After Worthy, 2011 (fig. 4), with fossil taxa from Olson, 1985, Feduccia, 1996, Holdaway, Worthy & Tennyson, 2001, and living species from Sibley & Monroe, 1990, Peterson, 2002-2004, with Finnish vernacular names from Lokki, Jännes & Nikander, 1992
Note:
Microcarbo, Stictocarbo, Leucocarbo, Hypoleucos, etc. have been treated as genera, however, neither Monroe & Sibley (1990) nor Peterson (2002-2004) recognized them. In recent phylogenetic analysis by Worthy. He reinstitutes Microcarbo as a separate genus of primitive cormorants, but also shows paraphyly of Phalacrocorax. To maintain monophyly of Phalacrocorax I have decied to treat Leptocarbo as a subgenus.
<==o Phalacrocoracidae Reichenbach, 1849 (Cormorants and shags; merimetsot) |?-o †Borvocarbo Mourer-Chauviré, Berthet & Hugueney, 2004; L.-Olig.-E.Mioc. Eu. | |-- †B. guilloti Mourer-Chauviré, Berthet & Hugueney, 2004; L.Olig. WEu. | |?- ?†B. stoeffelensis Mayr, 2007; L.Olig. CEu. | `?- ?†B. tardatus Göhlich & Mourer-Chauviré, 2010; E.Mioc. CEu. |?- †Oligocorax littoralis (Milne-Edwards, 1863) [=? †Phalacrocorax littoralis]; E.Mioc. W.-CEu. |?-o †Limicorallus Kurochkin, 1968; L.Olig. CAs., ?E.Mioc. CEu. | |-- †L. saiensis Kurochkin, 1968; L.Olig. CAs. | `?- ?†L. carbunculus Mayr, 2009; E.Mioc. CEu. |?- †Pliocarbo longipes Tugarinov, 1940 |?- †Valenticarbo praetermissus |--o †Nambashag Worthy, 2011; L.Olig.-E.Mioc. Aust. | |-- †N. billerooensis Worthy, 2011 | `-- †N. microglaucus Worthy, 2011 `--+--o †Nectornis | |-- †N. miocaenus (Milne-Edwards, 1867 in Milne-Edwards, 1967-1871a) [=? †Phalacrocorax miocaenus] E.Mioc. W.-CEu. | `-- †N. anatolicus (Mourer-Chauviré in Paicheler et al., 1978) [=? †Phalacrocorax anatolicus] M.Mioc. WAs. `--+--o Microcarbo Bonaparte, 1856 [Halietor Heine, 1860] (pygmy cormorants; pikkumerimetsot) | |?- M. coronatus (Wahlberg, 1855) Oefv.Vet.-Akad.Forh. 12 p.214 | |?- M. niger (Vieillot, 1817) (jaavanmerimetso) | |--o M. melanoleucos (Vieillot, 1817) (Vieillot, 1817) (australianmerimetso; Australian Shag) | | |-- M. m. brevicauda (Mayr, 1931) | | |-- M. m. brevirostris (Gould, 1837) (White-throated Cormorant) | | `-- M. m. melanoleucos (Vieillot, 1817) (Little Pied Cormorant) | `--+-- M. pygmeus (Pallas, 1773) (pikkumerimetso) | `--o M. africanus (Gmelin, 1789) (kaislamerimetso) | |-- M. a. africanus (Gmelin, 1789) | `-- M. a. pictilis (Bangs, 1918) `--o Phalacrocorax Brisson, 1760 [incl. Australocorax Lambrecht, 1931; Compsohalieus B. Brewer & Ridgway, 1884; Cormoranus Baillon, 1834; Dilophalieus Coues, 1903; Ecmeles Gistel, 1848; Euleucocarbo Voisin, 1973; Hydrocorax Vieillot, 1819 (non Brisson, 1760: preoccupied); Hypoleucus Reichenbach, 1852; Leucocarbo Bonaparte, 1857; Miocorax Lambrecht, 1933; Nannopterum Sharpe, 1899; Nesocarbo Voisin, 1973; Notocarbo Siegel-Causey, 1988; Pallasicarbo Coues, 1903; Paracorax Lambrecht, 1933; Poikilocarbo Boetticher, 1935; Stictocarbo Bonaparte, 1855; Viguacarbo Coues, 1903] (cormorants and shags; merimetsot) |-- P. sulcirostris (Brandt, 1837) (mustamerimetso) `--+--o P. brasilianus (Gmelin, 1789) [Phalacrocorax olivaceus (Humboldt, 1805); Phalacrocorax pampeanus Moreno & Mercerat, 1891; Phalacrocorax vigua] (oliivimerimetso) | |-- P. b. brasilianus (Gmelin, 1789) | `-- P. b. mexicanus (Brandt, 1837) |--o P. auritus (Lesson, 1831) (amerikanmerimetso) | |-- P. a. albociliatus Ridgway, 1884 | |-- P. a. auritus (Lesson, 1831) | |-- P. a. cincinatus (Brandt, 1837) | |-- P. a. floridanus (Audubon, 1835) | `-- P. a. heuretus Watson, Olson & JR Miller, 1991 `--+--o P. (Hypoleucos) varius (Gmelin, 1789) (Kaunomerimetso) | |-- P. v. hypoleucos (Brandt, 1837) | `-- P. v. varius (Gmelin, 1789) `--+--o (Phalacrocorax) [sensu stricto?] | |?- P. capillatus (Temminck & Schlegel, 1850) (japaninmerimetso) | |?- P. lucidus (Lichtenstein, 1823) (afrikanmerimetso) | `--o P. carbo (Linnaeus, 1758) (merimetso) | |-- P. c. carbo (Linnaeus, 1758) (merimetso) | |-- P. c. hanedae Kuroda, 1925 | |-- P. c. maroccanus Hartert, 1906 | |-- P. c. novaehollandiae Stephens, 1826 | `-- P. c. sinensis (Blumenbach, 1798) `--+-- P. capensis (Sparrman, 1788) (kapinmerimetso) `--+-- P. fuscescens (Vieillot, 1817) (tasmanianmerimetso; Tasmanian Shag) `--+-- P. nigrogularis Ogilvie-Grant & HO Forbes, 1899 (arabianmerimetso) `--+-- P. neglectus (Wahlberg, 1855) (matalikkomerimetso) `--+--+--+-- P. penicillatus (Brandt, 1837) (tyrskymerimetso) | | `-- P. gaimardi (Lesson & Garnot, 1828) [Poikilocarbo Boetticher, 1935] (Red-legged Cormorant; harmaamerimetso) | `--+--o (Stictocarbo) | | |?- †P. kumeyaay [Stictocarbo kumeyaay] | | |-- P. featherstoni Buller, 1873 (huppumerimetso) | | `--o P. punctatus (Sparrman, 1786) (partamerimetso) | | |-- P. p. punctatus (Sparrman, 1786) (Spotted Shag) | | `-- P. p. oliveri Mathews, 1930 (Blue Shag) | `-+--o P. aristotelis (Linnaeus, 1761) (karimetso) | | |-- P. a. aristotelis (Linnaeus, 1761) | | |-- P. a. desmarestii (Payraudeau, 1826) | | `-- P. a. riggenbachi Hartert, 1923 | |?- †P. wetmorei Brodkorb, 1955 | `--+-- P. urile (Gmelin, 1789) (beringinmerimetso) | |?- †P. perspicillatus Pallas, 1811 (silmälasimerimetso) | `--o P. pelegicus Pallas, 1811 (ulappamerimetso) | |-- P. p. pelagicus Pallas, 1811 | `-- P. p. resplendens Audubon, 1838 `--o (Leucocarbo) Bonaparte, 1856 [Nesocarbo; Notocarbo; Euleucocarbo] (Shags) |?- †P. gregorii |?- †P. vetustus |?-o L. albiventer (Lesson, 1831) (Imperial Shag, Blue-eyed Shag; Kuningasmerimetso) | |-- P. (L.) a. albiventer (Lesson, 1831) | `-- P. (L.) a. melanogenis (Blyth, 1860) |?- P. (L.) carunculatus (Gmelin, 1789) (maorimerimetso) |?- P. (L.) colensoi (Buller, 1888) (Auckland Islands Shag) |?- P. (L.) ranfurlyi (Ogilvie-Grant, 1901) (Bounty Islands Shag) |-- P. (L.) bougainvillii (Lesson, 1837 (guanomerimetso) `--+-- P. (L.) onslowi HO Forbes, 1893 (Chatham Islands Shag) `--+-- P. (L.) chalconotus (G.R. Gray, 1845) (Bronzed Shag) |-- P. (L.) nivalis (Falla, 1937) [P. (L.) atriceps nivalis] |-- P. (L.) purpurascens (Brandt, 1837) [Notocarbo purpurascens; Leucocarbo albiventer purpurascens] |-- P. (L.) verrucosus (Cabanis, 1875) (kergueleninmerimetso; Kerguele Islands Shag) `--+--o P. (L.) atriceps (King, 1828) (etelänmerimetso) | |-- P. (L.) a. atriceps (King, 1828) | |-- P. (L.) a. bransfieldensis (Murphy, 1936) (Antarctic Shag) | `-- P. (L.) a. georgianus (Lonnberg, 1906) (etelägeorgianmerimetso; South Georgia Shag) `-- P. (L.) campbelli (Filhol, 1878) [Nesocarbo campbelli] (sepelmerimetso)
Untreated Phalacrocoracidae
†P. sp. [Lee Creek Formation, North Carolina] [Olson & Rasmussen, 2001] †P. femoralis Miller, 1929; L.Mioc. SWNAm. †P. leptopus †P. goletensis †P. kennelli †P. chapalensis †P. macer †P. rogersi †P. macropus †P. pampeanus †P. destefani †P. mongoliensis †P. reliquus †P. intermedius (Milne-Edwards, 1867 in Milne-Edwards, 1967-1871a) E.-M.Mioc. W.-CEu. †P. brunhuberi †P. ibericus Villalta, 1963; L.Mioc. SWEu. †P. lautus Kurochkin & Ganea, 1972; L.Mioc. SEEu. †P. marinavis Shufeldt, 1915; L.Mioc. WNAm. †P. serdicensis Burchak-Abramiovic & Nikolov, 1984; L.Mioc. SEEu. †P. longipes (Tugarinov, 1940); L.Mioc.-E.Plioc. SEEu. †P. filyawi Emslie, 1995b; L. Plioc. SENA. †P. sp. [from Sarasota County, Florida] †P. idahensis Marsh, 1870 P. fuscicollis Stephens, 1826 (intianmerimetso) P. harrisi Rothschild, 1898 [Nannopterum Sharpe, 1899; Dilophalieus Coues, 1903] (Flightless Cormorant; galapagosinmerimetso) P. coronatus (Wahlberg, 1855) (töyhtömerimetso) P. magellanicus (Gmelin, 1789) (kalliomerimetso)
Reference(s):
- Emslie, S. D., 1995: A catastrophic death assemblage of a new species of cormorant and other seabirds from the Late Pliocene of Florida.
–Journal of Vertebrate Paleontology: Vol. 15, #2, pp. 313-330 - Feduccia, A., 1980: The Age of Birds.
–Harvard University Press, Cambridge, Massachusetts & London, England, 1980, IX-196 - Feduccia, A., 1997: The Origin and Evolution of Birds.
–Yale University Press, New Haven, 1996, x-420 - Holdaway, R. N., Worthy, T. H. & Tennyson, A. J. D., 2001: A working list of breeding bird species of the New Zealand region at first human contact.
–New Zealand Journal of Zoology: Vol. 28, pp. 119-187 - Olson, S. L., 1975: Paleornithology of St. Helena Island, South Atlantic Ocean.
–Smithsonian Contributions to Baleobiology: Vol. 23, pp. iv-49 - Olson, S. L., 1985: The fossil record of birds. 79-239.
in Farner, D., King, J. & Parkes, K. (eds.), 1985: Avian Biology. vol VIII.
–Academic Press, New York. 1985. - Olson, S. L. & Rasmussen, P. D., 2001: Miocene and Pliocene birds from the Lee Creek Mine, North Carolina. 233-365
in Ray, C. E. & Bohaska, D. J., (eds.) 2001: Geology and paleontology of the Lee Creek Mine, North Carolina, III.
–Smithsonian Contributions to Paleobiology: Nr. 90, iii-365 - Peterson, A. P., 2002-2011: Zoonomen Nomenclatural data of Birds. (Updated: 2011-10-27)
–iNet: Zoonomen Nomenclatural data: http://www.zoonomen.net/avtax/frame.html; [visited: 2011-12-07] - Sibley, C. G. & Monroe, B. L., 1990: Distribution and taxonomy of birds of the world.
–Yale University Press, New Haven & London, 1990. xxiv-1111. - Worthy, T. H., 2011: Descriptions and phylogenetic relationships of a new genus and two new species of Oligo-Miocene cormorants (Aves: Phalacrocoracidae) from Australia.
–Zoological Journal of the Linnean Society: Vol. 163, #1, pp. 277-314 [doi: 10.1111/j.1096-3642.2011.00693.x]