyptila Illiger 1811 BKB15, CC10 [incl. Attaprion Mathews 1933 CC10, Fulmariprion Mathews 1912 CC10, Heteroprion Mathews 1912 CC10, Prion Lesson 1828 non Lacépède 1799 CC10, Pseudoprion Coues 1866 CC10, Salviprion Mathews in Mathews & Hallstrom 1943 CC10; Pachyptilidae, Pachyptilinae]

Pachyptila belcheri (Mathews 1912) [=*Heteroprion belcheri; incl. H. belcheri falklandicus Mathews 1939, Pachyptila belcheri falklandica, H. belcheri lalfa Mathews 1939, Pa. (Heteroprion) belcheri orientalis Falla 1937, H. belcheri serventyi Mathews 1935, Pseudoprion turtur solanderi Mathews 1912, Procellaria turtur Mathews 1912 non Kuhl 1820] CC10

Pachyptila crassirostris (Mathews 1912) [=Pseudoprion turtur crassirostris, *Fulmariprion crassirostris] CC10

Pachyptila crassirostris crassirostris (Mathews 1912) [=P. (Pseudoprion) eatoni crassirostris; incl. Fulmariprion crassirostris antipodes Mathews & Hallstrom 1943] CC10

Pachyptila desolata (Gmelin 1789) JT12, CC10 [=Procellaria desolata CC10, Aestrelata desolata CC10, *Attaprion desolatus CC10, Daption desolatum CC10, Heteroprion desolatus CC10, Prion (Pseudoprion) desolata CC10, Prion desolatus CC10; incl. Heteroprion desolatus alexanderi Mathews & Iredale 1921 CC10, H. desolatus alter Mathews 1912 CC10, Pachyptila (Heteroprion) desolata alter CC10, Pa. desolata altera CC10, Pa. banksi Smith 1840 CC10, Attaprion descolatus banksi CC10, Pa. (Heteroprion) desolata banksi CC10, Heteroprion desolatus banksi CC10, Prion (Pseudoprion) banksii CC10, Procellaria banksii CC10, Prion dispar Bianchi 1913 CC10, Heteroprion desolatus dispar Iredlae 1938, Procellaria fasciata Bonnaterre 1791 CC10, H. desolatus georgia Mathews 1932 CC10, Attaprion desolatus georgia CC10, Pachyptila desolata georgia CC10, Pa. vittata georgicus Mathews 1933 CC10, Heteroprion desolatus heardi Mathews 1942 CC10, H. desolatus macquariensis Mathews 1912 CC10, Attaprion descolatus macquariensis CC10, H. desolatus mattingleyi Mathews 1912 CC10, A. desolatus mattingleyi CC10, H. desolatus peringueyi Mathews 1912 CC10, Prion rossi Bonaparte 1857 CC10]

Pachyptila salvini (Mathews 1912) JT12, CC10 [=Prion vittatus salvini CC10, Pa. (*Salviprion) salvini CC10]

Pachyptila salvini salvini (Mathews 1912) [=P. vittata salvini; incl. Heteroprion desolatus crozeti Mathews 1932, P. salvini crozeti, P. vittata crozeti, P. gouldi maui Mathews 1937, P. salvini muriwai Mathews & Hallstrom 1943, P. salvini whittelli Mathews 1938] CC10

Pachyptila turtur (Kuhl 1820) JT12, CC10 [=Procellaria turtur CC10, Prion (*Pseudoprion) turtur CC10; incl. Prion ariel Bonaparte 1857 CC10, Pachyptila ariel CC10, Procellaria ariel CC10, Pseudoprion ariel CC10, Ps. turtur armiger Mathews & Hallstrom 1943 CC10, Ps. turtur benchi Mathews & Hallstrom 1943 CC10, Prion brevirostris Gould 1855 CC10, Ps. turtur brevirostris CC10, Ps. turtur dertrum Mathews 1938 CC10, Ps. turtur eatoni Mathews 1912 CC10, Fulmariprion crassirostris eatoni CC10, Pachyptila (Pseudoprion) crassirostris eatoni CC10, Pa. turtur fallai Oliver 1930 CC10, Heteroprion belcheri fallai CC10, Pa. (Pseudoprion) turtur fallai CC10, Pseudoprion turtur huttoni Mathews 1912 CC10, Pa. (Pseudoprion) turtur huttoni CC10, Ps. turtur mangarei Mathews & Hallstrom 1943 CC10, Ps. turtur nova Mathews 1916 CC10, Ps. turtur oliveri Mathews 1932 CC10, Ps. turtur steadi Mathews 1932 CC10, Pachyptila turtur subantarctica Oliver 1955 CC10, Halobaena typica Bonaparte 1857 CC10]

Pachyptila vittata (Forster 1777) JT12, CC10 [=Procellaria vittata CC10, Pachyptila vittatus (l. c.) CC10, *Prion vittata CC10, *Prion vittatus CC10; incl. Prion australis Potts 1873 CC10, Pachyptila vittata balaena Mathews 1938 CC10, Procellaria forsteri Latham 1790 CC10, *Pa. forsteri CC10, Prion vittatus gouldi Mathews 1912 CC10, Pa. vittata gouldi CC10, Prion vittatus keyteli Mathews 1912 CC10, Pa. vittata keyteli CC10, Prion lamellirostris Bonaparte 1856 CC10, Procellaria latirostris Bonnaterre 1791 CC10, Pro. longirostris (l. c.) CC10, Prion magnirostris Gould 1862 CC10, Prion vittatus missus Mathews 1912 CC10, Pachyptila vittata missa CC10, Pa. vittata missus (l. c.) CC10]

*Pagodroma nivea (Forster 1777) [=Procellaria nivea, Daption niveum, Fulmarus (Pagrodroma) niveus, Thalassoica nivea] CC10

Pagodroma nivea major (Schlegel 1863) [=Procellaria nivea major non Pr. major Faber 1822; incl. Pa. confusa Mathews 1912] CC10

Pagodroma nivea nivea (Forster 1777) [incl. Pa. nivea alba Mathews 1928, Procellaria candida Peale 1848, Pa. nivea candida, Pa. nivea falklandica Mathews 1926, Procellaria nivea minor Schlegel 1863, Pa. nivea minor, Pa. novegeorgica Pagenstecher 1885, Pa. nivea novaegeorgica, Pa. nivea novageorgica, Pa. nivea pealei Mathews 1928] CC10

Pelecanoides Lacépède 1799 BKB15, CC10 [=Haladroma Illiger 1811 CC10, Halodroma Cuvier 1817 CC10, Onocralus Rafinesque 1815 CC10; incl. Pelagodyptes Murphy & Harper 1921 CC10, Porthmornis Murphy & Harper 1921 CC10, Puffinuria Lesson 1828 CC10; Haladromidae, Pelecanoidi, Pelecanoididae]

Pelecanoides georgicus Murphy & Harper 1916 JT12, CC10 [=Pele. georgica CC10, *Pelagodyptes georgicus CC10, Pele. urinatrix georgica CC10; incl. Pela. georgicus novus Mathews 1935 CC10]

*Pelecanoides urinatrix (Gmelin 1789) CC10, JT12, CC10 [=Procellaria urinatrix CC10, *Haladroma urinatrix CC10, *Halodroma urinatrix CC10, *Onocralus urinatrix CC10, Pelecanoides urinator CC10, Puffinuria urinatrix CC10]

Pelecanoides urinatrix berard (Gaimard 1823) [=Procellaria berard, Halodroma berardi (l. c.), Pelecanoides berardi (l. c.)] CC10

Pelecanoides urinatrix urinatrix (Gmelin 1789) [incl. Pe. urinatrix belcheri Mathews 1912, Procellaria tridactyla Forster in Lichtenstein 1844 non Pr. tridactyla Kuhl 1820, Pr. tridactylae Kuhl 1820] CC10

Procellaria Linnaeus 1758 BKB15, CC10 [=Cymatobolus Heine & Reichenow 1890 CC10, Cymbatobolus Checklist Committee 1990 CC10, Cymotobolus Mathews 1936 CC10, Majaqueus Reichenbach 1853 CC10; incl. Adamastor Bonarparte 1856 CC10, Priofinus Hombron & Jacquinot 18454 CC10]

*Procellaria aequinoctialis Linnaeus 1758 CC10, BKB15, CC10 [=*Cymatobolus aequinoctialis CC10, Fulmarus (*Majaqueus) aequinoctialis CC10, Puffinus aequinoctialis CC10; incl. Procellaria aequinoctialis brabournei Mathews 1912 CC10, Majaqueus fuliginosa Bonaparte 1856 CC10, Pr. fuliginosa Shaw in White 1790 non Gmelin 1789 CC10, Pr. aequinoctialis mixta Mathews 1912 CC10, Pr. nigra Forster in Lichtenstein 1844 non Pallas 1769 CC10, Pr. aequinoctialis steadi Mathews 1912 CC10]

Procellaria cinerea Gmelin 1789 BKB15, CC10 [=Cookilaria cinerea CC10, Adamastor cinerea (l. c.) CC10, A. cinereus CC10, *Priofinus cinereus CC10, Puffinus cinereus CC10; incl. Procellaria adamastor Schlegel 1863 CC10, Priofinus cinereus dydimus Mathews 1916 CC10, Pro. gelida Gmelin 1789 CC10, Adamastor gelidus CC10, Daption gelidum CC10, Fulmarus gelidus CC10, Rhantistes gelida CC10, Procellaria haesitata Forster in Lichtenstein 1844 CC10, *Adamastor haesitatus CC10, Aestrelata haesitata CC10, Procellaria melanura Bonnaterre 1791 CC10, Priofinus melanurus CC10, Pro. pallipes Mathews 1912 CC10, Adamastor typus Bonaparte 1857 CC10]

Procellaria ‘grisea’ Kuhl 1820 (n. d.) non Gmelin 1789 [=Aestrelata grisea, Fulmarus (Cookilaria) griseus] CC10

Procellaria parkinsoni Gray 1862 BKB15, CC10 [=Fulmarus (Majaqueus) parkinsoni CC10; incl. P. fuliginosa Gray 1862 nec Gmelin 1789 nec Shaw in White 1790 nec Forster in Lichtenstein 1844 CC10] CC10

Pseudobulweria rostrata (Peale 1848) CC10 [=Procellaria rostrata CC10, Aestrelata desolata rostrata CC10, Bulweria rostrata CC10, Oestrelata rostrata CC10, Pterodroma rostrata BKB15, Rhantistes rostrata CC10]

*Thalassoica antarctica (Gmelin 1789) [=Procellaria antarctica, *Aeipetes antarctica, Daption antarcticum, Fulmarus (Priocella) antarcticus, Priocella antarctica, *Thalassoeca antarctica; incl. Procellaria lugubris Tschudi 1856 non Bonaparte 1845] CC10

*Type species of generic name indicated

REFERENCES

[B93] Brooke, R. K. 1993. Annotated catalogue of the Aves type specimens in the South African Museum. Annals of the South African Museum 102 (10): 327–349.

[BKB15] Burleigh, J. G., R. T. Kimball & E. L. Braun. 2015. Building the avian tree of life using a large-scale, sparse supermatrix. Molecular Phylogenetics and Evolution 84: 53–63.

[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.

[CS77] Cramp, S., & K. E. L. Simmons (eds) 1977. Handbook of the Birds of Europe, the Middle East and North Africa: The Birds of the Western Palaearctic vol. 1. Ostrich to Ducks. Oxford University Press: Oxford.

[HSS13] Hirschfeld, E., A. Swash & R. Still. 2013. The World's Rarest Birds. Princeton University Press: Princeton (New Jersey).

[JT12] Jetz, W., G. H. Thomas, J. B. Joy, K. Hartmann & A. Ø. Mooers. 2012. The global diversity of birds in space and time. Nature 491: 444–448.

[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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