Cormorant - Systematics - Species in Phylogenetic Sequence

Species in Phylogenetic Sequence

This list attempts to follow a phylogenetic order. If the distinction into subfamilies would be upheld, the "blue-eyed" and related species would probably be the Leucocarboninae, and the groups that follow them the Phalacrocoracinae. The first two lineages (and possibly the Flightless Cormorant) are basal and cannot be assigned to either subfamily.

Basal lineage 1: "Microcormorants", proposed genus Microcarbo or Halietor ("Phalacrocoracinae"); the former genus name would be valid.

Small, short-billed subtropical to tropical marine and freshwater species from the Old World and Australia. They have black feet and almost all lack significant white feathers. They often have a diminutive frontal tuft.

• Little Pied Cormorant, Phalacrocorax melanoleucos
• Reed Cormorant, Phalacrocorax africanus
• Crowned Cormorant, Phalacrocorax coronatus
• Little Cormorant, Phalacrocorax niger
• Pygmy Cormorant, Phalacrocorax pygmaeus

Basal lineage 2: Red-legged Cormorant. Included in Leucocarbo or Stictocarbo ("Leucocarboninae")

Pacific coast of South America. This species apparently has no close living relatives. It has a highly apomorphic color pattern: naked red base of bill, red feet, and a white neck spot, and it is crestless. It seems to be convergent in some aspects with the punctatus superspecies. What seems sure by now is that this species must be placed in a distinct monotypic genus Poikilocarbo in almost any case, if any species are split from Phalacrocorax at all.

• Red-legged Cormorant, Phalacrocorax gaimardi

Blue-eyed shags and relatives: variously placed in Euleucocarbo, Hypoleucos, Leucocarbo, Notocarbo and Stictocarbo ("Leucocarboninae"), and the monotypic Nannopterum.

This reasonably well-supported marine clade contains 3 lineages:

1. One containing American species which are mainly black-footed, black-plumaged, and have yellow skin at the base of the bill as well as white display crests behind the eyes in breeding plumage. They occur in marine and freshwater habitats. The Flightless Cormorant of the Galápagos Islands also seems to belong here. Its wings have been reduced by evolution to tiny size, it is extremely apomorphic due to its flightlessness, and its plumage is entirely nondescript. If considered a distinct genus, they would get the name Dilophalieus or (more probably) Nannopterum, the old genus of the Flightless Cormorant.
2. The Rock Shag from southern South America with red skin at the bill base, pink feet, a frontal crest, and an apomorphic white ear-spot
3. A group of numerous close-knit forms from southern Pacific and subantarctic waters which are white below with pink feet but otherwise quite varying in appearance. It contains the King and Imperial complexes and the Guanay Cormorant. Almost all have some amount of white on the upperwing coverts, frontal crests, and blue eye-rings. The crested shags with yellow warts in front of the eyes belong to this group. The genus name Leucocarbo would apply to either this group, or the entire clade.

• Double-crested Cormorant or White-crested Cormorant, Phalacrocorax auritus
• Neotropic Cormorant or Olivaceous Cormorant, Phalacrocorax brasilianus
• Flightless Cormorant, Phalacrocorax harrisi
• Rock Shag, Phalacrocorax magellanicus

• Imperial Shag or Blue-eyed Shag, Phalacrocorax atriceps
  • White-bellied Shag, Phalacrocorax (atriceps) albiventer
  • Antarctic Shag, Phalacrocorax (atriceps) bransfieldensis
  • South Georgia Shag, Phalacrocorax (atriceps) georgianus
  • Heard Island Shag, Phalacrocorax (atriceps) nivalis
  • Crozet Shag, Phalacrocorax (atriceps) melanogenis
  • Kerguelen Shag, Phalacrocorax (atriceps) verrucosus
  • Macquarie Shag, Phalacrocorax (atriceps) purpurascens
• Guanay Cormorant, Phalacrocorax bougainvillii
• Rough-faced Shag or King Shag, Phalacrocorax carunculatus
• Bronze Shag, Phalacrocorax chalconotus
• Chatham Shag, Phalacrocorax onslowi
• Auckland Shag, Phalacrocorax colensoi
• Campbell Shag, Phalacrocorax campbelli
• Bounty Shag, Phalacrocorax ranfurlyi

North Pacific shags: spread between Compsohalieus ("Phalacrocoracinae") and Stictocarbo ("Leucocarboninae"). If a distinct genus, the former name would apply

A well-supported marine group ranging from the Bering Strait to California. They are black-footed and have white ornamental plumes strewn about the head and neck in breeding plumage. They tend to have prominent double crests.

• Brandt's Cormorant, Phalacrocorax penicillatus
• Spectacled Cormorant, Phalacrocorax perspicillatus - extinct (c.1850)
• Pelagic Cormorant or Baird's Cormorant, Phalacrocorax pelagicus
• Red-faced Cormorant, Phalacrocorax urile

Common Shag lineage: formerly in Compsohalieus ("Phalacrocoracinae") and Stictocarbo ("Leucocarboninae")

Black-footed smallish marine shags of Europe and southern Africa. Wahlberg's Cormorant is very tentatively placed here; it seems anatomically more similar to the P. fuscscens, but the more informative characters - the combination of frontal crest and lack of extensive naked skin at bill base in mid-sized Old World species - seem to place it here. If this is correct, they are probably very distantly related due to biogeography.

• European Shag, Phalacrocorax aristotelis
• Bank Cormorant, Phalacrocorax neglectus - tentatively placed here

Indian Ocean group: spread between Hypoleucos and Leucocarbo ("Leucocarboninae") and Compsohalieus ("Phalacrocoracinae"). Hypoleucos would be the correct genus name if they were split off.

A group of black-footed species occurring in tropical coastal or inland habitat between the Persian Gulf and Australia. Most species are tentatively assigned here, based on the combination of range, crestlessness, size, general lack of naked skin ornaments and the presence of some amount of white feathering in the ear region at least in breeding plumage. This clade is not too well supported, but this may be because the two presumed members included in recent research are quite dissimilar; the three unstudied ones are very similar to one or the other.

• Little Black Cormorant, Phalacrocorax sulcirostris
• Indian Cormorant, Phalacrocorax fuscicollis - tentatively placed here
• Socotra Cormorant, Phalacrocorax nigrogularis - tentatively placed here
• Australian Pied Cormorant or Yellow-faced Cormorant, Phalacrocorax varius
• Black-faced Cormorant, Phalacrocorax fuscescens - tentatively placed here

Spotted group: placed in Stictocarbo ("Leucocarboninae"); indeed, they would be the only members of this possibly distinct genus

A superspecies of the New Zealand region. Peculiarly apomorphic, with yellowish legs, prominent double crests, white ornamental plumes on the neck, a grey belly and spotted wings.

• Spotted Shag Phalacrocorax punctatus
• Pitt Cormorant or Featherstone's Shag Phalacrocorax featherstoni

Cape Cormorant: sometimes placed in Leucocarbo ("Leucocarboninae")

Highly plesiomorphic among its relatives; a species from the southern coasts of Africa. It is apparently close to the common ancestor of the next group and, perhaps apart from the all-black plumage, looks almost identical to that long-extinct bird.

• Cape Cormorant, Phalacrocorax capensis

True cormorants: these would be retained in Phalacrocorax no matter how the cormorants and shags are split up

They occur from the western Atlantic through the Old World into Australia, usually but not always in marine and temperate to subtropical habitat. They are characteristic, being large, with white cheek and thigh patches, ornamental plumes in the neck, a yellow naked bill base, black feet, and a shaggy nape crest.

• Great Cormorant, Phalacrocorax carbo
• White-breasted Cormorant, Phalacrocorax lucidus
• Japanese Cormorant, Phalacrocorax capillatus