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.
The spotted shag
Published 2 April 2025
Travel to any water body in New Zealand and you are likely to find shags among the more visible members of the bird fauna. No less than a dozen species of cormorant make their home in this country and its surrounding islands. Some represent local populations of more widespread species; others are found in New Zealand alone. Among the examples of the latter is the spotted shag Phalacrocorax punctatus.
The spotted shag is a marine species, nesting on cliffs and rocky outcrops around the North, South and Stewart Islands. It is more abundant in the southern part of its range, with populations around the northern part of the North Island being very small and localised. The spotted shag is easily distinguished from most other New Zealand cormorants by its predominantly grey plumage together with its orange bill and feet. Black specks on the grey back give the species its name. During the breeding season, the head is black with a pair of prominent crests on the head and nape; a patch of bright green bare skin is visible in front of the eye and a bright white stripe runs down the side of the head and neck. Outside the breeding season, the crests disappear, the colour of the bare skin dulls, and the white stripe becomes less obvious (Turbott 1956).
Cormorants have been regarded as a difficult group to classify owing to their structural uniformity, so their classification has shifted about over the years. Some have bypassed the issue by treating all shags and cormorants as a single genus Phalacrocorax; others have divided them between several genera. Within the New Zealand fauna, many authors have separated the spotted shag and its closest relative, the Pitt Island shag P. featherstoni, into their own genus Stictocarbo. The Pitt Island shag is an inhabitant of the Chatham Islands, east of the South Island, that closely resembles the spotted shag but is darker overall and lacks the white head-stripe. Some authors also included a number of cliff-dwelling marine shags from other parts of the world in Stictocarbo. However, recent phylogenetic analyses based primarily on molecular data have not supported the monophyly of the ‘cliff shags’. Instead, Kennedy & Spencer (2014) supported including P. punctatus and P. featherstoni within a broader clade of Indo-Australasian cormorants, which they included in Phalacrocorax.
Historically, two subspecies have been recognised for the spotted shag. The spotted shag Phalacrocorax punctatus punctatus was thought to inhabit the coasts of the North and northern South Islands, whereas the blue shag P. p. oliveri was found around the Fouveaux Strait between South and Stewart Islands. The blue shag was supposed to have a narrower white stripe on the head than the spotted shag. However, recognition of this distinction was confused by variation within the populations between individuals and by time of year. Analysis of the spotted shag’s molecular phylogeography by Rawlence et al. (2019) did not support recognition of the two subspecies as distinct. Instead, these authors identified the population of the northern North Island as the most isolated in the species. This population has declined rapidly in recent years though numbers in the southern part of the country remain robust. The species as a whole is not currently considered of concern conservation-wise, but its farther corners may still be cause for worry.
Systematics of Phalacrocorax
<==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
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