King shags Leucocarbo carunculatus, photographed by Kristian Stahl.

Belongs within: Neoaves.
Contains: Sulidae, Phalacrocorax.

The Sulida or Suloidea are the clade including Sulidae (gannets and boobies), Anhingidae (darters) and Phalacrocoracidae (shags or cormorants). All members of the clade are diving fish-eating birds. The darters or anhingas of the Anhingidae, all living species of which are placed in the single genus Anhinga, are a pantropical group of birds with particularly long and flexible necks; the neck is held back against the body when approaching prey, then darted forward to impale the prey on their long straight beak.

Synapomorphies (from Mayr 2004): Os palatinum a flat horizontal plate; vomer absent; processus parocciptales protruding in caudal direction; quadratum with processus orbitalis reduced; eggs incubated beneath feet; sky-pointing/wing-waving and hop displays present.

<==Sulida [Sulae, Suloidea]
    |--Sulidae LZ07
    `--+--Borvocarbo Mourer-Chauviré et al. 2004 M05
       `--Phalacrocoracoidea LZ07
            |--Anhingidae [Anhinginae, Plotinae, Plottinae, Ptynginae] LZ07
            |    |--Protoplotus beauforti VR72
            |    |--Macranhinga paranensis M04
            |    `--Anhinga Ridgway 1887 M02 (see below for synonymy)
            |         |  i. s.: A. laticeps VR72
            |         |         A. melanogaster Pennant 1769 R85 [=Plotus melanogaster S66]
            |         |         A. nana VR72
            |         |         A. pannonica (Lambrecht 1916) [=Plotus pannonicus] M02
            |         |         A. parva VR72
            |         |--A. rufa (Dausin 1802) JT12, WS48 [=Plotus rufus WS48, A. melanogaster rufa HR96]
            |         `--+--A. anhinga JT12 [=Plotus anhinga G30]
            |            `--A. novaehollandiae (Gould 1847) JT12, WS48 (see below for synonymy)
            `--Phalacrocoracidae [Halieidae] LZ07
                 |  i. s.: Nannopterum harrisi FP64
                 |         Nectornis Cheneval 1984 M02
                 |           |--*N. miocaenus (Milne-Edwards 1867) (see below for synonymy) M02
                 |           `--N. anatolicus M02
                 |--Phalacrocorax LZ07
                 `--Leucocarboninae HWT01
                      |--Stictocarbo featherstoni (Buller 1873) HWT01, WH02
                      `--Leucocarbo Bonaparte 1857 B94
                           |--L. atriceps (King 1828) R85
                           |    |--L. a. atriceps M03
                           |    `--L. a. nivalis M03
                           |--L. campbelli (Filhol 1878) WH02 [=Nesocarbo campbelli HWT01]
                           |--L. carunculatus (Gmelin 1789) WH02 [incl. L. chalconotus (Gray 1845) WH02, R85]
                           |--L. colensoi (Buller 1888) WH02 [=Euleucocarbo colensoi HWT01]
                           |--L. onslowi (Forbes 1893) WH02 [=Euleucocarbo onslowi HWT01]
                           |--L. purpurascens (Brandt 1837) WH02 (see below for synonymy)
                           `--L. ranfurlyi (Ogilvie-Grant 1901) WH02 [=Euleucocarbo ranfurlyi HWT01]

Anhinga Ridgway 1887 M02 [incl. Plotus Linné 1766 B94, Plottus (l. c.) B94, Ptynx Möhring 1752 (pre-Linnean) non Blyth 1840 B94]

Anhinga novaehollandiae (Gould 1847) JT12, WS48 [=Plotus novaehollandiae WS48, A. melanogaster novaehollandiae HR96, A. rufa novaehollandiae WS48; incl. P. novaehollandiae derbyi Mathews 1912 WS48]

Leucocarbo purpurascens (Brandt 1837) WH02 [=L. atriceps purpurascens R85, Notocarbo purpurascens HWT01]

*Nectornis miocaenus (Milne-Edwards 1867) [=Graculus miocaenus, Oligocorax miocaenus, Phalacrocorax miocaenus] M02

*Type species of generic name indicated


[B94] Bock, W. J. 1994. History and nomenclature of avian family-group names. Bulletin of the American Museum of Natural History 222: 1-281.

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

[G30] Graham, G. M. 1930. The natural history of the Victoria Nyanza. Proceedings of the Linnean Society of London 141: 43-44.

[HR96] Heather, B. D., & H. A. Robertson. 1996. The Field Guide to the Birds of New Zealand. Viking: Auckland.

[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.

[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.

[LZ07] Livezey, B. C., & R. L. Zusi. 2007. Higher-order phylogeny of modern birds (Theropoda, Aves: Neornithes) based on comparative anatomy. II. Analysis and discussion. Zoological Journal of the Linnean Society 149 (1): 1-95.

[M04] Mayr, G. 2004. Tertiary plotopterids (Aves, Plotopteridae) and a novel hypothesis on the phylogenetic relationships of penguins (Spheniscidae). Journal of Zoological Systematics and Evolutionary Research 43: 61-71.

[M05] Mayr, G. 2005. The Paleogene fossil record of birds in Europe. Biological Reviews 80: 515-542.

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

[M03] Morcombe, M. 2003. Field Guide to Australian Birds, 2nd ed. Steve Parish Publishing.

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

[S66] Schlegel, H. 1866. Communication from, on mammals and birds collected in Madagascar. Proceedings of the Zoological Society of London 1866: 419-426.

[VR72] Vickers Rich, P. 1972. A fossil avifauna from the Upper Miocene Beglia Formation of Tunisia. Notes du Service Géologique 35: 29-66.

[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.

[WH02] Worthy, T. H., & R. N. Holdaway. 2002. The Lost World of the Moa: Prehistoric life of New Zealand. Indiana University Press: Bloomington (Indiana).

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