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SULIFORMES Frigatebirds, gannets, darters, and cormorants

The information presented here is identical to that contained in the fifth edition of the Checklist of the Birds of New Zealand (Checklist Committee 2022). To access a pdf version of the Checklist click here.

Symbols and Abbreviations
➤ Indicates a species (cf. subspecies)
† Indicates an extinct taxon
NMNZ, Museum of New Zealand Te Papa Tongarewa, Wellington

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Order SULIFORMES: Frigatebirds, Gannets, Darters, and Cormorants

We follow the recommendations of Ericson et al. (2006), Hackett et al. (2008), Burleigh et al. (2015), Prum et al. (2015), and Kuhl et al. (2021) in separating this clade of waterbirds from Pelecaniformes. We follow Bock (1994), Clements et al. (2019), and Chesser et al. (2020) in using the name Suliformes. The family sequence follows Clements et al. (2019) and Chesser et al. (2020).

Family FREGATIDAE Degland & Gerbe: Frigatebirds

Fregatinae Degland & Gerbe, 1867: Ornithologie européenne 2(11): 357 – Type genus Fregata La Cépède, 1799.

Genus Fregata La Cépède

Fregata La Cépède, 1799: Tableaux Method. Mamm. Oiseaux: 15 – Type species (by subsequent designation) Pelecanus Aquilus Linnaeus = Fregata aquila (Linnaeus).

Tachypetes Vieillot, 1816: Analyse Nouv. Ornith. Elem.: 63 – Type species (by monotypy) Pelecanus minor Gmelin = Fregata minor (Gmelin).

Atagen G.R. Gray, 1841: List Gen. Birds (2nd edition): 101 – Type species (by original designation) Pelecanus minor Gmelin = Fregata minor (Gmelin).

Parvifregata Mathews, 1920: Birds Australia (Suppl. 1) 1: 64 – Type species (by original designation) Atagen ariel G.R. Gray = Fregata ariel (G.R. Gray).

Fregata minor (Gmelin)
Great Frigatebird

Pelecanus minor Gmelin, 1789: Syst. Nat., 13th edition 1(2): 572. Based on the “Lesser Frigate Pelican” of Latham 1785, Gen. Synop. Birds 3(2): 590 – no locality = Christmas Island, Indian Ocean (fide Lowe 1924, Novit. Zool. 31: 306).

Five subspecies commonly accepted, breeding on islands in the following regions (Dorst & Mougin 1979; del Hoyo et al. 1992; Dickinson 2003): F. m. aldabrensis Mathews, 1914 (west Indian Ocean), F. m. minor (Indian Ocean and South China Sea), F. m. palmerstoni (Pacific Ocean), F. m. ridgwayi Mathews, 1914 (east Pacific Ocean), and F. m. nicolli Mathews, 1914 (Atlantic Ocean). However, Marchant & Higgins (1990) recognised no subspecies. The common name great frigatebird is adopted for this species, following the discussion by Marchant & Higgins (1990).

Fregata minor palmerstoni (Gmelin)
Great Frigatebird

Pelecanus Palmerstoni Gmelin, 1789: Syst. Nat., 13th edition 1(2): 573. Based on the “Palmerston Frigate Pelican” of Latham 1785, Gen. Synop. Birds 3(2): 592 – Palmerston Island, Pacific Ocean.

Fregata aquila; Anon. 1870, Cat. Colonial Mus.: 76. Not Pelecanus aquilus Linnaeus, 1758.

Tachypetes aquila; Buller 1888 (Mar.), History of the Birds of N.Z., 2nd edition 2 (part 5): 182. Not Pelecanus aquilus Linnaeus, 1758.

Tachypetes aquilus; Cheeseman 1891, Trans. Proc. N.Z. Inst. 23: 223. Not Pelecanus aquilus Linnaeus, 1758.

Fregata aquila palmerstoni (Gmelin); Mathews & Iredale 1913, Ibis 1 (10th series): 417.

Fregata minor peninsulae Mathews, 1923: Bull. Brit. Ornith. Club 44: 15 – Raine Island, Queensland, Australia.

Fregata minor palmerstoni (Gmelin); Peters 1931, Check-list Birds World 1: 96.

Fregata minor (Gmelin) subspecies; Checklist Committee 1953, Checklist N.Z. Birds: 31.

Fregata minor; Checklist Committee 1970, Annot. Checklist Birds N.Z.: 33. Not Pelecanus minor Gmelin, 1789.

Fregata minor peninsulae Mathews; Checklist Committee 1990, Checklist Birds N.Z.: 86.

About 18 records from the New Zealand region, the most southerly being Westport (Oliver 1930; Checklist Committee 1953; Edgar 1972b; G. Taylor & Parrish 1991; Medway 2000a; Fryer 2004). The largest flock was eight over the Meyer Islands and Raoul Island, Kermadec Islands / Rangitāhua, Mar. 2016 (Miskelly, Crossland et al. 2017). The earliest record was 1861 (not 1855) – see explanation in Miskelly et al. (2022). Former attribution of New Zealand birds to F. m. peninsulae (Checklist Committee 1990) followed Condon (1975), but that name is a junior synonym of F. m. palmerstoni (see Dorst & Mougin 1979).

Fregata ariel (G.R. Gray)
Lesser Frigatebird

Three subspecies generally accepted (Dorst & Mougin 1979; del Hoyo et al. 1992; Dickinson 2003): F. a. ariel (Indian and Pacific Oceans), F. a. iredalei Mathews, 1914 (west Indian Ocean), and F. a. trinitatis Miranda-Ribeiro, 1919 (Atlantic Ocean), although their distinctiveness has been questioned (Marchant & Higgins 1990).

Fregata ariel ariel (G.R. Gray)
Lesser Frigatebird

Atagen ariel G.R. Gray, 1845: Gen. Birds 3: 669, pl. 185 – no locality = Raine Island, northern Queensland, Australia (fide Mathews 1914, Austral Avian Rec. 2: 121).

Fregata minor; Buller 1873 (Mar.), History of the Birds of N.Z., 1st edition (part 5): 342. Not Pelecanus minor Gmelin, 1789.

Tachypetes minor; Buller 1888 (Mar.), History of the Birds of N.Z., 2nd edition 2 (part 5): 185. Not Pelecanus minor Gmelin, 1789.

Fregata ariel ariel (G.R. Gray); Mathews & Iredale 1913, Ibis 1 (10th series): 418.

Fregata ariel (G.R. Gray); Checklist Committee 1990, Checklist Birds N.Z.: 87.

Breeds on islands off Queensland, New Caledonia, Fiji, and elsewhere in the tropical Pacific Ocean. Straggles to the New Zealand region: about 32 records since 1907, mostly in the north but once south to Otago (Turbott 1952; Checklist Committee 1953; Oliver 1955; Hudson 1963; Lockstone 1967; Edgar 1971b, 1972b; Marchant & Higgins 1990; G. Taylor & Parrish 1991; Powlesland et al. 1992; Pierce 1992; Guest 1992; Medway 2000a; Miskelly, Crossland et al. 2013, 2019) and probably twice to the Chatham Islands (Imber 1994). New Zealand records are considered to be of the closest breeding subspecies (Checklist Committee 1953, 1970; Oliver 1955).

Family SULIDAE Reichenbach: Gannets and Boobies

Sulinae Reichenbach, 1849: Avium Syst. Nat.: 6 – Type genus Sula Brisson, 1760.

Morus and Sula are considered generically distinct (Olson 1985b; van Tets et al. 1988; BOU Records Committee 1991; Friesen & Anderson 1997; J. Nelson 2005).

Genus Morus Vieillot

Morus Vieillot, 1816: Analyse Nouv. Ornith. Elem.: 63 – Type species (by monotypy) Buffon’s “Fou de Bassan” = Pelecanus bassanus Linnaeus = Morus bassanus (Linnaeus).

Moris J.R. Forster, 1817: Synop. Cat. Brit. Birds: 59. Unnecessary nomen novum for Morus Vieillot, 1816.

Sulita Mathews, 1915: Austral Avian Rec. 2: 123. Unnecessary nomen novum for Morus Vieillot, 1816.

Morus capensis (Lichtenstein)
Cape Gannet

Dysporus capensis Lichtenstein, 1823: Verzeich. Doubl., Berlin: 86 – Cape of Good Hope, South Africa.

Sula capensis (Lichtenstein); Medway 2001, Notornis 48: 62.

Morus capensis (Lichtenstein); Parrish 2002, Notornis 49: 102.

Breeds on islands off coasts of southern Africa, ranging north in winter and straggling further afield (Dorst & Mougin 1979; C. Robertson & Stephenson 2005). The first New Zealand record was a bird at Cape Kidnappers Dec. 1997–Sep. 2005; it successfully bred with an Australasian gannet and a hybrid chick returned to the colony in 2005 (Medway 2001a; Stephenson 2002, 2003; C. Robertson & Stephenson 2005). Subsequently recorded at Farewell Spit, Dec. 2015 (Miskelly, Crossland et al. 2017).

Morus serrator (G.R. Gray)
Australasian Gannet | Tākapu

Sula Australis Gould, 1841: Proc. Zool. Soc. London 1840 (8): 177 – Tasmanian Seas. Junior primary homonym of Sula australis Stephens, 1826.

Pelecanus serrator G.R. Gray, 1843: in E. Dieffenbach, Travels in N.Z. 2: 200 – no locality – vicinity of the Three Kings Islands (fide Medway 1993, Notornis 40: 69).

Sula serrator (G.R. Gray); G.R. Gray 1845, in Richardson & J.E. Gray (Eds), Zool. Voy. ‘Erebus’ & ‘Terror’, Birds 1(8): 19.

Jula [sic] australis Gould; Ellman 1861, Zoologist 19: 7472. Not Sula australis Stephens, 1826.

Dysporus serrator (G.R. Gray); Finsch 1867, Journ. für Ornith. 15: 339.

Sula serrata; Finsch 1882, Ibis 6 (4th series): 402. Unjustified emendation.

Sula serrator serrator (G.R. Gray); Mathews 1913, Austral Avian Rec. 2: 63.

Sula serrator dyotti Mathews, 1913: Austral Avian Rec. 2: 63 – Tasmania, Australia.

Morus serrator serrator (G.R. Gray); Mathews 1913, List Birds Australia: 98.

Sulita serrator serrator (G.R. Gray); Mathews & Iredale 1921, Man. Birds of Australia 1: 77.

Sulita serrator rex Mathews & Iredale, 1921: Man. Birds of Australia 1: 77 – New Zealand.

Moris serrator (G.R. Gray); Oliver 1930, New Zealand Birds, 1st edition: 207.

Sula bassana serrator (G.R. Gray); Checklist Committee 1953, Checklist N.Z. Birds: 28.

Morus serrator rex; Howard & Moore 1980, Complete Checklist Birds World: 60.

Morus serrator (G.R. Gray); Checklist Committee 1990, Checklist Birds N.Z.: 79.

Pelecanus Sectator Solander, 1993: in Medway, Notornis 40: 66 – “Ocean which washes northern Australia. S.lat. 36–33 W.Long. 185–187”, error for vicinity of the Three Kings Islands (fide Medway 1993, Notornis 40: 69).

Sula serrator rex (Mathews & Iredale); Medway 1993, Notornis 40: 66.

Australia (breeding on islets off Tasmania and south-east Australia), Philip and Nepean Islands (Norfolk Island group) and New Zealand (Marchant & Higgins 1990; J. Nelson 2005). In New Zealand, breeds on outlying islands from Manawatāwhi / Three Kings Islands to Hauraki Gulf, Bay of Plenty, and Tolaga Bay on the east coast of the North Island; Oaia, Motutara (Sugarloaf Rock at Muriwai Beach), and Karewa / Gannet Islands on the North Island west coast (Wodzicki et al. 1984). Cape Kidnappers (south end of Hawke Bay) and two colonies at Muriwai, opposite Motutara Island, are the only North Island mainland colonies (Wodzicki et al. 1984; B. Greene 2003). In the South Island, colonies are at Waimaru Bay (Pelorus Sound), Arapawa Island, Farewell Spit, The Nuggets, and Little Solander Island (Marchant & Higgins 1990; D. Brown & Wilson 2004). Adult birds range widely in New Zealand seas during winter; juveniles and some adults disperse to coastal waters of Australia, as far west as the Indian Ocean, with vagrants to South Africa and north to New Caledonia (Wodzicki & Stein 1958; Wodzicki 1967; Marchant & Higgins 1990; J. Moore 1999; C. Robertson & Stephenson 2005). Stragglers reach the Chatham Islands (including one breeding record; Imber 1994; Miskelly et al. 2006), Snares / Tini Heke, Auckland Islands / Maukahuka, and Campbell Island / Motu Ihupuku (Kinsky 1969; Miskelly et al. 2001a; Miskelly, Elliott et al. 2020). Midden and Holocene deposits: North, South, and Chatham Islands (Millener 1991).

Genus Sula Brisson

Sula Brisson, 1760: Ornithologie 1: 60; 6: 494 – Type species (by tautonymy) Sula leucogaster Boddaert.

Dysporus Illiger, 1811: Prodromus Syst. Mamm. Avium: 279. Unnecessary nomen novum for Sula Brisson, 1760.

Hemisula Mathews, 1913: Austral Avian Rec. 2: 55 – Type species (by original designation) Sula leucogaster rogersi Mathews.

Parasula Mathews, 1913: Austral Avian Rec. 2: 55 – Type species (by original designation) Sula dactylatra bedouti Mathews.

In his list of New Zealand birds, Gray (1862: 250) included “Sula fiber”, with the following localities: “Enderby’s Island; Lord Howe’s Island?”. We are unable to identify this species, so regard it as a nomen dubium.

Sula sula (Linnaeus)
Red-footed Booby

Pelecanus Sula Linnaeus, 1766, Syst. Nat., 12th edition, 1: 218 – “in Pelago indico” = Barbados, West Indies (fide Grant & Mackworth-Praed 1933, Bull. Brit. Ornith. Club 53: 187).

Sula sula (Linnaeus); Verreaux & Des Murs 1860: Rev. et Mag. de Zool.: 442.

Three subspecies are recognised: S. s. sula breeding in the West Indies, Hispaniola, Puerto Rico, Virgin Islands, Dominica, Grenadines, Belize, Venezuela, on Fernando de Noronha, and possibly Ascension Island; S. s. rubripes breeding on many Indian Ocean and Pacific Ocean islands; S. s. websteri breeding on Revillagigedo Islands off Mexico, Cocos Island off Costa Rica, and Galápagos Islands (Dorst & Mougin 1979). Four records from New Zealand (Miskelly, Crossland et al. 2021). The first birds of this species from New Zealand (two at the Kermadec Islands / Rangitāhua in Mar.–Apr. 2016; Miskelly, Crossland et al. 2017) were not identified to subspecies.

F. Hutton (1871: 49) listed “Dysporus piscator L. – red-legged gannet”, but the taxon Pelecanus piscator Linnaeus, 1758 is a nomen dubium. Therefore, Hutton’s citation cannot be assigned to any subspecies of Sula sula.

Sula sula rubripes Gould
Indo-Pacific Red-footed Booby

Sula rubripes Gould, 1838: Synop. Birds Australia 4, App.: 7 – “New South Wales” = Raine Island, northern Queensland (fide Mathews 1915, Birds Australia 4: 210).

Sula nicolli Grant & Mackworth-Praed, 1933: Bull. Brit. Ornith. Club 53: 118 – Gloriosa Island, Indian Ocean.

Sula sula rubripes Gould; del Hoyo et al. 1992: Handb. Birds World 1: 325.

A bird considered to be of this subspecies was found dead on Pakatoa Island, Hauraki Gulf, in May 2017 (Miskelly, Crossland et al. 2021; Auckland Museum specimen LB15822).

Sula sula websteri Rothschild
Eastern Pacific Red-footed Booby

Sula websteri Rothschild, 1898: Bull. Brit. Ornith. Club 54: 52 – “Clarion Island, Galapagos and the neighbouring seas” = Clarion Island, Revillagigedo Islands (fide Dorst & Mougin 1979, in Peters, Check-list Birds World 1 (2nd edition): 186).

Sula piscatrix websteri Rothschild; Beebe 1951, Galápagos: World’s End, 2nd edition: 323, fig. 70.

Sula sula websteri Rothschild; del Hoyo et al. 1992: Handb. Birds World 1: 325.

A bird considered to be of this subspecies was present at the Muriwai gannet colony, west Auckland, for 5 weeks in early 2017 (Miskelly, Crossland et al. 2019).

Sula leucogaster (Boddaert)
Brown Booby

Pelecanus Leucogaster Boddaert, 1783: Tables des Planches Enluminéez d’Histoire Naturelle de M. d’Aubenton: 57, pl. 973 – Cayenne, South America.

Breeds on tropical islands of the Indian, Pacific, and Atlantic Oceans, and the Caribbean Sea. Four subspecies; three outside New Zealand region (Dorst & Mougin 1979, Carboneras 1992b): breeding on Atlantic Ocean and Caribbean islands (S. l. leucogaster) and on eastern Pacific islands (S. l. brewsteri Goss, 1888, and S. l. etesiaca Thayer & Bangs, 1905). All subspecies considered doubtfully distinct by some authors (e.g. Marchant & Higgins 1990).

Sula leucogaster plotus (J.R. Forster)
Brown Booby

Pelecanus Plotus J.R. Forster, 1844: in Lichtenstein Descrip. Animalium: 278 – near New Caledonia.

Sula fusca; Hamilton 1889, Trans. Proc. N.Z. Inst. 21: 128. Not Sula fusca Vieillot, 1825.

Sula sula; Buller 1906, Suppl. Birds N.Z. 2: 50. Not Pelecanus sula Linnaeus, 1766.

Hemisula leucogaster plotus (Forster); Mathews 1913, List Birds Australia: 99.

Sula leucogaster rogersi Mathews, 1913: Austral Avian Rec. 1: 189 – Bedout Island, Western Australia.

Sula leucogaster; Oliver 1930, New Zealand Birds, 1st edition: 205. Not Pelecanus Leucogaster Boddaert, 1783.

Sula leucogaster plotus (Forster); Checklist Committee 1953, Checklist N.Z. Birds: 28.

Breeds on tropical islands in the Pacific and Indian Oceans, also the Red Sea and Gulf of Aden (Dorst & Mougin 1979). Reaches New Zealand waters probably every year (e.g. Powlesland & Powlesland 1993; Veitch et al. 2004; Miskelly, Crossland et al. 2017), rarely south to about 45°S (Pierce 1969). Sometimes these vagrants reside in an area for several months (e.g. Gaze 1975; Guest 1992; G. Taylor & Parrish 1992, 1994a) and may associate with gannets (e.g. Stein 1952; Hawkins et al. 1992; Hawkins & Cook 1994; Miskelly, Crossland et al. 2017). A banded juvenile from Johnston Atoll was collected offshore from Okarito in 1986 (NMNZ OR.023569).

Sula dactylatra Lesson
Masked Booby

Sula dactylatra Lesson, 1831: Traité d’Ornith. 8: 601 – Ascension Island, Atlantic Ocean.

Also known as the blue-faced booby. Four subspecies; all but one (S. d. tasmani) outside the New Zealand region (Dorst & Mougin 1979; Pitman & Jehl 1998): Atlantic Ocean and Caribbean islands (S. d. dactylatra), Indian Ocean and Arabian Sea islands (S. d. melanops Heuglin, 1859), and islands in the eastern Indian Ocean, the Banda and Coral Seas and much of the Pacific Ocean (S. d. personata Gould, 1846). A fifth race, from the eastern Indian Ocean, – S. d. bedouti Mathews, 1913 – is recognised by some authors (e.g. Marchant & Higgins 1990; R. O’Brien & Davies 1990; Dickinson 2003). The Nazca booby (Sula granti Rothschild, 1902), breeding on some eastern Pacific islands, was formerly regarded as a race of S. dactylatra (Pitman & Jehl 1998). Masked boobies straggle south to northern New Zealand, but their subspecific identity or identities have not been determined: Karewa / Gannet Island, 1883; west of North Cape (Otou), 1964; two, Firth of Thames, 1977–78 (Powlesland & Pickard 1992); near Hamilton, Jul. 1983 (J. Moore 1985a); beach-wrecked near Dargaville, May 1988 (Powlesland & Pickard 1992); Paraparaumu Beach, Sep. 1995 (NMNZ OR.025810); Nine Mile Beach, Buller (Scofield 2008). Recorded as Holocene remains at the Chatham Islands (Millener 1991).

Sula dactylatra tasmani van Tets, Meredith, Fullagar & Davidson
Masked Booby

Sula piscator; G.R. Gray 1862, Ibis 4: 250. Not Pelecanus piscator Linnaeus, 1758 = nomen dubium.

Sula cyanops; Cheeseman 1889, Trans. Proc. N.Z. Inst. 21: 121. Not Dysporus cyanops Sundevall, 1837.

Parasula dactylatra personata; Mathews 1913, List Birds Australia: 99. Not Sula personata Gould, 1846.

Sula dactylatra; Oliver 1930, New Zealand Birds, 1st edition: 203. Not Sula dactylatra Lesson, 1831.

Sula dactylatra personata; Checklist Committee 1953, Checklist N.Z. Birds: 28. Not Sula personata Gould, 1846.

Sula tasmani van Tets, Meredith, Fullagar & Davidson, 1988: Notornis 35: 45 – Norfolk Island.

Sula dactylatra fullagari O’Brien & Davies, 1990: Marine Ornithology 18: 2 – Lord Howe Island.

Sula dactylatra tasmani van Tets, Meredith, Fullagar & Davidson; McAllan, Curtis, Hutton & Cooper 2004, Australian Field Ornithology 21(Suppl.): 6.

Local subspecies S. d. tasmani breeds on islands in the Lord Howe, Norfolk Island, and Kermadec Islands / Rangitāhua groups (R. O’Brien & Davies 1990). This subspecies was first described from Holocene material as an extinct species (van Tets et al. 1988) but is now considered to be conspecific with the extant masked booby (Holdaway & Anderson 2001; Holdaway et al. 2001; Worthy & Holdaway 2002; McAllan et al. 2004; Priddel et al. 2005). Ranges mainly north to the east coast of Australia, New Caledonia, Vanuatu (Marchant & Higgins 1990; Priddel et al. 2005; Anon. 2006a), Fiji (Anon. 2004a), and Tonga (J. Moore 1999). Sometimes called the Tasman booby (e.g. van Tets et al. 1988).

Family ANHINGIDAE Lesson: Darters

Anhingas Lesson, 1831: Traité d’Ornith. 8: 598 – Type genus Anhinga Brisson, 1760.

Genus Anhinga Brisson

Anhinga Brisson, 1760: Ornithologie 1: 60; 6: 476 – Type species (by tautonymy and monotypy) Plotus anhinga Linnaeus = Anhinga anhinga (Linnaeus).

Plotus Gunnerus, 1761: Trondh. Selskabs Skrifter 1: 263 – Type species (by subsequent designation) Plotus anhinga Linnaeus = Anhinga anhinga (Linnaeus). Suppressed and invalid (fide ICZN 1973, Opinion 999. Bull. Zool. Nomenclature 30(2): 80).

Anhinga melanogaster Pennant
Darter

Anhinga melanogaster Pennant, 1769: Indian Zool.: 13, pl. 12 – Sri Lanka, and Java, Indonesia.

Three or four subspecies usually recognised (Marchant & Higgins 1990; del Hoyo et al. 1992; Johnsgard 1993; Christidis & Boles 1994; Dickinson 2003): A. m. melanogaster (India to south-east Asia, Sumatra, Java, Borneo, Philippines, and Sulawesi), A. m. rufa (Daudin, 1802) (Africa and Middle East), A. m. novaehollandiae (New Guinea and Australia), and sometimes A. m. vulsini Bangs, 1918 (Madagascar).

Anhinga melanogaster novaehollandiae (Gould)
Australian Darter

Plotus Novae-Hollandiae Gould, 1847: Proc. Zool. Soc. London 1847 (15): 34 – “Southern coast of Australia” = New South Wales (fide Mathews 1913, List Birds Australia: 97).

Plotus novae-hollandiae Gould; Hutton 1904, Index Faunae N.Z.: 39.

Plotus novaehollandiae novaehollandiae Gould; Mathews & Iredale 1913, Ibis 1 (10th series): 416.

Anhinga novae-hollandiae (Gould); Stidolph 1927, Emu 26: 215.

Anhinga novaehollandiae (Gould); Oliver 1930, New Zealand Birds, 1st edition: 201.

Anhinga rufa novaehollandiae (Gould); Checklist Committee 1953, Checklist N.Z. Birds: 31.

Anhinga melanogaster novaehollandiae (Gould); Cramp et al. 1977, Birds Western Palearctic 1: 223.

Anhinga melanogaster rufa; Checklist Committee 1990, Checklist Birds N.Z.: 133. Not Plotus rufus Daudin, 1802.

Australia and New Guinea. Four or five stragglers recorded in New Zealand: possibly seen at Lake Ohau in “summer” 1862; one dead Hokitika, Jan. 1874 (Buller 1875a; Oliver 1955; van Tets & Scarlett 1972). Also live birds: Waiharara, Northland, Dec. 1992 (Medway 2000a); Lake Daniell, Nelson, Nov. 2003 (Scofield 2005a); and Wellington Harbour, Dec. 2003 (Medway 2004b). This race is sometimes considered to be a full species (Marchant & Higgins 1990; del Hoyo et al. 1992).

Family PHALACROCORACIDAE Reichenbach: Cormorants and Shags | Kawau

Phalacrocoracidae Reichenbach, 1849: Avium Syst. Nat.: 6 – Type genus Phalacrocorax Brisson, 1760.

We follow the classification system proposed by Kennedy & Spencer (2014). This includes placement of little shag in the genus Microcarbo, and spotted shag and Pitt Island shag in the genus Phalacrocorax.

Genus Microcarbo Bonaparte

Microcarbo Bonaparte, 1855: Consp. Av. 2: 177. Nomen nudum.

Microcarbo Bonaparte, 1856: Compt. Rend. Séa. Acad. Sci., Paris 43: 577 – Type species (by original designation) Pelecanus pygmaeus Pallas = Microcarbo pygmaeus (Pallas).

Haliëtor Heine, 1860: Journ. für Ornith. 8: 202 – Type species (by original designation) Pelecanus pygmaeus Pallas = Microcarbo pygmaeus (Pallas).

Microcarbo melanoleucos (Vieillot)
Little Pied Cormorant

Three subspecies: M. m. melanoleucos in south-east Asia, New Guinea, Australia, and some western Pacific Islands, M. m. brevicauda (Mayr, 1931) on Rennell Island, Solomon Islands, and M. m. brevirostris in New Zealand (Dorst & Mougin 1979; Marchant & Higgins 1990). Some authors consider M. m. brevirostris to be the same taxon as M. m. melanoleucos (e.g. Dowding & Taylor 1987; M. Taylor 1987) but others regard it as a distinct species (see Marchant & Higgins 1990; del Hoyo et al. 1992). One record from Chatham Island in 1988–89 of uncertain subspecies (Miskelly et al. 2006).

Microcarbo melanoleucos melanoleucos (Vieillot)
Little Pied Cormorant

Hydrocorax melanoleucos Vieillot, 1817: Nouv. Dict. Hist. Nat., nouv. éd. 8: 88 – “Australasie”, restricted to New South Wales, Australia (fide Mathews 1912, Novit. Zool. 18(3): 241).

Phalacrocorax flavirhynchus Gould, 1838: Proc. Zool. Soc. London 1837 (5): 157 – New South Wales, Australia.

Graucalus flavirostris G.R. Gray, 1843: in E. Dieffenbach, Travels in N.Z. 2: 201. Unjustified emendation.

Graculus melanoleucos (Vieillot); G.R. Gray 1862, Ibis 4: 251.

Phalacrocorax melanoleucos (Vieillot); Buller 1888 (Mar.), History of the Birds of N.Z., 2nd edition 2 (part 5): 173.

Phalacrocorax melanoleucos melanoleucos (Vieillot); Marchant & Higgins 1990, HANZAB 1: 902.

Microcarbo melanoleucos (Vieillot); Christidis & Boles 2008, Syst. Taxon. Australian Birds: 20, 103.

Australia, south-east Asia, New Guinea, and some western Pacific Islands (Marchant & Higgins 1990). Bred on Campbell Island / Motu Ihupuku 1967–69 (Kinsky 1969; Marchant & Higgins 1990; del Hoyo et al. 1992). At least seven pied birds on Snares Islands / Tini Heke in 1976–77 were considered to have originated in Australia (Sagar 1977). Several other records from Snares Islands / Tini Heke were also pied birds: 1975 and sightings 1982–85 (Horning 1976; Miskelly et al. 2001a; NMNZ OR.018703). Vagrant little shags on Norfolk Island in 1978, 1979, and 1996–97 were apparently pied (J. Moore 1981, 1999; Schodde et al. 1983), so most likely from Australia. Occasional immigration to mainland New Zealand from Australia has been suggested by several authors (Buller 1887–88; Dowding & Taylor 1987; Marchant & Higgins 1990; Secker 1994).

Microcarbo melanoleucos brevirostris (Gould)
Little Shag | Kawaupaka

Phalacrocorax brevirostris Gould, 1837: Proc. Zool. Soc. London 1837 (5): 26 – no locality = New Zealand (fide Mathews & Iredale 1913, Ibis 1 (10th series): 415).

Gracalus brevirostris (Gould); G.R. Gray 1845, in Richardson & J.E. Gray (Eds), Zool. Voy. ‘Erebus’ & ‘Terror’, Birds 1(8): 20.

Gracalus melanoleucos; G.R. Gray 1845, in Richardson & J.E. Gray (Eds), Zool. Voy. ‘Erebus’ & ‘Terror’, Birds 1(8): 20. Not Hydrocorax melanoleucos Vieillot, 1817.

Carbo flavagula Peale, 1848: U.S. Expl. Exped. 8: 270, 336 – Bay of Islands, Northland.

Haliaeus brevirostris (Gould); Bonaparte 1855, Consp. Gen. Avium 2: 178.

Halieus brevirostris (Gould); Bonaparte 1856, Compt. Rend. Séa. Acad. Sci., Paris 43: 577.

Carbo brevirostris (Gould); Cassin 1858, U.S. Expl. Exped. Ornithology: 375.

Pelecanus carboides; Ellman 1861, Zoologist 19: 7472. Not Phalacrocorax carboides Gould, 1838.

Graculus brevirostris (Gould); G.R. Gray 1862, Ibis 4: 252.

Phalacrocorax finschii Sharpe, 1875: in Richardson & J.E. Gray (Eds), Zool. Voy. ‘Erebus’ & ‘Terror’, Birds – 1 (Appendix): 34 – New Zealand.

Graculus finschi (Sharpe); Finsch 1876, Trans. Proc. N.Z. Inst. 8: 203. Unjustified emendation.

Microcarbo melanoleucos brevirostris (Gould); Mathews & Iredale 1913, Ibis 1 (10th series): 415.

Phalacrocorax melanoleucus; Oliver 1930, New Zealand Birds, 1st edition: 184. Not Hydrocorax melanoleucos Vieillot, 1817. Unjustified emendation.

Haliëtor melanoleucos brevirostris (Gould); Peters 1931, Check-list Birds World 1: 93.

Microcarbo brevirostris (Gould); Mathews 1936, Suppl. Birds Norfolk & Lord Howe Islands: 143.

Phalacrocorax melanoleucos brevirostris Gould; Checklist Committee 1953, Checklist N.Z. Birds: 29.

Breeds from North Cape (Otou) to Stewart Island / Rakiura and on nearby offshore islands. Vagrant to Snares Islands / Tini Heke in the 1960s, where it may have bred (Warham 1967; Warham & Keeley 1969). Apparently this subspecies also reached the Auckland Islands / Maukahuka in 1942, and Campbell Island / Motu Ihupuku in 1958 (Bailey & Sorensen 1962; Miskelly, Elliott et al. 2020). A dimorphic subspecies with some intermediate variants; the relative scarcity of the latter suggests that the white-breasted phase may be maintained by occasional immigration of Ph. m. melanoleucos from Australia (Dowding & Taylor 1987). In Holocene deposits and middens in North, South, and Stewart Islands (e.g. Horn 1983; Worthy 1998c, 1998d).

Genus Phalacrocorax Brisson

Phalacrocorax Brisson, 1760: Ornithologie 1: 60; 6: 511 – Type species (by tautonymy) Phalacrocorax Brisson = Pelecanus carbo Linnaeus = Phalacrocorax carbo (Linnaeus).

Carbo La Cépède, 1799: Tableaux Method. Mamm. Oiseaux: 15 – Type species (by tautonymy) Pelecanus carbo Linnaeus.

Halieus Illiger, 1811: Prodromus Syst. Mamm. Avium: 279. Unnecessary nomen novum for Phalacrocorax Brisson, 1760.

Hydrocorax Vieillot, 1816: Analyse Nouv. Ornith. Elem.: 63 – Type species (by subsequent designation) Pelecanus carbo Linnaeus. Junior homonym of Hydrocorax Brisson, 1760.

Haliaeus Schinz, 1825: in Cuvier’s Thierreich 4: 570. Unjustified emendation.

Graucalus G.R. Gray, 1841: List Gen. Birds (2nd edition): 101 – Type species (by original designation) Pelecanus carbo Linnaeus, 1758. Junior homonym of Graucalus Cuvier, 1816.

Gracalus G.R. Gray, 1845: in Richardson & J.E. Gray (Eds), Zool. Voy. ‘Erebus’ & ‘Terror’, Birds 1(8): 19. Unjustified emendation.

Ecmeles Gistel, 1848: Naturg. Thierreichs: 9. Unnecessary nomen novum for Hydrocorax Vieillot, 1816.

Graculus Reichenbach, 1850: Novit. Synop. Avium: no 2304, pl. 278 – Type species (by original designation) Carbo javanicus Horsfield = Phalacrocorax niger (Vieillot).

Hypoleucus Reichenbach, 1853: Avium Syst. Nat. 2(1): 7 – Type species (by original designation) Pelecanus varius Gmelin = Phalacrocorax varius (Gmelin).

Stictocarbo Bonaparte, 1855: Compt. Rend. Séa. Acad. Sci., Paris 41: 1115 – Type species (by subsequent designation) Pelecanus punctatus Sparrman = Phalacrocorax punctatus (Sparrman).

Enygrotheres Heine & Reichenow, 1890: Nom. Mus. Hein. Ornith.: 353 – Type species (by original designation) Pelecanus punctatus Sparrman = Phalacrocorax punctatus (Sparrman).

Phalacrocorax carbo (Linnaeus)
Great Cormorant

Pelecanus carbo Linnaeus, 1758: Syst. Nat., 10th edition 1: 133 – Europe, restricted to the “rock nesting form of the north Atlantic Ocean” (fide Hartert 1920, Vögel Pal. Fauna: 1387).

Widespread. Six subspecies widely accepted: Europe and North America, Ph. c. carbo; Europe and Asia, Ph. c. sinensis (Blumenbach, 1798); Japan, Ph. c. hanedae Kuroda, 1925; north-west Africa, Ph. c. maroccanus Hartert, 1906; Africa, Ph. c. lucidus (Lichtenstein, 1823); and Australasia, Ph. c. novaehollandiae (see Dorst & Mougin 1979; del Hoyo et al. 1992; Dickinson 2003). An additional race from north-east Africa (Ph. c. lugubris Rüppell, 1845) is sometimes accepted (Dickinson 2003). The form in Australasia (Ph. c. novaehollandiae) is sometimes considered to be a full species (e.g. Buller 1905–06; Siegel-Causey 1988; del Hoyo et al. 1992) with one subspecies in Australia, Ph. n. novaehollandiae – not Ph. n. carboides Gould, 1838 contra Marchant & Higgins (1990: 810) and del Hoyo et al. (1992) – and another in New Zealand, Ph. n. steadi (Mathews & Iredale, 1913) (Dorst & Mougin 1979). However, Pelecanus major Ellman, 1861 is a senior synonym for the New Zealand form. The African race Ph. c. lucidus is also sometimes considered to be a full species (del Hoyo et al. 1992). The six widely used races are accepted here, pending a full taxonomic review of the group.

Phalacrocorax carbo novaehollandiae Stephens
Black Shag | Māpunga

Phalacrocorax Novae Hollandiae Stephens, 1826: in Shaw, General Zool. 13(1): 93 – New South Wales, Australia.

Phalacrocorax carboides Gould, 1838: Proc. Zool. Soc. London 1837 (5): 156 – Tasmania, Australia.

Graucalus carboides (Gould); G.R. Gray 1843, in E. Dieffenbach, Travels in N.Z. 2: 201.

Gracalus carboides (Gould); G.R. Gray 1845, in Richardson & J.E. Gray (Eds), Zool. Voy. ‘Erebus’ & ‘Terror’, Birds 1(8): 20.

Pelecanus flavirostris? Forster; Ellman 1861, Zoologist 19: 7472. Not Phalacrocorax flavirostris G.R. Gray, 1843.

Pelecanus major Ellman, 1861: Zoologist 19: 7472 – New Zealand.

Graculus carboïdes (Gould); G.R. Gray 1862, Ibis 4: 251.

Graculus carbo; Finsch 1870, Journ. für Ornith. 18: 375. Not Pelecanus carbo Linnaeus, 1758.

Gaculus [sic] novaehollandiae (Stephens); G.R. Gray 1871, Hand-list Birds 3: 127.

Phalacrocorax (Graculus) carbo; Potts 1872, Zoologist Jun. 1872: 3092.

Phalacrocorax carbo; Sharpe 1875, in Richardson & J.E. Gray (Eds), Zool. Voy. ‘Erebus’ & ‘Terror’, Birds –1 (Appendix): 34. Not Pelecanus carbo Linnaeus, 1758.

Phalacrocorax novae-zealandiae var. major Forbes, 1892: Trans. Proc. N.Z. Inst. 24: 189 – no locality = Te Aute Swamp, Hawke’s Bay (fide Worthy 2000, Journ. Royal Soc. N.Z. 30: 10). Junior secondary homonym of Pelecanus major Ellman, 1861.

Phalacrocorax major (Ellman); Rothschild 1905, Proc. IVth International Ornith. Congress: 195.

Carbo carbo steadi Mathews & Iredale, 1913: Ibis 1 (10th series): 411 – New Zealand.

Phalacrocorax carbo steadi (Mathews & Iredale); Oliver 1930, New Zealand Birds, 1st edition: 176.

Phalacrocorax carbosteadi; Stead 1932, Life Histories New Zealand Birds: 1. Unjustified emendation.

Phalacrocorax carbo novaehollandiae Stephens; Checklist Committee 1953, Checklist N.Z. Birds: 29.

Breeds throughout the main islands of New Zealand and on the Chatham Islands; straggling to Norfolk, Kermadec, Snares / Tini Heke, Auckland / Maukahuka, Campbell / Motu Ihupuku, and Macquarie Islands; also resident in Australia, Solomon Islands, and New Caledonia (Marchant & Higgins 1990; Barré & Géraux 2004; Veitch et al. 2004; Miskelly, Elliott et al. 2020). Common in Holocene deposits and middens, North and South Islands; a few Holocene records from Stewart Island / Rakiura and Chatham Islands (Millener 1991; Worthy 1998c).

Phalacrocorax varius (Gmelin)
Pied Cormorant

Two subspecies: Ph. v. varius in New Zealand and Ph. v. hypoleucos (Brandt, 1837) in Australia (Marchant & Higgins 1990).

Phalacrocorax varius varius (Gmelin)
Pied Shag | Kāruhiruhi

Pelecanus varius Gmelin, 1789: Syst. Nat., 13th edition 1(2): 576. Based on the “Pied Shag” of Latham 1785, Gen. Synop. Birds 3(2): 605 – Queen Charlotte Sound, Marlborough.

Graucalus varius (Gmelin); G.R. Gray 1843, in E. Dieffenbach, Travels in N.Z. 2: 201.

Pelecanus pica J.R. Forster, 1844: in M.H.C. Lichtenstein, Descrip. Animalium: 104 – Queen Charlotte Sound, Marlborough.

Gracalus varius (Gmelin); G.R. Gray 1845, in Richardson & J.E. Gray (Eds), Zool. Voy. ‘Erebus’ & ‘Terror’, Birds 1(8): 19.

Carbo fucosus Peale, 1848: U.S. Expl. Exped. 8: 268, 336 – Bay of Islands, Northland.

Hypoleucus varius (Gmelin); Reichenbach 1853, Avium Syst. Nat. 2(1): 7.

Carbo hypoleucus; Cassin 1858, U.S. Expl. Exped. Ornithology: 372. Not Carbo hypoleucos Brandt, 1837.

Carbo leucogaster; Cassin 1858, U.S. Expl. Exped. Ornithology: 372. Not Hydrocorax leucogaster Vieillot, 1817.

Graculus varius (Gmelin); G.R. Gray 1862, Ibis 4: 251.

Phalacrocorax varius (Gmelin); Buller 1873 (Mar.), History of the Birds of N.Z., 1st edition (part 5): 328.

Graculus leucogaster; Finsch 1882, Ibis 6 (4th series): 402. Not Hydrocorax leucogaster Vieillot, 1817.

Hypoleucus varius varius (Gmelin); Mathews & Iredale 1913, Ibis 1 (10th series): 411.

Phalacrocorax varius varius (Gmelin); Oliver 1930, New Zealand Birds, 1st edition: 181.

Breeds on coasts, harbours, estuaries, and offshore islands, from Manawatāwhi / Three Kings Group to Stewart Island / Rakiura (Marchant & Higgins 1990; C. Robertson et al. 2007). Straggles to Snares Islands / Tini Heke (Miskelly et al. 2001a). Midden and Late Pleistocene or Holocene deposits, North, South, and Stewart Islands (Millener 1991; Worthy 1998c; Worthy & Grant-Mackie 2003).

Phalacrocorax sulcirostris (Brandt)
Little Black Shag | Kawau Tūī

Carbo sulcirostris Brandt, 1837: Bull. l’Acad. Imp. Sci., St Petersburg 3: 56 – “Terrae australes”, restricted to New South Wales, Australia (fide Mathews & Iredale 1913, Ibis 1 (10th series): 415).

Carbo purpuragula Peale, 1848: U.S. Expl. Exped. 8: 269, 336 – Manua Bay, Raglan, Waikato.

Microcarbo stictocephalus Bonaparte, 1857: Consp. Gen. Avium 2: 178 – New South Wales, Australia.

Graculus stictocephalus (Bonaparte); G.R. Gray 1862, Ibis 4: 252.

Graculus sulcirostris Brandt [sic]; Anon. 1870, Cat. Colonial Mus.: 76.

Graculus sulcirostris (Brandt); Finsch 1872, Journ. für Ornith. 20: 258.

Graculus? sulcirostris (Brandt); Finsch 1874, Journ. für Ornith. 22: 174, 214.

Phalacrocorax purpuragula (Peale); Sharpe 1875, in Richardson & J.E. Gray (Eds), Zool. Voy. ‘Erebus’ & ‘Terror’, Birds – 1 (Appendix): 34.

Phalacrocorax sulcirostris (Brandt); Hutton 1904, Index Faunae N.Z.: 33.

Mesocarbo ater purpuragula (Peale); Mathews & Iredale 1913, Ibis 1 (10th series): 415.

Mesocarbo sulcirostris purpuragula (Peale); Mathews 1929, Ibis 5 (12th series): 699.

Breeds in Indonesia, New Guinea, Australia, and New Zealand (Marchant & Higgins 1990). In New Zealand, widely distributed on lakes and estuaries of the North Island; numbers increasing (e.g. H. Robertson 1992) and now regularly seen in the South Island (e.g. Edgar 1972b; Bull et al. 1985; W. Cooper 1991; O’Donnell & West 1994, 1995; O’Donnell 2001; Pollock 2003); rarely south to Stewart Island / Rakiura (Bull et al. 1985). Breeds at scattered sites from Northland to Wairarapa, especially in Waikato and Rotorua districts (Oliver 1955; Marchant & Higgins 1990; H. Robertson 1992; Powlesland et al. 1993); disperses in winter, especially to the coast (Powlesland et al. 1993). Vagrant at Norfolk Island, Snares Islands / Tini Heke, and Auckland Islands / Maukahuka (Marchant & Higgins 1990; J. Moore 1999; Miskelly et al. 2015; Miskelly, Elliott et al. 2020) and probably at the Kermadec Islands / Rangitāhua (Morrison 1979; Veitch et al. 2004). The species probably arrived in New Zealand after human colonisation; no Holocene or midden records (Holdaway et al. 2001).

Phalacrocorax punctatus (Sparrman)
Spotted Shag | Kawau Tikitiki

Pelicanus [sic] punctatus Sparrman, 1786: Mus. Carlsonianum 1: no X, pl. 10 – Queen Charlotte Sound, Marlborough.

Pelecanus naevius Gmelin, 1789: Syst. Nat., 13th edition 1(2): 575 – Queen Charlotte Sound, Marlborough.

Phalacrocorax naevius (Gmelin); Cuvier 1817, Règne Anim. 1: 525.

Hydrocorax dilophus Vieillot, 1817: Nouv. Dict. Hist. Nat., nouv. éd. 8: 85. Unnecessary nomen novum for Pelecanus naevius Gmelin, 1789.

Graucalus punctatus (Sparrman); G.R. Gray 1843, in E. Dieffenbach, Travels in N.Z. 2: 201.

Gracalus punctatus (Forster) [sic]; G.R. Gray 1845, in Richardson & J.E. Gray (Eds), Zool. Voy. ‘Erebus’ & ‘Terror’, Birds 1(8): 20.

Graculus naevius (Gmelin); Reichenbach 1850, Vollst. Naturg. Schwim. Aves Natatores: pl. 33, fig. 369.

Stictocarbo punctatus (Sparrman); Bonaparte 1856, Compt. Rend. Séa. Acad. Sci., Paris 43: 574.

Pelecanus cirrhatus Gray; Ellman 1861, Zoologist: 19: 7472. Not Pelecanus cirrhatus Gmelin, 1789.

Graculus punctatus (Sparrman); G.R. Gray 1862, Ibis 4: 252.

Phalacrocorax punctatus (Sparrman); Hutton 1904, Index Faunae N.Z.: 33.

Phalacrocorax punctatus; Ogilvie-Grant 1905, Ibis 5 (8th series): 567. Not Pelecanus punctatus Sparrman, 1786.

Stictocarbo punctatus sassi Mathews, 1930. Bull. Brit. Ornith. Club 50: 19 – North Island.

Stictocarbo steadi Oliver, 1930: Trans. N.Z. Inst. 61: 139 – Otago. Junior secondary homonym of Carbo steadi Mathews & Iredale, 1913.

Stictocarbo [punctatus] steadi Oliver; Mathews 1931, Ibis 1 (13th series): 45.

Phalacrocorax oliveri Mathews, 1931: Bull. Brit. Ornith. Club 51: 18. Nomen novum for Stictocarbo steadi Oliver, 1930.

Phalacrocorax punctatus oliveri Mathews, 1930 [sic]; Peters 1931, Check-list Birds World 1: 92.

Phalacrocorax (Stictocarbo) punctatus (Sparrman); Checklist Committee 1953, Checklist N.Z. Birds: 30.

Phalacrocorax (Stictocarbo) punctatus steadi; Checklist Committee 1953, Checklist N.Z. Birds: 31. Not Carbo steadi Mathews & Iredale, 1913.

Stictocarbo punctatus (Sparrman); Checklist Committee 1970, Annot. Checklist Birds N.Z.: 33.

Stictocarbo punctatus steadi; Checklist Committee 1970, Annot. Checklist Birds N.Z.: 33. Not Carbo steadi Mathews & Iredale, 1913.

Leucocarbo punctatus (Sparrman); van Tets 1976, Proceedings 16th International Orn. Congress: 122.

Phalacrocorax punctatus (Sparrman); Marchant & Higgins 1990, HANZAB 1: 838.

Leucocarbo punctatus oliveri (Mathews); Johnsgard 1993, Cormorants, Darters and Pelicans: 315.

Stictocarbo punctatus oliveri (Mathews); Tennyson & Bartle 2008, Tuhinga 19: 195.

In the North Island, breeds on islands at the east end of Waiheke Island, and on Wellington islands (Kapiti, Makaro / Ward, Matiu / Somes, and Mokopuna) (Tennyson 1991b; H. Robertson 1992; Miskelly 2000; Cotter & Nicholson 2005; Rawlence, Rayner et al. 2019). In the South Island, breeds in the Marlborough Sounds, on Banks Peninsula, in Otago, along the South Island West Coast, (including The Steeples, Te Miko, Perpendicular Point, Motukiekie Rocks, Point Elizabeth, and Open Bay Islands), and on Stewart Island / Rakiura and its inshore islands; on Codfish / Whenua Hou, Omaui, Pig, and Centre Islands (Stead 1948; Oliver 1955; Kinsky 1970b; Edgar 1972b; W. Cooper 1991; W. Cooper & McClelland 1992; O’Donnell & West 1992; W. Cooper 1994; C. Robertson et al. 2007). Near Auckland, formerly bred on other islands of the inner Hauraki Gulf (The Noises, Motukawao group), Auckland west coast (Te Henga (Bethells Beach), Oaia Island), Waikato coast (Girdwood Point near Waikaretu) (Turbott 1956b; Checklist Committee 1970; A. Rowe et al. 2000; Rawlence, Rayner et al. 2019). Vagrant inland and to Snares Islands / Tini Heke and Macquarie Island (Buller 1887–88: 154; Ogilvie-Grant 1905; Oliver 1955; G. Taylor & Parrish 1991; Miskelly et al. 2001a). Common in midden and Late Pleistocene or Holocene deposits of North, South, and Stewart Islands (Millener 1991; Worthy 1998c; Worthy & Grant-Mackie 2003).

We follow Rawlence, Rayner et al. (2019) in not recognising any subspecies.

Phalacrocorax featherstoni Buller
Pitt Island Shag | Kawau o Rangihaute

Graculus africanus; Hutton 1872, Ibis 2 (3rd series): 249. Not Pelecanus africanus Gmelin, 1788.

Phalacrocorax africanus; Hutton 1873, Trans. Proc. N.Z. Inst. 5: 224. Not Pelecanus africanus Gmelin, 1788.

Phalacrocorax featherstoni Buller, 1873: Ibis 3 (3rd series): 90 – Chatham Islands.

Graculus Featherstoni (Buller); Finsch 1874, Journ. für Ornith. 22: 174, 215.

Stictocarbo featherstoni (Buller); Mathews & Iredale 1913, Ibis 1 (10th series): 414.

Phalacrocorax (Stictocarbo) punctatus featherstoni Buller; Checklist Committee 1953, Checklist N.Z. Birds: 31.

Stictocarbo punctatus featherstoni (Buller); Checklist Committee 1970, Annot. Checklist Birds N.Z.: 33.

Leucocarbo featherstoni (Buller); van Tets 1976, Proc. 16th International Ornith. Congress: 122.

Phalacrocorax punctatus featherstoni Buller; Dorst & Mougin 1979, in Peters, Check-list Birds World 1 (2nd edition): 172.

Stictocarbo featherstoni (Buller); Siegel-Causey 1988, Condor 90: 892.

Leucocarbo (punctatus) featherstoni (Buller); Johnsgard 1993, Cormorants, Darters and Pelicans: 321.

Chatham Islands (M. Bell & Bell 2000b; Bester & Charteris 2005). Recorded from Holocene dune and midden deposits (Millener 1991). Rangihaute is a proper noun and should have an initial capital.

Genus Leucocarbo Bonaparte

Leucocarbo Bonaparte, 1857: Consp. Gen. Avium 2: 176 – Type species (by subsequent designation) Carbo bougainvillii Lesson = Leucocarbo bougainvillii (Lesson).

Euleucocarbo Voisin, 1973: Notornis 20: 268 – Type species (by original designation) Leucocarbo (Euleucocarbo) carunculatus (Gmelin). As a subgenus of Leucocarbo.

Nesocarbo Voisin, 1973: Notornis 20: 268 – Type species (by original designation) Leucocarbo (Nesocarbo) campbelli (Filhol). As a subgenus of Leucocarbo.

Notocarbo Siegel-Causey, 1988: Condor 90: 891 – Type species (by original designation) Notocarbo atriceps atriceps (King) = Leucocarbo atriceps atriceps (King).

The species sequence is based on phylogenetic trees in Rawlence, Till et al. (2017) and Rawlence et al. (2022).

Leucocarbo ranfurlyi (Ogilvie-Grant)
Bounty Island Shag

Phalacrocorax ranfurlyi Ogilvie-Grant, 1901: Bull. Brit. Ornith. Club 11: 66 – Bounty Islands.

Hypoleucus campbelli ranfurlyi (Ogilvie-Grant); Mathews & Iredale 1913, Ibis 1 (10th series): 413.

Phalacrocorax carunculatus ranfurlyi Ogilvie-Grant; Peters 1931, Check-list Birds World 1: 91.

Phalacrocorax (Leucocarbo) campbelli ranfurlyi Ogilvie-Grant; Checklist Committee 1953, Checklist N.Z. Birds: 30.

Leucocarbo campbelli ranfurlyi (Ogilvie-Grant); Checklist Committee 1970, Annot. Checklist Birds N.Z.: 33.

Leucocarbo ranfurlyi (Ogilvie-Grant); van Tets 1976, Proc. 16th International Ornith. Congress: 122.

Phalacrocorax campbelli ranfurlyi Ogilvie-Grant; Dorst & Mougin 1979, in Peters, Check-list Birds World 1 (2nd edition): 177.

Euleucocarbo ranfurlyi (Ogilvie-Grant); Siegel-Causey 1988, Condor 90: 892.

Leucocarbo (carunculatus) ranfurlyi (Ogilvie-Grant); Johnsgard 1993, Cormorants, Darters and Pelicans: 283.

Breeds only on the Bounty Islands; possible vagrant at Antipodes Islands (Warham & Bell 1979; C. Robertson & van Tets 1982; G. Taylor 2000a).

➤ Leucocarbo septentrionalis Rawlence, Till, Easton, Spencer, Schuckard, Melville, Scofield, Tennyson, Rayner, Waters & Kennedy
Kohatu Shag | Kawau Kōhatu

Leucocarbo carunculatus; Worthy 1996, New Zealand Journ. Zool. 23(1): 95. Not Pelecanus carunculatus Gmelin, 1789.

Leucocarbo septentrionalis Rawlence, Till, Easton, Spencer, Schuckard, Melville, Scofield, Tennyson, Rayner, Waters & Kennedy, 2017: Molec. Phylogen. Evolution 115: 207 – Tokerau Beach, Doubtless Bay, Northland.

This extinct species was described by Rawlence, Till et al. (2017). It is known only from coastal Holocene bone deposits in Northland (Rawlence, Till et al. 2017). The Māori name should have a macron as shown here (and as gifted by Ngāti Kuri) – use of a macron was not permitted in the journal where the species was first named.

Leucocarbo carunculatus (Gmelin)
New Zealand King Shag | Kawau Pāteketeke

Pelecanus carunculatus Gmelin, 1789: Syst. Nat., 13th edition 1(2): 576. Based on the “Carunculated Shag” of Latham 1785, Gen. Synop. Birds 3(2): 603 – Queen Charlotte Sound, Marlborough.

Pelecanus cirrhatus Gmelin, 1789: Syst. Nat., 13th edition 1(2): 576 – Queen Charlotte Sound, Marlborough.

Hydrocorax cirratus (Gmelin); Vieillot 1817, Nouv. Dict. Hist. Nat., nouv. éd. 8: 84. Unjustified emendation.

Pelecanus cirratus Gmelin; Dumont 1818, in Levrault, Dict. Sci. Nat. 10: 453. Unjustified emendation.

Phalacrocorax cirrhatus (Gmelin); Stephens 1826, in Shaw, General Zool. 13(1): 95.

Graucalus carunculatus (Gmelin); G.R. Gray 1843, in E. Dieffenbach, Travels in N.Z. 2: 201.

Graucalus cirrhatus (Gmelin); G.R. Gray 1843, in E. Dieffenbach, Travels in N.Z. 2: 201.

Gracalus cirrhatus (Gmelin); G.R. Gray 1845, in Richardson & J.E. Gray (Eds), Zool. Voy. ‘Erebus’ & ‘Terror’, Birds 1(8): 19.

Graculus cirrhatus (Gmelin); G.R. Gray 1862, Ibis 4: 251.

Phalacrocorax carunculatus (Gmelin); Buller 1873 (Mar.), History of the Birds of N.Z., 1st edition (part 5): 332.

Graculus carunculatus (Gmelin); Finsch 1874, Journ. für Ornith. 22: 174.

Phalacrocorax finschii Buller, 1876: Trans. Proc. N.Z. Inst. 8: 197, 417 – Queen Charlotte Sound, Marlborough. Junior primary homonym of Phalacrocorax finschii Sharpe, 1875 = Microcarbo melanoleucos brevirostris (Gould).

Phalacrocorax cirrhatus (Gmelin); Hutton 1879, Trans. Proc. N.Z. Inst. 6: 336. In part.

Hypoleucus carunculatus carunculatus (Gmelin); Mathews & Iredale 1913, Ibis 1 (10th series): 412.

Phalacrocorax (Leucocarbo) carunculatus carunculatus (Gmelin); Checklist Committee 1953, Checklist N.Z. Birds: 29.

Leucocarbo carunculatus carunculatus (Gmelin); Checklist Committee 1970, Annot. Checklist Birds N.Z.: 32.

Leucocarbo carunculatus (Gmelin); van Tets 1976, Proc. 16th International Ornith. Congress: 122.

Phalacrocorax carunculatus carunculatus (Gmelin); Dorst & Mougin 1979, in Peters, Check-list Birds World 1 (2nd edition): 176.

Euleucocarbo carunculatus (Gmelin); Siegel-Causey 1988, Condor 90: 892.

Breeds only on islands on the western side of Cook Strait (i.e. outer Marlborough Sounds): White Rocks, Sentinel Rock, Duffers Reef, North Trio, Stewart (Tekuru Kuru; Marlborough), and Rahuinui Islands (A. Nelson 1971; Medway 1987; Schuckard 1994, 2006). Vagrant to Adele Island, Tasman Bay, and Farewell Spit (Bull et al. 1985; Miskelly, Crossland et al. 2017). May have reached the North Island: Lake Horowhenua in Jul. 1966 (Edgar 1972b) and Wellington Harbour in Jul. 2002 (Medway 2002f; Parrish 2006). Records of this species at Oamaru (Marchant & Higgins 1990) refer to the Otago shag. Holocene deposits and middens in the northern South Island (e.g. Worthy 1998d). Leucocarbo shag bones from Wairarapa were confirmed to be this species based on DNA analyses (McFadgen 2003; Rawlence, Till et al. 2017).

Leucocarbo onslowi (Forbes)
Chatham Island Shag | Papua

Graculus carunculatus; Hutton 1872, Ibis 2 (3rd series): 249. Not Pelecanus carunculatus Gmelin, 1789.

Phalacrocorax carunculatus; Hutton 1873, Trans. Proc. N.Z. Inst. 5: 224. Not Pelecanus carunculatus Gmelin, 1789.

Graculus carunculatus; Finsch 1874, Journ. für Ornith. 22: 213. Not Pelecanus carunculatus Gmelin, 1789.

Phalacrocorax cirrhatus; Hutton 1879, Trans. Proc. N.Z. Inst. 6: 336. In part.

Phalacrocorax imperialis; Buller 1888 (Mar.), History of the Birds of N.Z., 2nd edition 2 (part 5): 153. Not Phalacrocorax imperialis King, 1831.

Phalacrocorax onslowi Forbes, 1893: Ibis 5 (6th series): 533 – Chatham Islands.

Phalacrocorax rothschildi Forbes, 1893: Ibis 5 (6th series): 537 – Chatham Islands.

Hypoleucus carunculatus onslowi (Forbes); Mathews & Iredale 1913, Ibis 1 (10th series): 412.

Phalacrocorax (Leucocarbo) carunculatus onslowi Forbes; Checklist Committee 1953, Checklist N.Z. Birds: 30.

Leucocarbo carunculatus onslowi (Forbes); Checklist Committee 1970, Annot. Checklist Birds N.Z.: 32.

Leucocarbo onslowi (Forbes); van Tets 1976, Proc. 16th International Ornith. Congress: 122.

Phalacrocorax carunculatus onslowi Forbes; Dorst & Mougin 1979, in Peters, Check-list birds World 1 (2nd edition): 176.

Euleucocarbo onslow (Forbes); Siegel-Causey 1988, Condor 90: 892. Unjustified emendation.

Leucocarbo (carunculatus) onslowi (Forbes); Johnsgard 1993, Cormorants, Darters and Pelicans: 275.

Chatham Islands (M. Bell & Bell 2000b; Bester & Charteris 2005). Recorded from both Holocene deposits and middens (Millener 1991).

Leucocarbo chalconotus (G.R. Gray)
Otago Shag | Matapo

Graucalus auritus; G.R. Gray 1843, in E. Dieffenbach, Travels in N.Z. 2: 201. Not Carbo auritus Lesson, 1831.

Gracalus chalconotus G.R. Gray, 1845: in Richardson & J.E. Gray (Eds), Zool. Voy. ‘Erebus’ & ‘Terror’, Birds 1(8): 20, pl. 21 – Otago.

Graculus glaucus Reichenbach, 1850: Avium Syst. Nat.: pl. 49, fig. 2553. Based on the “Cormoran glauque” of Hombron & Jacquinot, 1845 in Dumont d’Urville, Voyage Pôle Sud, Zoologie, Atlas pl. 31, fig. 1 – Otago.

Phalacrocorax glaucus Hombron & Jacquinot; Pucheran 1853, in Dumont d’Urville, Voyage Pôle Sud, Zoologie, 3: 127.

Graculus chalconotus (G.R. Gray); G.R. Gray 1862, Ibis 4: 252.

Phalacrocorax chalconotus (G.R. Gray); Buller 1873 (Mar.), History of the Birds of N.Z., 1st edition (part 5): 334.

Phalacrocorax glaucus Hombron & Jacquinot; Buller 1888 (Mar.), History of the Birds of N.Z., 2nd edition 2 (part 5): 163.

Phalacrocorax huttoni Buller, 1888 (Mar.): History of the Birds of N.Z., 2nd edition 2 (part 5): 174 – near Dunedin.

Hypoleucus chalconotus (G.R. Gray); Mathews & Iredale 1913, Ibis 1 (10th series): 413.

Hypoleucus huttoni (Buller); Mathews 1936, Suppl. Birds Norfolk & Lord Howe Islands: 141.

Phalacrocorax (Leucocarbo) carunculatus chalconotus (G.R. Gray); Checklist Committee 1953, Checklist N.Z. Birds: 29. In part.

Leucocarbo carunculatus chalconotus (G.R. Gray); Checklist Committee 1970, Annot. Checklist Birds N.Z.: 32. In part.

Leucocarbo chalconotus (G.R. Gray); van Tets 1976, Proc. 16th International Ornith. Congress: 122.

Phalacrocorax carunculatus chalconotus (G.R. Gray); Dorst & Mougin 1979, in Peters, Check-list Birds World 1 (2nd edition): 176.

Euleucocarbo chalconotus (G.R. Gray); Siegel-Causey 1988, Condor 90: 892.

Leucocarbo chalconotus (G.R. Gray); Checklist Committee 1990, Checklist Birds N.Z.: 83. In part.

Phalacrocorax chalconotus (G.R. Gray); Marchant & Higgins 1990, HANZAB 1: 876. In part.

Leucocarbo (carunculatus) chalconotus (G.R. Gray); Johnsgard 1993, Cormorants, Darters and Pelicans: 271.

Inhabits coastal waters and breeds only from near Oamaru south to The Sisters, Catlins coast (Rawlence et al. 2016). Ranges north to Banks Peninsula (Crossland 2021). Midden and Late Pleistocene or Holocene bone deposits, Cape Wanbrow, Otago, and Marfells Beach, Marlborough (Worthy 1998d; Worthy & Grant-Mackie 2003; Rawlence, Till et al. 2017).

The dimorphic Leucocarbo shags of Otago, Southland, and Stewart Island were considered a single species (Stewart Island shag L. chalconotus) until Rawlence et al. (2016) demonstrated that the Otago population was sister to the monomorphic Chatham Island shag (L. onslowi), meaning that either all three populations were conspecific, or that they should all be considered full species. We follow Rawlence et al. (2016) in separating the northern and southern populations of Stewart Island shag (sensu Checklist Committee 2010) as full species. The type specimen for L. chalconotus was from Otago (G.R. Gray 1845), and so this name is retained for the larger northern species, with Otago shag as the recommended English name.

Leucocarbo stewarti (Ogilvie-Grant)
Foveaux Shag | Mapo

Phalacrocorax colensoi Buller, 1888 (Mar.): History of the Birds of N.Z., 2nd edition 2 (part 5): 161 – Bluff, Southland. Not Phalacrocorax colensoi Buller, 1888 – Auckland Islands.

Phalacrocorax stewarti Ogilvie-Grant, 1898: Cat. Birds Brit. Mus. 26: 385 – Bluff, Southland.

Hypoleucus campbelli stewarti (Ogilvie-Grant); Mathews & Iredale 1913, Ibis 1 (10th series): 413.

Phalacrocorax huttoni Buller; Oliver 1930, New Zealand Birds, 1st edition: 191. In part.

Phalacrocorax chalconotus (G.R. Gray); Oliver 1930, New Zealand Birds, 1st edition: 192. In part.

Phalacrocorax (Leucocarbo) carunculatus chalconotus (G.R. Gray); Checklist Committee 1953, Checklist N.Z. Birds: 29. In part.

Leucocarbo carunculatus chalconotus (G.R. Gray); Checklist Committee 1970, Annot. Checklist Birds N.Z.: 32. In part.

Leucocarbo chalconotus (G.R. Gray); Checklist Committee 1990, Checklist Birds N.Z.: 83. In part.

Phalacrocorax chalconotus (G.R. Gray); Marchant & Higgins 1990, HANZAB 1: 876. In part.

Foveaux Strait and Stewart Island / Rakiura; midden and Late Pleistocene or Holocene bone deposits from Stewart Island (Worthy 1998c).

See comments under L. chalconotus above. The type locality for L. stewarti was Bluff (Ogilvie-Grant 1898, p.386), not Stewart Island / Rakiura (fide Checklist Committee 2010; Rawlence et al. 2016). We follow Rawlence et al. (2016) in recommending that this smaller southern species be known as the Foveaux shag.

Leucocarbo colensoi (Buller)
Auckland Island Shag | Kawau o Motu Maha

Phalacrocorax colensoi Buller, 1888 (Mar.): History of the Birds of N.Z., 2nd edition 2 (part 5): 161 – Auckland Islands. In part.

Phalacrocorax campbelli Ogilvie-Grant, 1905: Ibis 5 (8th series): 573 – Auckland Islands. In part.

Hypoleucus campbelli colensoi (Buller); Mathews & Iredale 1913, Ibis 1 (10th series): 413.

Phalacrocorax carunculatus colensoi Buller; Peters 1931, Check-list Birds World 1: 91.

Phalacrocorax (Leucocarbo) campbelli colensoi Buller; Checklist Committee 1953, Checklist N.Z. Birds: 30.

Leucocarbo campbelli colensoi (Buller); Checklist Committee 1970, Annot. Checklist Birds N.Z.: 33.

Leucocarbo colensoi (Buller); van Tets 1976, Proc. 16th International Ornith. Congress: 122.

Phalacrocorax campbelli colensoi Buller; Dorst & Mougin 1979, in Peters, Check-list Birds World 1 (2nd edition): 177.

Euleucocarbo colensoi (Buller); Siegel-Causey 1988, Condor 90: 892.

Leucocarbo (carunculatus) colensoi (Buller); Johnsgard 1993, Cormorants, Darters and Pelicans: 279.

Breeds only on the Auckland Islands / Maukahuka (G. Taylor 2000a; Miskelly, Elliott et al. 2020). One resident at Snares Islands / Tini Heke 1994–2001 (Miskelly et al. 2001a,b). Natural and midden remains have been found in dunes on Enderby Island (Anderson 2005; Tennyson 2020a).
Motu Maha is a proper noun and should have initial capitals.

Leucocarbo campbelli (Filhol)
Campbell Island Shag

Urile campbelli Filhol, 1878: Bull. Soc. Philomath. Paris 7(2): 132 – Campbell Island.

Phalacrocorax magellanicus; Hutton 1879, Trans. Proc. N.Z. Inst. 11: 338. Not Pelecanus magellanicus Gmelin, 1789.

Phalacrocorax nycthemerus; Hutton 1880, Proc. Linn. Soc. New South Wales 4: 357. Not Phalacrocorax nycthemerus Cabanis, 1855 = nomen dubium.

Phalacrocorax campbelli (Filhol); Buller 1906, Suppl. Birds N.Z. 2: 39.

Hypoleucus campbelli campbelli (Filhol); Mathews & Iredale 1913, Ibis 1 (10th series): 412.

Phalacrocorax carunculatus campbelli (Filhol); Peters 1931, Check-list Birds World 1: 91.

Hypoleucus campbelli (Filhol); Mathews 1936, Suppl. Birds Norfolk & Lord Howe Islands: 135.

Phalacrocorax (Leucocarbo) campbelli campbelli (Filhol); Checklist Committee 1953, Checklist N.Z. Birds: 30.

Leucocarbo campbelli campbelli (Filhol); Checklist Committee 1970, Annot. Checklist Birds N.Z.: 33.

Leucocarbo campbelli (Filhol); van Tets 1976, Proc. 16th International Ornith. Congress: 122.

Phalacrocorax campbelli campbelli (Filhol); Dorst & Mougin 1979, in Peters, Check-list Birds World 1 (2nd edition): 177.

Breeds only on Campbell Island / Motu Ihupuku (G. Taylor 2000a). Vagrant at Antipodes Islands (Tennyson et al. 2002).

Leucocarbo purpurascens (Brandt)
Macquarie Island Shag

Carbo purpurascens Brandt, 1837: Bull. l’Acad. Imp. Sci., St Petersburg 3: 56 – no locality = Macquarie Island (fide Rothschild 1898, Bull. Brit. Ornith. Club 8: 21).

Phalacrocorax carunculatus (Gmelin); Buller 1888 (Mar.), History of the Birds of N.Z., 2nd edition 2 (part 5): 155. In part.

Phalacrocorax traversi Rothschild, 1898: Bull. Brit. Ornith. Club 8: 21 – Macquarie Island.

Hypoleucus carunculatus traversi (Rothschild); Mathews & Iredale 1913, Ibis 1 (10th series): 412.

Phalacrocorax atriceps traversi Rothschild; Peters 1931, Check-list Birds World 1: 92.

Hypoleucus atriceps purpurascens (Brandt); Mathews 1936, Suppl. Birds Norfolk & Lord Howe Islands: 134.

Phalacrocorax (Leucocarbo) albiventer purpurascens (Brandt); Checklist Committee 1953, Checklist N.Z. Birds: 30.

Leucocarbo albiventer purpurascens (Brandt); Checklist Committee 1970, Annot. Checklist Birds N.Z.: 33.

Phalacrocorax atriceps purpurascens (Brandt); Devillers & Terschuren 1978, Le Gerfaut 68: 76.

Phalacrocorax albiventer purpurascens (Brandt); Dorst & Mougin 1979, in Peters, Check-list Birds World 1 (2nd edition): 176.

Leucocarbo atriceps purpurascens (Brandt); Checklist Committee 1990, Checklist Birds N.Z.: 84.

Phalacrocorax purpurascens (Brandt); Marchant & Higgins 1990, HANZAB 1: 867.

Leucocarbo purpurascens (Brandt); Holdaway et al. 2001, New Zealand Journ. Zool. 28(2): 129, 177.

Breeds only on Macquarie Island and adjacent Bishop and Clerk Islets (Brothers 1985). Vagrant to Auckland Islands / Maukahuka (two specimens collected in 1901; Miskelly & Cooper 2020). Recognised as a full species following Marchant & Higgins (1990) and Holdaway et al. (2001).