Phalacrocorax

 Double-crested cormorants Phalacrocorax auritus, photographed by Larry Thompson.

Belongs within: Sulida.

Phalacrocorax is a genus of cormorants or shags. There is no formal distinction between these names though there has been a historical tendency for the name ‘cormorant’ to apply to freshwater forms and ‘shag’ to marine birds. However, this usage has not been invariable and many species may be found in either habitat.

The circumscription of Phalacrocorax varies significantly between authors. Some would use it to cover all or almost all shags whereas others restrict it to the larger black-footed species related to the great cormorant P. carbo. A group of southern species with pink feet and white venters, including the imperial shag P. atriceps and the guanay cormorant P. bougainvillii, is often separated as the genus Leucocarbo.

<==Phalacrocorax Brisson 1760CC10 (see below for synonymy)
    |--P. gaimardiBKB15 [=P. (Sticticarbo) gaimardiHD09; incl. P. cirrigerF43]
    `--+--+--P. melanoleucos (Vieillot 1817)BKB15, CC10 (see below for synonymy)
       |  |    |--P. m. melanoleucos (see below for synonymy)CC10
       |  |    |--P. m. brevicauda Mayr 1931CC10
       |  |    `--P. m. brevirostris Gould 1837 (see below for synonymy)CC10
       |  `--P. pygmaeus (Pallas 1773)BKB15, CS77 [=Pelecanus pygmaeusCC10, *Halietor pygmaeusCC10, *Microcarbo pygmaeusCC10]
       `--+--+--P. aristotelis (Linnaeus 1761)BKB15, M02 (see below for synonymy)
          |  |    |--P. a. aristotelisCS77
          |  |    |--P. a. desmarestii (Payraudeau 1826)CS77
          |  |    `--P. a. riggenbachi Hartert 1923CS77
          |  `--+--+--P. penicillatusBKB15
          |     |  `--+--P. urile (Gmelin 1788)BKB15, S05 (see below for synonymy)
          |     |     `--+--P. capillatusBKB15
          |     |        `--P. pelagicusBKB15
          |     |             |--P. p. pelagicusS05
          |     |             `--P. p. robustus Sharpe 1899S05
          |     `--+--P. neglectusBKB15
          |        `--+--P. nigrogularis Ogilvie-Grant & Forbes 1899BKB15, CS77
          |           `--+--+--*P. carbo (Linnaeus 1758)CC10, BKB15, CC10 (see below for synonymy)
          |              |  |    |--P. c. carboCC10
          |              |  |    |--P. c. hanedae Kuroda 1925CC10
          |              |  |    |--P. c. lucidus (Lichtenstein 1823)CC10 [incl. P. carbo patricki Williams 1966CS77]
          |              |  |    |--P. c. lugubris Rüppell 1845CC10
          |              |  |    |--P. c. maroccanus Hartert 1906CC10
          |              |  |    |--P. c. novaehollandiae Stephens in Shaw 1826CC10 (see below for synonymy)
          |              |  |    `--P. c. sinensis (Blumenbach 1798)CC10
          |              |  `--P. capensisBKB15
          |              `--+--+--P. featherstoni Buller 1873BKB15, CC10 (see below for synonymy)
          |                 |  `--P. punctatus (Sparrman 1786)BKB15, CC10 (see below for synonymy)
          |                 |       |--P. p. punctatus (see below for synonymy)CC10
          |                 |       `--P. p. oliveri Mathews 1931 (see below for synonymy)CC10
          |                 `--+--P. sulcirostris (Brandt 1837)BKB15, CC10 (see below for synonymy)
          |                    `--+--P. fuscescens (Vieillot 1817)BKB15, WS48 (see below for synonymy)
          |                       `--P. varius (Gmelin 1789)BKB15, CC10 (see below for synonymy)
          |                            |--P. v. variusCC10 (see below for synonymy)
          |                            |--P. v. hypoleucos (Brandt 1837) [=Carbo hypoleucos]CC10
          |                            `--P. v. nitidus Serventy 1940WS48
          `--+--+--P. atriceps (King 1828)BKB15, R85 [=Leucocarbo atricepsR85, *Notocarbo atricepsCC10]
             |  |    |--P. a. atricepsM03
             |  |    `--'Leucocarbo’ a. nivalisM03
             |  `--+--P. harrisiBKB15
             |     `--+--P. auritus (Lesson 1831)BKB15, CC10 [=Carbo auritusCC10, Graucalus auritusCC10]
             |        `--P. brasilianusBKB15
             `--+--P. magellanicus (Gmelin 1789)BKB15, CC10 [=Pelecanus magellanicusCC10, Ph. (Leucocarbo) magellanicusHD09]
                `--+--P. bougainvilliiBKB15 [=Carbo bougainvilliiCC10, *Leucocarbo bougainvilliiCC10, HD09]
                   `--+--P. ranfurlyi Ogilvie-Grant 1901BKB15, CC10 (see below for synonymy)
                      |--+--P. albiventerBKB15 [=P. (Leucocarbo) albiventerHD09]
                      |  |--P. georgianusBKB15
                      |  `--P. purpurascens (Brandt 1837)BKB15, CC10 (see below for synonymy)
                      `--+--+--P. campbelli (Filhol 1878)BKB15, CC10 (see below for synonymy)
                         |  `--P. colensoi Buller 1888BKB15, CC10 (see below for synonymy)
                         `--+--P. chalconotus (Gray 1845)BKB15, CC10 (see below for synonymy)
                            `--P. onslowi Forbes 1893BKB15, CC10 (see below for synonymy)
Phalacrocorax incertae sedis:
  P. africanus (Gmelin 1788) [=Pelecanus africanus, Graculus africanus]CC10
    |--P. a. africanusCS77
    `--P. a. pictilis Bangs 1918CS77
  P. brasiliensisF43
  P. carunculatus (Gmelin 1789)JT12, CC10 (see below for synonymy)
  P. coronatus (Wahlberg 1855)CS77
  P. filamentosusT89
  P. fuscicollisJT12
  P. gouldiHD09
  P. graculusS05
  P. ibericus Villalta 1963 [=P. ibericum]M02
  P. imperialis King 1831CC10
  P. intermedius (Milne-Edwards 1867) (see below for synonymy)M02
  P. lautus Kuročkin & Ganea 1972M02
  ‘Hydrocorax’ leucogaster Vieillot 1817 [=Carbo leucogaster, Graculus leucogaster]CC10
  P. littoralis (Milne-Edwards 1863) [=Graculus littoralis, *Oligocorax littoralis]M02
  P. longipes (Tugarinov 1940) [=*Pliocarbo longipes]M02
  P. nigerJT12 [incl. Carbo javanicusCC10, *Graculus javanicusCC10, Phalacrocorax javanicaE42]
  P. nycthemerus Cabanis 1855 (n. d.)CC10
  P. olivaceusM03
  P. perspicillatusFP64
  P. serdicensis Burčak-Abramovič & Nikolov 1984M02
  P. verrucosus [=P. (Leucocarbo) verrucosus]HD09
  P. viguaS18
  P. wetmoreiFP64
Nomen nudum: Carbo risgoviensis Fraas in Engel 1908 [=Phalacrocorax risgoviensis]M02

Phalacrocorax Brisson 1760CC10 [=Carbo Lacépède 1799CC10, Ecmeles Gistel 1848CC10, Gracalus Gray 1845CC10, Graucalus Gray 1841 non Cuvier 1816CC10, Haliaeus Schinz 1825CC10, Halieus Illiger 1811CC10, Hydrocorax Vieillot 1816 non Brisson 1760CC10; incl. Botaurites Ammon 1918M02, Enygrotheres Heine & Reichenow 1890CC10, Euleucocarbo Voisin 1973CC10, Graculus Reichenbach 1850CC10, Halietor Heine 1860CC10, Hypoleucus Reichenbach 1853CC10, Leucocarbo Bonaparte 1857CC10, Microcarbo Bonaparte 1856CC10, Nesocarbo Voisin 1973CC10, Notocarbo Siegel-Causey 1988CC10, Oligocorax Brodkorb 1952M02, Paracorax Lambrecht 1933M02, Pliocarbo Tugarinov 1940M02, Stictocarbo Bonaparte 1855CC10; Carboninae, Halieinae, Phalacrocoracinae]

Phalacrocorax aristotelis (Linnaeus 1761)BKB15, M02 [=Pelecanus aristotelisM02; incl. Ph. destefanii Regàlia 1902M02, *Paracorax destefaniiM02, Ph. destefaniM02]

Phalacrocorax campbelli (Filhol 1878)BKB15, CC10 [=Urile campbelliCC10, Hypoleucus campbelliCC10, Leucocarbo (*Nesocarbo) campbelliCC10, Phalacrocorax carunculatus campbelliCC10]

*Phalacrocorax carbo (Linnaeus 1758)CC10, BKB15, CC10 [=Pelecanus carboCC10, *Carbo carboCC10, *Ecmeles carboCC10, Graculus carboCC10, *Graucalus carboCC10, *Halieus carboCC10, *Hydrocorax carboCC10; incl. Carbo cormoranus Smirnow 1901S05, Phalacrocorax cormoranusE42]

Phalacrocorax carbo novaehollandiae Stephens in Shaw 1826CC10 [=Gaculus (l. c.) novaehollandiaeCC10; incl. Ph. carboides Gould 1838CC10, Gracalus carboidesCC10, Graculus carboidesCC10, Graucalus carboidesCC10, Pelecanus carboidesCC10, Phalacrocorax novaehollandiae carboidesCC10, Pe. major Ellman 1861CC10, Ph. novaezealandiae var. major Forbes 1892CC10, Carbo carbo steadi Mathews & Iredale 1913CC10, Ph. novaehollandiae steadiCC10, C. carbo westralis Mathews 1912WS48]

Phalacrocorax carunculatus (Gmelin 1789)JT12, CC10 [=Pelecanus carunculatusCC10, Graculus carunculatusCC10, Graucalus carunculatusCC10, Hypoleucus carunculatusCC10, Leucocarbo (*Euleucocarbo) carunculatusCC10; incl. Pe. cirrhatus Gmelin 1789CC10, Gracalus cirrhatusCC10, Graculus cirrhatusCC10, Graucalus cirrhatusCC10, Hydrocorax cirratusCC10, Pelecanus cirratusCC10, Phalacrocorax cirrhatusCC10, Ph. finschii Buller 1876 non Sharpe 1875CC10]

Phalacrocorax chalconotus (Gray 1845)BKB15, CC10 [=Gracalus chalconotusCC10, Euleucocarbo chalconotusCC10, Graculus chalconotusCC10, Hypoleucus chalconotusCC10, Phalacrocorax (Leucocarbo) carunculatus chalconotusCC10; incl. Graculus glaucus Riechenbach 1850CC10, Phalacrocorax glaucusCC10, Ph. huttoni Buller 1888CC10, H. huttoniCC10, Ph. stewarti Ogilvie-Grant 1898CC10, Hypoleucus campbelli stewartiCC10, Ph. (Leucocarbo) stewartiHD09]

Phalacrocorax colensoi Buller 1888BKB15, CC10 [=Phalacrocorax colensoiCC10, Euleucocarbo colensoiCC10, Hypoleucus campbelli colensoiCC10, Phalacrocorax (Leucocarbo) campbelli colensoiCC10, P. carunculatus colensoiCC10]

Phalacrocorax featherstoni Buller 1873BKB15, CC10 [=Graculus featherstoniCC10, Leucocarbo featherstoniCC10, Phalacrocorax (Stictocarbo) punctatus featherstoniCC10]

Phalacrocorax fuscescens (Vieillot 1817)BKB15, WS48 [=Hydrocorax fuscescensWS48; incl. Carbo gouldi tunneyi Mathews 1912WS48]

Phalacrocorax intermedius (Milne-Edwards 1867) [=Graculus intermedius, Miocorax intermedius; incl. *Botaurites avitus Ammon 1918, Ardea brunhuberi Ammon 1918, Phalacrocorax brunhuberi, P. praecarbo Ammon 1918]M02

Phalacrocorax melanoleucos (Vieillot 1817)BKB15, CC10 [=Hydrocorax melanoleucosCC10, Gracalus melanoleucosCC10, Graculus melanoleucosCC10, Microcarbo melanoleucosCC10, Phalacrocorax melanoleucusCC10]

Phalacrocorax melanoleucos brevirostris Gould 1837 [=Carbo brevirostris, Gracalus brevirostris, Graculus brevirostris, Haliaeus brevirostris, Halietor melanoleucos brevirostris, Halieus brevirostris, Microcarbo melanoleucos brevirostris; incl. Phalacrocorax finschii Sharpe 1875, Graculus finschi, Carbo flavagula Peale 1848]CC10

Phalacrocorax melanoleucos melanoleucos (Vieillot 1817) [incl. Ph. flavirhynchus Gould 1838, Graucalus flavirostris Gray in Dieffenbach 1843, Pelecanus flavirostris, Phalacrocorax flavirostris]CC10

Phalacrocorax onslowi Forbes 1893BKB15, CC10 [=Euleucocarbo onslowCC10, Hypoleucus carunculatus onslowiCC10, Leucocarbo onslowiCC10, Phalacrocorax (Leucocarbo) carunculatus onslowiCC10; incl. P. rothschildi Forbes 1893CC10]

Phalacrocorax punctatus (Sparrman 1786)BKB15, CC10 [=Pelicanus (l. c.) punctatusCC10, *Enygrotheres punctatusCC10, Gracalus puncatusCC10, Graculus punctatusCC10, Graucalus punctatusCC10, Leucocarbo punctatusCC10, *Stictocarbo punctatusCC10]

Phalacrocorax punctatus oliveri Mathews 1931 [=Leucocarbo punctatus oliveri, Stictocarbo punctatus oliveri, S. steadi Oliver 1930 non Carbo carbo steadi Mathews & Iredale 1913, Phalacrocorax (Stictocarbo) punctatus steadi]CC10

Phalacrocorax punctatus punctatus (Sparrman 1786) [incl. Hydrocorax dilophus Vieillot 1817, Pelecanus naevius Gmelin 1789, Graculus naevius, Phalacrocorax naevius, Stictocarbo punctatus sassi Mathews 1930]CC10

Phalacrocorax purpurascens (Brandt 1837)BKB15, CC10 [=Carbo purpurascensCC10, Hypoleucus atriceps purpurascensCC10, Leucocarbo atriceps purpurascensCC10, Notocarbo purpurascensHWT01, Phalacrocorax (Leucocarbo) albiventer purpurascensCC10, P. atriceps purpurascensCC10; incl. P. traversi Rothschild 1898CC10, Hypoleucus carunculatus traversiCC10, P. atriceps traversiCC10, P. (Leucocarbo) traversiHD09]

Phalacrocorax ranfurlyi Ogilvie-Grant 1901BKB15, CC10 [=Euleucocarbo ranfurlyiCC10, Hypoleucus campbelli ranfurlyiCC10, Leucocarbo ranfurlyiCC10, Phalacrocorax (Leucocarbo) campbelli ranfurlyiCC10, P. carunculatus ranfurlyiCC10]

Phalacrocorax sulcirostris (Brandt 1837)BKB15, CC10 [=Carbo sulcirostrisCC10, Graculus sulcirostrisCC10; incl. C. purpuragula Peale 1848CC10, Mesocarbo ater purpuragulaCC10, Me. sulcirostris purpuragulaCC10, Phalacrocorax purpuragulaCC10, Microcarbo stictocephalus Bonaparte 1857CC10, Graculus stictocephalusCC10]

Phalacrocorax urile (Gmelin 1788)BKB15, S05 [=Pelecanus urileS05; incl. Ph. bicristatus Pall. 1826S05, Pe. violaceus Gmelin 1788S05, Ph. violaceusS05]

Phalacrocorax varius (Gmelin 1789)BKB15, CC10 [=Pelecanus variusCC10, Gracalus variusCC10, Graculus variusCC10, Graucalus variusCC10, *Hypoleucus variusCC10]

Phalacrocorax varius varius (Gmelin 1789)CC10 [incl. Carbo fucosus Peale 1848CC10, C. varius perthi Mathews 1912WS48, Pelecanus pica Forster in Lichtenstein 1844CC10]

*Type species of generic name indicated

References

[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.

[E42] Ewer, W. 1842. List of a collection of birds from India. Proceedings of the Zoological Society of London 10: 91–93.

[FP64] Fisher, J., & R. T. Peterson. 1964. The World of Birds: A comprehensive guide to general ornithology. Macdonald: London.

[F43] Fraser, L. 1843. On the collection of birds brought to England by Mr. Bridges. Proceedings of the Zoological Society of London 11: 108–121.

[HWT01] Holdaway, R. N., T. H. Worthy & A. J. D. Tennyson. 2001. A working list of breeding bird species of the New Zealand region at first human contact. New Zealand Journal of Zoology 28: 119–187.

[HD09] Hutton, F. W., & J. Drummond. 1909. The Animals of New Zealand: An account of the Dominion’s air-breathing vertebrates 3rd ed. Whitcombe and Tombs Limited.

[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.

[M03] Mayr, G. 2003. The phylogenetic affinities of the shoebill (Balaeniceps rex). Journal für Ornithologie 144: 157–175.

[M02] Mlíkovský, J. 2002. Cenozoic Birds of the World. Part 1: Europe. Ninox Press: Praha.

[R85] Robertson, C. J. R. (ed.) 1985. Reader’s Digest Complete Book of New Zealand Birds. Reader’s Digest: Sydney.

[S05] Schalow, H. 1905. Die Vögel der Arktis. In: Römer, F., & F. Schaudinn (eds) Fauna Arctica. Eine Zusammenstellun der arktischen Tierformen, mit besonder Berücksichtigung des Spitzbergen-Gebietes auf Grund der Ergebnisse der Deutschen Expedition in das Nördliche Eismeer im Jahre 1898 vol. 4 pp. 79–288. Gustav Fischer: Jena.

[S18] Stone, W. 1918. Birds of the Panama Canal Zone, with special reference to a collection made by Mr. Lindsey L. Jewel. Proceedings of the Academy of Natural Sciences of Philadelphia 70: 239–280.

[T89] Takeshita, N. 1989. Nihon no Yachoo. Kogakukan: Tokyo.

[WS48] Whittell, H. M., & D. L. Serventy. 1948. A systematic list of the birds of Western Australia. Public Library, Museum and Art Gallery of Western Australia, Special Publication 1: 1–126.

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