Red-footed booby Sula sula, copyright Drew Avery.

Belongs within: Aequornithia.
Contains: Phalacrocorax.

The Sulida or Suloidea are the clade including Sulidae (gannets and boobies), Anhingidae (darters) and Phalacrocoracidae (shags or cormorants). All members of this 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. The shags or cormorants of the Phalacrocoracidae have relatively short necks and beaks with a hooked tip and capture prey by diving from the surface and pursuing it underwater. The late Oligocene Oligocrax stoeffelensis resembled a small cormorant in overall appearance and may have represented either a stem-group phalacrocoracid or a stem member of the Phalacrocoracidae + Anhingidae clade (Mayr 2009).

The Sulidae contain the gannets and boobies, marine birds that capture fish by plunge diving. Sulids nest in colonies, sometimes very densely. The gannets of the genus Morus are found in temperate waters of the Northern and Southern Hemispheres, while the boobies of the genus Sula are tropical in distribution. Abbott's booby Papasula abbotti is an endangered species endemic to Christmas Island in the eastern Indian Ocean.

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]
    |--+--Protoplotus beauforti Lambrecht 1931 M09
    |  `--Phalacrocoracoidea M09
    |       |  i. s.: Piscator Harrison & Walker 1976 M09, M02
    |       |           `--*P. tenuirostris Harrison & Walker 1976 M02
    |       |--Borvocarbo Mourer-Chauviré et al. 2004 M05
    |       |    `--B. guilloti Mourer-Chauviré et al. 2004 M09
    |       `--+--Anhingidae [Anhinginae, Plotinae, Plottinae, Ptynginae] JT12
    |          |    |--Macranhinga paranensis M04
    |          |    `--Anhinga Brisson 1760 CC10 (see below for synonymy)
    |          |         |  i. s.: A. laticeps VR72
    |          |         |         A. melanogaster Pennant 1769 CC10 [=Plotus melanogaster S66]
    |          |         |           |--A. m. melanogaster CC10
    |          |         |           `--A. m. vulsini Bangs 1918 CC10
    |          |         |         A. nana VR72
    |          |         |         A. pannonica (Lambrecht 1916) [=Plotus pannonicus] M02
    |          |         |         A. parva VR72
    |          |         |         A. subvolans (Brodkorb 1956) U93
    |          |         |--A. rufa (Dausin 1802) JT12, CC10 (see below for synonymy)
    |          |         `--+--*A. anhinga CC10, JT12 [=Plotus anhinga CC10]
    |          |            |    |--A. a. anhinga E52
    |          |            |    `--A. a. leucogaster E52
    |          |            `--A. novaehollandiae (Gould 1847) JT12, CC10 (see below for synonymy)
    |          `--+--Oligocrax stoeffelensis (Mayr 2007) PB15, M09 [=Borvocarbo stoeffelensis M09]
    |             `--Phalacrocoracidae [Halieidae, Leucocarboninae] JT12
    |                  |--Nannopterum harrisi CS77
    |                  |--Phalacrocorax JT12
    |                  |--Limicorallus saiensis Kurochkin 1968 M09
    |                  `--Nectornis Cheneval 1984 M02
    |                       |--*N. miocaenus (Milne-Edwards 1867) (see below for synonymy) M02
    |                       `--N. anatolicus M02
    `--Sulidae [Dysporidae, Enkurosulidae, Pseudosulidae, Sulariidae, Sulinae] JT12
         |  i. s.: Enkurosula Kašin 1977 [=Pseudosula Harrison 1975 non Boetticher 1955] M02
         |           `--*E. pygmaea (Milne-Edwards 1874) [=Sula pygmaea, Microsula pygmaea, *Pseudosula pygmaea] M02
         |         Eostega Lambrecht 1929 M02
         |           `--*E. lebedinskyi Lambrecht 1929 M02, M09
         |         Empheresula Harrison 1975 [=Parasula Harrison 1975 non Mathews 1913] M02
         |           `--*E. arvernensis (Milne-Edwards 1867) [=Sula arvernensis, *Parasula arvernensis] M02
         |         Sarmatosula Grigorescu & Kessler 1977 M02
         |           `--*S. dobrogensis Grigorescu & Kessler 1977 M02
         |         Prophalacrocorax Harrison 1975 M02
         |           `--*P. ronzoni (Gervais 1849) M02, M09 [=Mergus ronzoni M02, Sula ronzoni M02]
         |--Masillastega Mayr 2002 M05
         |    `--M. rectirostris Mayr 2002 M09
         `--+--+--Papasula abbotti JT12
            |  `--Morus Vieillot 1816 JT12, CC10 [=Moris Forster 1817 CC10, Sulita Mathews 1915 CC10]
            |       |  i. s.: M. olsoni Grigorescu & Kessler 1988 M02
            |       |--*M. bassanus (Linnaeus 1758) CC10, JT12, CS77 (see below for synonymy)
            |       `--+--M. capensis (Lichtenstein 1823) JT12, CC10 [=Dysporus capensis CC10, Sula capensis CC10]
            |          `--M. serrator (Gray in Dieffenbach 1843) JT12, CC10 (see below for synonymy)
            `--Sula Brisson 1760 JT12, CC10 (see below for synonymy)
                 |  i. s.: S. abbotti CS77
                 |         S. cyanops (Sundevall 1837) [=Dysporus cyanops] CC10
                 |         S. fiber Gray 1862 (n. d.) CC10
                 |         S. fusca Vieillot 1825 CC10
                 |         S. guano FP64
                 |         S. piscator (Linnaeus 1758) (n. d.) [=Pelecanus piscator] CC10
                 |--S. sula (Linnaeus 1766) JT12, CC10 [=Pelecanus sula CC10]
                 |    |--S. s. sula I92
                 |    `--S. s. rubripes Gould 1838 I92
                 `--+--*S. leucogaster (Boddaert 1783) CC10, JT12, CC10 [=Pelecanus leucogaster CC10]
                    |    |--S. l. leucogaster CC10
                    |    |--S. l. brewsteri Goss 1888 CC10
                    |    |--S. l. etesiaca Thayer & Bangs 1905 CC10
                    |    `--S. l. plotus (Forster in Lichtenstein 1844) (see below for synonymy) CC10
                    `--+--+--S. dactylatra Lesson 1831 JT12, CC10
                       |  |    |--S. d. dactylatra CC10
                       |  |    |--S. d. bedouti Mathews 1913 [=*Parasula bedouti] CC10
                       |  |    |--S. d. melanops Heuglin 1859 CC10
                       |  |    |--S. d. personata Gould 1846 [=Parasula dactylatra perstonata] CC10
                       |  |    `--S. d. tasmani van Tets, Meredith et al. 1988 (see below for synonymy) CC10
                       |  `--S. granti Rothschild 1902 JT12, CC10
                       `--+--S. nebouxii JT12
                          `--S. variegata JT12

Nomen nudum: Sula rozieri Lucazeau 1959 M02

Anhinga Brisson 1760 CC10 [incl. Plotus Linné 1766 B94, Plottus (l. c.) B94, Ptynx Möhring 1752 (pre-Linnean) non Blyth 1840 B94]

Anhinga novaehollandiae (Gould 1847) JT12, CC10 [=Plotus novaehollandiae CC10, A. melanogaster novaehollandiae CC10, A. rufa novaehollandiae CC10; incl. P. novaehollandiae derbyi Mathews 1912 WS48]

Anhinga rufa (Dausin 1802) JT12, CC10 [=Plotus rufus WS48, A. melanogaster rufa CC10; incl. A. melanogaster chantrei (Oustalet 1882) CS77]

*Morus bassanus (Linnaeus 1758) CC10, JT12, CS77 [=Pelecanus bassanus CC10, Sula bassana CS77, *Sulita bassana CC10]

Morus serrator (Gray in Dieffenbach 1843) JT12, CC10 [=Pelecanus serrator CC10, Dysporus serrator CC10, Moris serrator CC10, Sula serrata CC10, Sula bassana serrator CC10, Sulita serrator CC10; incl. Sula australis Gould 1841 non Stephens 1826 CC10, Jula (l. c.) australis CC10, Sula serrator dyotti Mathews 1913 CC10, Sulita serrator rex Mathews & Iredale 1921 CC10, Morus serrator rex CC10, Sula serrator rex CC10, Pelecanus sectator Solander in Medway 1993 CC10]

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

Sula Brisson 1760 JT12, CC10 [=Dysporus Illiger 1811 CC10; incl. Hemisula Mathews 1913 CC10, Parasula Mathews 1913 CC10]

Sula dactylatra tasmani van Tets, Meredith et al. 1988 [incl. S. dactylatra fullagari O’Brien & Davies 1990] CC10

Sula leucogaster plotus (Forster in Lichtenstein 1844) [=Pelecanus plotus, Hemisula leucgaster plotus; incl. S. leucogaster rogersi Mathews 1913, *H. rogersi] CC10

*Type species of generic name indicated


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[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.

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[PB15] Prum, R. O., J. S. Berv, A. Dornburg, D. J. Field, J. P. Townsend, E. M. Lemmon & A. R. Lemmon. 2015. A comprehensive phylogeny of birds (Aves) using targeted next-generation DNA sequencing. Nature 526: 569–573.

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

[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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Last updated: 1 December 2021.

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