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 1760 CC10 (see below for synonymy)
    |--P. gaimardi BKB15 [=P. (Sticticarbo) gaimardi HD09]
    `--+--+--P. melanoleucos (Vieillot 1817) BKB15, CC10 (see below for synonymy)
       |  |    |--P. m. melanoleucos (see below for synonymy) CC10
       |  |    |--P. m. brevicauda Mayr 1931 CC10
       |  |    `--P. m. brevirostris Gould 1837 (see below for synonymy) CC10
       |  `--P. pygmaeus (Pallas 1773) BKB15, CS77 (see below for synonymy)
       `--+--+--P. aristotelis (Linnaeus 1761) BKB15, M02 (see below for synonymy)
          |  |    |--P. a. aristotelis CS77
          |  |    |--P. a. desmarestii (Payraudeau 1826) CS77
          |  |    `--P. a. riggenbachi Hartert 1923 CS77
          |  `--+--+--P. penicillatus BKB15
          |     |  `--+--P. urile (Gmelin 1789) BKB15, I92
          |     |     `--+--P. capillatus BKB15
          |     |        `--P. pelagicus BKB15
          |     `--+--P. neglectus BKB15
          |        `--+--P. nigrogularis Ogilvie-Grant & Forbes 1899 BKB15, CS77
          |           `--+--+--*P. carbo (Linnaeus 1758) CC10, BKB15, CC10 (see below for synonymy)
          |              |  |    |--P. c. carbo CC10
          |              |  |    |--P. c. hanedae Kuroda 1925 CC10
          |              |  |    |--P. c. lucidus (Lichtenstein 1823) CC10 [incl. P. carbo patricki Williams 1966 CS77]
          |              |  |    |--P. c. lugubris Rüppell 1845 CC10
          |              |  |    |--P. c. maroccanus Hartert 1906 CC10
          |              |  |    |--P. c. novaehollandiae Stephens in Shaw 1826 CC10 (see below for synonymy)
          |              |  |    `--P. c. sinensis (Blumenbach 1798) CC10
          |              |  `--P. capensis BKB15
          |              `--+--+--P. featherstoni Buller 1873 BKB15, 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. varius CC10 (see below for synonymy)
          |                            |--P. v. hypoleucos (Brandt 1837) [=Carbo hypoleucos] CC10
          |                            `--P. v. nitidus Serventy 1940 WS48
          `--+--+--P. atriceps (King 1828) BKB15, R85 [=Leucocarbo atriceps R85, *Notocarbo atriceps CC10]
             |  |    |--P. a. atriceps M03
             |  |    `--'Leucocarbo’ a. nivalis M03
             |  `--+--P. harrisi BKB15
             |     `--+--P. auritus (Lesson 1831) BKB15, CC10 [=Carbo auritus CC10, Graucalus auritus CC10]
             |        `--P. brasilianus BKB15
             `--+--P. magellanicus (Gmelin 1789) BKB15, CC10 (see below for synonymy)
                `--+--P. bougainvillii BKB15 [=Carbo bougainvillii CC10, *Leucocarbo bougainvillii CC10, HD09]
                   `--+--P. ranfurlyi Ogilvie-Grant 1901 BKB15, CC10 (see below for synonymy)
                      |--+--P. albiventer BKB15 [=P. (Leucocarbo) albiventer HD09]
                      |  |--P. georgianus BKB15
                      |  `--P. purpurascens (Brandt 1837) BKB15, CC10 (see below for synonymy)
                      `--+--+--P. campbelli (Filhol 1878) BKB15, CC10 (see below for synonymy)
                         |  `--P. colensoi Buller 1888 BKB15, CC10 (see below for synonymy)
                         `--+--P. chalconotus (Gray 1845) BKB15, CC10 (see below for synonymy)
                            `--P. onslowi Forbes 1893 BKB15, CC10 (see below for synonymy)

Phalacrocorax incertae sedis:
  P. africanus (Gmelin 1788) [=Pelecanus africanus, Graculus africanus] CC10
    |--P. a. africanus CS77
    `--P. a. pictilis Bangs 1918 CS77
  P. carunculatus (Gmelin 1789) JT12, CC10 (see below for synonymy)
  P. coronatus (Wahlberg 1855) CS77
  P. filamentosus T89
  P. fuscicollis JT12
  P. gouldi HD09
  P. ibericus Villalta 1963 [=P. ibericum] M02
  P. imperialis King 1831 CC10
  P. intermedius (Milne-Edwards 1867) (see below for synonymy) M02
  P. lautus Kuročkin & Ganea 1972 M02
  ‘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. niger JT12 [incl. Carbo javanicus CC10, *Graculus javanicus CC10]
  P. nycthemerus Cabanis 1855 (n. d.) CC10
  P. olivaceus M03
  P. perspicillatus FP64
  P. serdicensis Burčak-Abramovič & Nikolov 1984 M02
  P. verrucosus [=P. (Leucocarbo) verrucosus] HD09
  P. vigua S18
  P. wetmorei FP64

Nomen nudum: Carbo risgoviensis Fraas in Engel 1908 [=Phalacrocorax risgoviensis] M02

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

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

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

*Phalacrocorax carbo (Linnaeus 1758) CC10, BKB15, CC10 [=Pelecanus carbo CC10, *Carbo carbo CC10, *Ecmeles carbo CC10, Graculus carbo CC10, *Graucalus carbo CC10, *Halieus carbo CC10, *Hydrocorax carbo CC10]

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

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

Phalacrocorax chalconotus (Gray 1845) BKB15, CC10 [=Gracalus chalconotus CC10, Euleucocarbo chalconotus CC10, Graculus chalconotus CC10, Hypoleucus chalconotus CC10, Phalacrocorax (Leucocarbo) carunculatus chalconotus CC10; incl. Graculus glaucus Riechenbach 1850 CC10, Phalacrocorax glaucus CC10, Ph. huttoni Buller 1888 CC10, H. huttoni CC10, Ph. stewarti Ogilvie-Grant 1898 CC10, Hypoleucus campbelli stewarti CC10, Ph. (Leucocarbo) stewarti HD09]

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

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

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

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 magellanicus (Gmelin 1789) BKB15, CC10 [=Pelecanus magellanicus CC10, Ph. (Leucocarbo) magellanicus HD09]

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

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

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

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 purpurascens CC10, Hypoleucus atriceps purpurascens CC10, Leucocarbo atriceps purpurascens CC10, Notocarbo purpurascens HWT01, Phalacrocorax (Leucocarbo) albiventer purpurascens CC10, P. atriceps purpurascens CC10; incl. P. traversi Rothschild 1898 CC10, Hypoleucus carunculatus traversi CC10, P. atriceps traversi CC10, P. (Leucocarbo) traversi HD09]

Phalacrocorax pygmaeus (Pallas 1773) BKB15, CS77 [=Pelecanus pygmaeus CC10, *Halietor pygmaeus CC10, *Microcarbo pygmaeus CC10]

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

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

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

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

*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 4^th 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.

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

[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 3^rd ed. Whitcombe and Tombs Limited.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[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.

[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.

Last updated: 1 December 2021.