HOME
The Info List - Cormorant


--- Advertisement ---



3–43, see text

Synonyms

Australocorax Lambrecht, 1931 Compsohalieus B. Brewer & Ridgway, 1884 Cormoranus Baillon, 1834 Dilophalieus Coues, 1903 Ecmeles Gistel, 1848 Euleucocarbo Voisin, 1973 Halietor Heine, 1860 Hydrocorax Vieillot, 1819 (non Brisson, 1760: preoccupied) Hypoleucus Reichenbach, 1852 Leucocarbo Bonaparte, 1857 Microcarbo
Microcarbo
Bonaparte, 1856 Miocorax Lambrecht, 1933 Nannopterum Sharpe, 1899 Nesocarbo Voisin, 1973 Notocarbo Siegel-Causey, 1988 Pallasicarbo Coues, 1903 Paracorax Lambrecht, 1933 Poikilocarbo Boetticher, 1935 Pliocarbo Tugarinov, 1940 Stictocarbo Bonaparte, 1855 Viguacarbo Coues, 1903 (but see text)

Phalacrocoracidae
Phalacrocoracidae
is a family of some 40 species of aquatic birds commonly known as cormorants and shags. Several different classifications of the family have been proposed recently, and the number of genera is disputed. There is no consistent distinction between "cormorants" and "shags" as these appellations have been assigned to different species in these genera at various points in time. Cormorants and shags are medium-to-large birds, with body weight in the range of 0.35–5 kilograms (0.77–11.02 lb) and wing span of 45–100 centimetres (18–39 in). The majority of species have dark feathers. The bill is long, thin and hooked. Their feet have webbing between all four toes. All species are fish-eaters, catching the prey by diving from the surface. They are excellent divers, and under water they propel themselves with their feet with help from their wings; some cormorant species have been found to dive as deep as 45 metres. They have relatively short wings due to their need for economical movement underwater, and consequently have the highest flight costs of any bird.[1] Cormorants nest in colonies around the shore, on trees, islets or cliffs. They are coastal rather than oceanic birds, and some have colonised inland waters – indeed, the original ancestor of cormorants seems to have been a fresh-water bird. They range around the world, except for the central Pacific islands.

Contents

1 Names 2 Description 3 Habitat 4 Behaviour 5 Systematics

5.1 Species
Species
in HBW taxonomic sequence 5.2 Species
Species
in phylogenetic sequence 5.3 Evolution
Evolution
and fossil record

6 In human culture

6.1 Cormorant
Cormorant
fishing 6.2 In folklore, literature, and art

7 See also 8 Footnotes 9 References 10 External links

Names[edit] No consistent distinction exists between cormorants and shags. The names 'cormorant' and 'shag' were originally the common names of the two species of the family found in Great Britain, Phalacrocorax carbo (now referred to by ornithologists as the great cormorant) and P. aristotelis (the European shag). "Shag" refers to the bird's crest, which the British forms of the great cormorant lack. As other species were discovered by English-speaking sailors and explorers elsewhere in the world, some were called cormorants and some shags, depending on whether they had crests or not. Sometimes the same species is called a cormorant in one part of the world and a shag in another, e.g., the great cormorant is called the black shag in New Zealand
New Zealand
(the birds found in Australasia
Australasia
have a crest that is absent in European members of the species). Van Tets (1976) proposed to divide the family into two genera and attach the name "cormorant" to one and "shag" to the other, but this flies in the face of common usage and has not been widely adopted. The scientific genus name is Latinised Ancient Greek, from φαλακρός (phalakros, "bald") and κόραξ (korax, "raven").[2] This is often thought to refer to the creamy white patch on the cheeks of adult great cormorants, or the ornamental white head plumes prominent in Mediterranean birds of this species, but is certainly not a unifying characteristic of cormorants. "Cormorant" is a contraction derived either directly from Latin
Latin
corvus marinus, "sea raven" or through Brythonic Celtic. Cormoran
Cormoran
is the Cornish name of the sea giant in the tale of Jack the Giant Killer. Indeed, "sea raven" or analogous terms were the usual terms for cormorants in Germanic languages
Germanic languages
until after the Middle Ages. The French explorer André Thévet
André Thévet
commented in 1558, "...the beak [is] similar to that of a cormorant or other corvid," which demonstrates that the erroneous belief that the birds were related to ravens lasted at least to the 16th century. Description[edit]

Great cormorant
Great cormorant
with hooked bill

Cormorants and shags are medium-to-large seabirds. They range in size from the pygmy cormorant (Phalacrocorax pygmaeus), at as little as 45 cm (18 in) and 340 g (12 oz), to the flightless cormorant (Phalacrocorax harrisi), at a maximum size 100 cm (39 in) and 5 kg (11 lb). The recently extinct spectacled cormorant (Phalacrocorax perspicillatus) was rather larger, at an average size of 6.3 kg (14 lb). The majority, including nearly all Northern Hemisphere species, have mainly dark plumage, but some Southern Hemisphere species are black and white, and a few (e.g. the spotted shag of New Zealand) are quite colourful. Many species have areas of coloured skin on the face (the lores and the gular skin) which can be bright blue, orange, red or yellow, typically becoming more brightly coloured in the breeding season. The bill is long, thin, and sharply hooked. Their feet have webbing between all four toes, as in their relatives. Habitat[edit]

Imperial shags in Beagle Channel

They are coastal rather than oceanic birds, and some have colonised inland waters – indeed, the original ancestor of cormorants seems to have been a fresh-water bird, judging from the habitat of the most ancient lineage. They range around the world, except for the central Pacific islands. Behaviour[edit] All are fish-eaters, dining on small eels, fish, and even water snakes. They dive from the surface, though many species make a characteristic half-jump as they dive, presumably to give themselves a more streamlined entry into the water. Under water they propel themselves with their feet, though some also propel themselves with their wings (see the picture,[3] commentary [4] and existing reference video [5]). Some cormorant species have been found, using depth gauges, to dive to depths of as much as 45 metres.

Wing-drying behaviour

After fishing, cormorants go ashore, and are frequently seen holding their wings out in the sun. All cormorants have preen gland secretions that are used ostensibly to keep the feathers waterproof. Some sources[6] state that cormorants have waterproof feathers while others say that they have water permeable feathers.[7][8] Still others suggest that the outer plumage absorbs water but does not permit it to penetrate the layer of air next to the skin.[9] The wing drying action is seen even in the flightless cormorant but commonly in the Antarctic shags[10] and red-legged cormorants. Alternate functions suggested for the spread-wing posture include that it aids thermoregulation,[11] digestion, balances the bird or indicates presence of fish. A detailed study of the great cormorant concludes that it is without doubt[12] to dry the plumage.[13][14] Cormorants are colonial nesters, using trees, rocky islets, or cliffs. The eggs are a chalky-blue colour. There is usually one brood a year. The young are fed through regurgitation. They typically have deep, ungainly bills, showing a greater resemblance to those of the pelicans, to which they are related, than is obvious in the adults. Systematics[edit] The cormorants are a group traditionally placed within the Pelecaniformes
Pelecaniformes
or, in the Sibley–Ahlquist taxonomy, the expanded Ciconiiformes. This latter group is certainly not a natural one, and even after the tropicbirds have been recognised as quite distinct, the remaining Pelecaniformes
Pelecaniformes
seem not to be entirely monophyletic. Their relationships and delimitation – apart from being part of a "higher waterfowl" clade which is similar but not identical to Sibley and Ahlquist's "pan-Ciconiiformes" – remain mostly unresolved. Notwithstanding, all evidence agrees that the cormorants and shags are closer to the darters and Sulidae
Sulidae
(gannets and boobies), and perhaps the pelicans or even penguins, than to all other living birds.[15] In recent years, three preferred treatments of the cormorant family have emerged: either to leave all living cormorants in a single genus, Phalacrocorax, or to split off a few species such as the imperial shag complex (in Leucocarbo) and perhaps the flightless cormorant. Alternatively, the genus may be disassembled altogether and in the most extreme case be reduced to the great, white-breasted and Japanese cormorants.[16] Pending a thorough review of the Recent and prehistoric cormorants, the single-genus approach[17] is followed here for three reasons: first, it is preferable to tentatively assigning genera without a robust hypothesis. Second, it makes it easier to deal with the fossil forms, the systematic treatment of which has been no less controversial than that of living cormorants and shags. Third, this scheme is also used by the IUCN,[18] making it easier to incorporate data on status and conservation. In accordance with the treatment there, the imperial shag complex is here left unsplit as well, but the king shag complex has been.

Occipital crest or os nuchale in Phalacrocorax carbo

The cormorants and the darters have a unique bone on the back of the top of the skull known as the os nuchale or occipital style which was called a xiphoid process in early literature. This bony projection provides anchorage for the muscles that increase the force with which the lower mandible is closed[19][20] and this bone and the highly developed muscles over it, the M. adductor mandibulae caput nuchale are unique to the families Phalacrocoracidae
Phalacrocoracidae
and Anhingidae.[21][22] Several evolutionary groups are still recognizable. However, combining the available evidence suggests that there has also been a great deal of convergent evolution; for example the cliff shags are a convergent paraphyletic group. The proposed division into Phalacrocorax sensu stricto (or subfamily "Phalacrocoracinae") cormorants and Leucocarbo sensu lato (or "Leucocarboninae") shags[23] does indeed have some degree of merit.[24] The resolution provided by the mtDNA 12S rRNA
12S rRNA
and ATPase
ATPase
subunits six and eight sequence data[24] is not sufficient to properly resolve several groups to satisfaction; in addition, many species remain unsampled, the fossil record has not been integrated in the data, and the effects of hybridisation – known in some Pacific species especially – on the DNA sequence
DNA sequence
data are unstudied. Species
Species
in HBW taxonomic sequence[edit]

Cormorant
Cormorant
(species unknown) begins its dive

Immature Phalacrocorax atriceps albiventer

Phalacrocorax niger
Phalacrocorax niger
in Hyderabad, India

Guanay cormorant
Guanay cormorant
(Phalacrocorax bougainvillii) at Weltvogelpark Walsrode

This sequence follows the Handbook of the Birds of the World.[17]

Double-crested cormorant
Double-crested cormorant
or white-crested cormorant, Phalacrocorax auritus Neotropic cormorant
Neotropic cormorant
or olivaceous cormorant, Phalacrocorax brasilianus (or Phalacrocorax olivaceus) Little black cormorant, Phalacrocorax sulcirostris Great cormorant
Great cormorant
or black shag, Phalacrocorax carbo White-breasted cormorant, Phalacrocorax lucidus Indian cormorant, Phalacrocorax fuscicollis Cape cormorant, Phalacrocorax capensis Socotra cormorant, Phalacrocorax nigrogularis Bank cormorant
Bank cormorant
or Wahlberg's cormorant, Phalacrocorax neglectus Japanese cormorant
Japanese cormorant
or Temminck's cormorant, Phalacrocorax capillatus Brandt's cormorant, Phalacrocorax penicillatus Spectacled cormorant, Phalacrocorax perspicillatus – extinct (c. 1850) European shag
European shag
or common shag, Phalacrocorax aristotelis Pelagic cormorant
Pelagic cormorant
or Baird's cormorant, Phalacrocorax pelagicus Red-faced cormorant, Phalacrocorax urile Rock shag, Phalacrocorax magellanicus Australian pied cormorant
Australian pied cormorant
or yellow-faced cormorant, Phalacrocorax varius Black-faced cormorant, Phalacrocorax fuscescens Red-legged cormorant
Red-legged cormorant
or red-footed shag, Phalacrocorax gaimardi Spotted shag
Spotted shag
Phalacrocorax punctatus Pitt shag
Pitt shag
or Featherstone's shag Phalacrocorax featherstoni Flightless cormorant, Phalacrocorax harrisi Guanay cormorant, Leucocarbo bougainvillii New Zealand
New Zealand
king shag or rough-faced shag, Leucocarbo carunculatus Otago shag
Otago shag
or bronze shag, Leucocarbo chalconotus Foveaux shag
Foveaux shag
or bronze shag, Leucocarbo stewarti Chatham shag, Leucocarbo onslowi Auckland shag, Leucocarbo colensoi Campbell shag, Leucocarbo campbelli Bounty shag, Leucocarbo ranfurlyi Imperial shag
Imperial shag
or blue-eyed shag, Leucocarbo atriceps Antarctic shag, Leucocarbo bransfieldensis South Georgia shag, Leucocarbo georgianus Heard Island shag, Leucocarbo nivalis Crozet shag, Leucocarbo melanogenis Kerguelen shag, Leucocarbo verrucosus Macquarie shag, Leucocarbo purpurascens Little pied cormorant, Microcarbo
Microcarbo
melanoleucos Reed cormorant
Reed cormorant
or long-tailed cormorant, Microcarbo
Microcarbo
africanus Crowned cormorant, Microcarbo
Microcarbo
coronatus Little cormorant, Microcarbo
Microcarbo
niger Pygmy cormorant, Microcarbo
Microcarbo
pygmaeus

Species
Species
in phylogenetic sequence[edit]

Little cormorant, Microcarbo
Microcarbo
niger

Red-legged cormorant
Red-legged cormorant
(Phalacrocorax gaimardi)

This list attempts to follow a phylogenetic order.[25] 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", genus Microcarbo
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
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, Microcarbo
Microcarbo
melanoleucos Reed cormorant, Microcarbo
Microcarbo
africanus Crowned cormorant, Microcarbo
Microcarbo
coronatus Little cormorant, Microcarbo
Microcarbo
niger Pygmy cormorant, Microcarbo
Microcarbo
pygmeus

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 colour pattern - naked red base of bill, red feet, and a white neck spot -, and is crestless. It seems to be convergent in some aspects with the punctatus superspecies ("Stictocarbo" proper). What seems sure 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.[26]

Red-legged cormorant, Phalacrocorax gaimardi

The double-crested cormorant's crests are normally not visible

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 three lineages: One containing American species which are mainly black-footed, blackish-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 a 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. 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 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 sometimes to this group.

Double-crested cormorant
Double-crested cormorant
or white-crested cormorant, Phalacrocorax auritus Neotropic cormorant
Neotropic cormorant
or olivaceous cormorant, Phalacrocorax brasilianus Flightless cormorant, Phalacrocorax harrisi Rock shag, Phalacrocorax magellanicus

The following are more accepted in the group:

Guanay cormorant, Leucocarbo bougainvillii

Imperial shag
Imperial shag
or blue-eyed shag, Leucocarbo atriceps

White-bellied shag, Leucocarbo albiventer

Antarctic shag, Leucocarbo bransfieldensis South Georgia shag, Leucocarbo georgianus Heard Island shag, Leucocarbo nivalis Crozet shag, Leucocarbo melanogenis Kerguelen shag, Leucocarbo verrucosus Macquarie shag, Leucocarbo purpurascens Guanay cormorant, Leucocarbo bougainvillii Rough-faced shag
Rough-faced shag
or king shag, Leucocarbo carunculatus Otago shag
Otago shag
or bronze shag, Leucocarbo chalconotus Foveaux shag
Foveaux shag
or bronze shag, Leucocarbo stewarti Chatham shag, Leucocarbo onslowi Auckland shag, Leucocarbo colensoi Campbell shag, Leucocarbo campbelli Bounty shag, Leucocarbo ranfurlyi

Brandt's cormorant
Brandt's cormorant
(Phalacrocorax penicillatus) – crestless, but with ornamental plumes

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
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
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. The Bank cormorant
Bank 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
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[27]

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

Little black cormorant, Phalacrocorax sulcirostris

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[24] are quite dissimilar; the three unstudied ones are very similar to one or the other.[17]

Little black cormorant, Phalacrocorax sulcirostris Indian cormorant, Phalacrocorax fuscicollis – tentatively placed here Socotra cormorant, Phalacrocorax nigrogularis – tentatively placed here Australian pied cormorant
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
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
Spotted shag
Phalacrocorax punctatus Pitt shag
Pitt shag
or Featherstone's shag Phalacrocorax featherstoni

Great cormorant
Great cormorant
drying its wings

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
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

Evolution
Evolution
and fossil record[edit] Cormorants seem to be a very ancient group, with similar ancestors reaching back to the time of the dinosaurs. In fact, the earliest known modern bird, Gansus yumenensis, had essentially the same structure. The details of the evolution of the cormorant are mostly unknown. Even the technique of using the distribution and relationships of a species to figure out where it came from, biogeography, usually very informative, does not give very specific data for this probably rather ancient and widespread group. However, the closest living relatives of the cormorants and shags are the other families of the suborder Sulae—darters and gannets and boobies—which have a primarily Gondwanan distribution. Hence, at least the modern diversity of Sulae
Sulae
probably originated in the southern hemisphere. While the Leucocarbonines are almost certainly of southern Pacific origin—possibly even the Antarctic which, at the time when cormorants evolved, was not yet ice-covered—all that can be said about the Phalacrocoracines is that they are most diverse in the regions bordering the Indian Ocean, but generally occur over a large area. Similarly, the origin of the family is shrouded in uncertainties. Some Late Cretaceous fossils have been proposed to belong with the Phalacrocoracidae: A scapula from the Campanian- Maastrichtian
Maastrichtian
boundary, about 70 mya (million years ago), was found in the Nemegt Formation
Nemegt Formation
in Mongolia; it is now in the PIN collection.[28] It is from a bird roughly the size of a spectacled cormorant, and quite similar to the corresponding bone in Phalacrocorax. A Maastrichtian
Maastrichtian
(Late Cretaceous, c. 66 mya) right femur, AMNH
AMNH
FR 25272 from the Lance Formation
Lance Formation
near Lance Creek, Wyoming, is sometimes suggested to be the second-oldest record of the Phalacrocoracidae; this was from a rather smaller bird, about the size of a long-tailed cormorant.[29] As the Early Oligocene
Early Oligocene
Sula" ronzoni cannot be assigned to any of the suloid families—cormorants and shags, darters, and gannets and boobies—with certainty, the best interpretation is that the Phalacrocoracidae
Phalacrocoracidae
diverged from their closest ancestors in the Early Oligocene, perhaps some 30 million years ago, and that the Cretaceous fossils represent ancestral suloids, "pelecaniforms" or "higher waterbirds"; at least the last lineage is generally believed to have been already distinct and undergoing evolutionary radiation at the end of the Cretaceous. What can be said with near certainty is that AMNH FR 25272 is from a diving bird that used its feet for underwater locomotion; as this is liable to result in some degree of convergent evolution and the bone is missing indisputable neornithine features, it is not entirely certain that the bone is correctly referred to this group.[30] During the late Paleogene, when the family presumably originated, much of Eurasia was covered by shallow seas, as the Indian Plate
Indian Plate
finally attached to the mainland. Lacking a detailed study, it may well be that the first "modern" cormorants were small species from eastern, south-eastern or southern Asia, possibly living in freshwater habitat, that dispersed due to tectonic events. Such a scenario would account for the present-day distribution of cormorants and shags and is not contradicted by the fossil record; as remarked above, a thorough review of the problem is not yet available.

Double-crested cormorant

Two distinct genera of prehistoric cormorants are widely accepted today, if Phalacrocorax is used for all living species:

Limicorallus (Indricotherium middle Oligocene
Oligocene
of Chelkar-Teniz, Kazakhstan) Nectornis (Late Oligocene/Early Miocene of Central Europe – Middle Miocene of Bes-Konak, Turkey) – includes Oligocorax miocaenus

The proposed genus Oligocorax appears to be paraphyletic – the European species have been separated in Nectornis, and the North American ones are placed in the expanded Phalacrocorax. A Late Oligocene
Oligocene
fossil cormorant foot from Enspel, Germany, sometimes placed herein, would then be referable to Nectornis if it proves not to be too distinct. All these early European species might belong to the basal group of "microcormorants", as they conform with them in size and seem to have inhabited the same habitat: subtropical coastal or inland waters. Limicorallus, meanwhile, was initially believed to be a rail or a dabbling duck by some. There are also undescribed remains of apparent cormorants from the Quercy
Quercy
Phosphorites of Quercy
Quercy
(France), dating to some time between the Late Eocene
Late Eocene
and the mid-Oligocene. Some other Paleogene remains are sometimes assigned to the Phalacrocoracidae, but these birds seem quite intermediate between cormorants and darters (and lack clear autapomorphies of either). Thus, they may be quite basal members of the Palacrocoracoidea. The taxa in question are:

Piscator ( Late Eocene
Late Eocene
of England) "Pelecaniformes" gen. et sp. indet. (Jebel Qatrani Early Oligocene
Early Oligocene
of Fayum, Egypt) – similar to Piscator? Borvocarbo ( Late Oligocene of C Europe)

The supposed Late Pliocene/Early Pleistocene "Valenticarbo" is a nomen dubium and given its recent age probably not a separate genus. The remaining species are, in accordance with the scheme used in this article, all placed in the modern genus Phalacrocorax:

Phalacrocorax marinavis ( Oligocene
Oligocene
– Early Miocene of Oregon, US) – formerly Oligocorax Phalacrocorax littoralis (Late Oligocene/Early Miocene of St-Gérand-le-Puy, France) – formerly Oligocorax, might belong into Nectornis Phalacrocorax intermedius (Early – Middle Miocene of C Europe) – includes P. praecarbo, Ardea/P. brunhuberi and Botaurites avitus Phalacrocorax macropus (Early Miocene – Pliocene of north-west US) Phalacrocorax ibericus (Late Miocene of Valles de Fuentiduena, Spain) Phalacrocorax lautus (Late Miocene of Golboçica, Moldavia) Phalacrocorax serdicensis (Late Miocene of Hrabarsko, Bulgaria) Phalacrocorax femoralis (Modelo Late Miocene/Early Pliocene of WC North America) – formerly Miocorax Phalacrocorax sp. (Late Miocene/Early Pliocene of Lee Creek Mine, US) Phalacrocorax longipes (Late Miocene – Early Pliocene of the Ukraine) – formerly Pliocarbo Phalacrocorax goletensis (Early Pliocene – Early Pleistocene of Mexico) Phalacrocorax wetmorei (Bone Valley Early Pliocene of Florida) Phalacrocorax sp. (Bone Valley Early Pliocene of Polk County, Florida, US)[31] Phalacrocorax leptopus (Juntura Early/Middle Pliocene of Juntura, Malheur County, Oregon, US) Phalacrocorax idahensis (Middle Pliocene – Pleistocene of Idaho, US) Phalacrocorax destefanii[verification needed] (Late Pliocene of Italy) – formerly Paracorax Phalacrocorax filyawi (Pinecrest Late Pliocene of Florida, US) – may be P. idahensis Phalacrocorax kumeyaay (San Diego Late Pliocene of California, US) Phalacrocorax macer (Late Pliocene of Idaho, US) Phalacrocorax mongoliensis (Late Pliocene of W Mongolia) Phalacrocorax rogersi (Late Pliocene – Early Pleistocene of California, US) Phalacrocorax kennelli (San Diego Pliocene of California, US) Phalacrocorax sp. "Wildhalm"[verification needed] (Pliocene) – may be same as P. longipes[32] Phalacrocorax chapalensis (Late Pliocene/Early Pleistocene of Jalisco, Mexico Phalacrocorax gregorii (Late Pleistocene of Australia) – possibly not a valid species Phalacrocorax vetustus (Late Pleistocene of Australia) – formerly Australocorax, possibly not a valid species Phalacrocorax reliquus Phalacrocorax sp. (Sarasota County, Florida, US) – may be P. filawyi/idahensis

The former "Phalacrocorax" (or "Oligocorax") mediterraneus is now considered to belong to the bathornithid Paracrax antiqua.[33] "P." subvolans was actually a darter (Anhinga). In human culture[edit] Cormorant
Cormorant
fishing[edit] Main article: Cormorant
Cormorant
fishing

A Chinese fisherman with his two cormorants

Humans have used cormorants' fishing skills in various places in the world. Archaeological evidence suggests that cormorant fishing was practiced in Ancient Egypt, Peru, Korea and India, but the strongest tradition has remained in China and Japan, where it reached commercial-scale level in some areas.[34] In Japan, cormorant fishing is called ukai (鵜飼). Traditional forms of ukai can be seen on the Nagara River
Nagara River
in the city of Gifu, Gifu
Gifu, Gifu
Prefecture, where cormorant fishing has continued uninterrupted for 1300 years, or in the city of Inuyama, Aichi. In Guilin, China, cormorants are famous for fishing on the shallow Lijiang River. In Gifu, the Japanese cormorant
Japanese cormorant
(P. capillatus) is used; Chinese fishermen often employ great cormorants (P. carbo).[35] In Europe, a similar practice was also used on Doiran Lake in the region of Macedonia.[36] In a common technique, a snare is tied near the base of the bird's throat, which allows the bird only to swallow small fish. When the bird captures and tries to swallow a large fish, the fish is caught in the bird's throat. When the bird returns to the fisherman's raft, the fisherman helps the bird to remove the fish from its throat. The method is not as common today, since more efficient methods of catching fish have been developed, but is still practiced as a cultural tradition.[35][34] In folklore, literature, and art[edit] Further information: Birds in culture Cormorants feature in heraldry and medieval ornamentation, usually in their "wing-drying" pose, which was seen as representing the Christian cross, and symbolizing nobility and sacrifice. For John Milton
John Milton
in Paradise Lost, the cormorant symbolizes greed: perched atop the Tree of Life, Satan
Satan
took the form of a cormorant as he spied on Adam and Eve during his first intrusion into Eden.[37] In some Scandinavian areas, they are considered good omen; in particular, in Norwegian tradition spirits of those lost at sea come to visit their loved ones disguised as cormorants.[37] For example, the Norwegian municipalities of Røst, Loppa
Loppa
and Skjervøy
Skjervøy
have cormorants in their coat of arms. The symbolic liver bird of Liverpool is commonly thought to be a cross between an eagle and a cormorant.

Cormorants catching Fish. Hanging silk scroll by Yūhi, Middle Edo period, Japan, 1755

In 1853, a woman wearing a dress made of cormorant feathers was found on San Nicolas Island, off the southern coast of California. She had sewn the feather dress together using whale sinews. She is known as the Lone Woman of San Nicolas and was later baptised "Juana Maria" (her original name is lost). The woman had lived alone on the island for 18 years before being rescued. When removed from San Nicolas, she brought with her a green cormorant dress she made; this dress is reported to have been removed to the Vatican.[citation needed] The bird has inspired numerous writers, including Amy Clampitt, who wrote a poem called "The Cormorant
Cormorant
in its Element". The species she described may have been the pelagic cormorant, which is the only species in the temperate U.S. with the "slim head ... vermilion-strapped" and "big black feet" that she mentions.[citation needed] A cormorant representing Blanche Ingram
Blanche Ingram
appears in the first of the fictional paintings by Jane in Charlotte Brontë's novel Jane Eyre.[citation needed] The cormorant served as the hood ornament for the Packard
Packard
automobile brand.[38] Cormorants (and books about them written by a fictional ornithologist) are a recurring fascination of the protagonist in Jesse Ball's 2018 novel Census. See also[edit]

Anhinga Cormorant
Cormorant
culling

Footnotes[edit]

^ Elliott KH, Ricklefs RE, Gaston AJ, Hatch SA, Speakman JR, Davoren GK. 2013. High flight costs and low dive costs support the biomechanical hypothesis for flightlessness in penguins. PNAS 110:9380-9384. [1] ^ Jobling, James A (2010). The Helm Dictionary of Scientific Bird Names. London: Christopher Helm. p. 301. ISBN 978-1-4081-2501-4.  ^ "www.nwdiveclub.com/download/file.php?id=22712&mode=view". nwdiveclub.com.  ^ "Birds diving beyond 50ft down and going horizontally there?! - Northwest Dive Club".  ^ "WCS Newsroom".  ^ Cramp S, Simmons KEL (1977) Handbook of the Birds of the Western Palearctic Volume 1, Oxford University Press ISBN 0-19-857358-8 ^ Rijke AM (1968). "The water repellency and feather structure of cormorants, Phalacrocoracidae". J. Exp. Biol. 48: 185–189.  ^ Marchant S. M.; Higgins, P. J. (1990). Handbook of Australian, New Zealand and Antarctic Birds. Vol 1A. Oxford University Press.  ^ Hennemann, W. W., III (1984). "Spread-winged behaviour of double-crested and flightless cormorants Phalacrocorax auritus and P. harrisi: wing drying or thermoregulation?". Ibis. 126 (2): 230–239. doi:10.1111/j.1474-919X.1984.tb08002.x. CS1 maint: Multiple names: authors list (link) ^ Cook, Timothee R; Guillaume Leblanc (2007). "Why is wing-spreading behaviour absent in blue-eyed shags?" (PDF). Animal
Animal
Behaviour. 74 (3): 649–652. doi:10.1016/j.anbehav.2006.11.024.  ^ Curry-Lindahl, K (1970). "Spread-wing postures in Pelecaniformes
Pelecaniformes
and Ciconiiformes" (PDF). Auk. 87 (2): 371–372. doi:10.2307/4083936. JSTOR 4083936.  ^ Sellers, R. M. (1995). "Wing-spreading behavior of the cormorant Phalacrocorax carbo" (PDF). Ardea. 83: 27–36.  ^ Nelson, J. Bryan (2005). Pelicans, Cormorants and Their Relatives: Pelecanidae, Sulidae, Phalacrocoracidae, Anhingidae, Fregatidae, Phaethontidae. Oxford University Press. pp. 162–163. ISBN 0-19-857727-3.  ^ Bernstein, N. P; S J Maxson (1982). "Absence of Wing-spreading Behavior in the Antarctic Blue-eyed Shag (Phalacrocorax Atriceps Bransfieldensis)" (PDF). The Auk. 99 (3): 588–589.  ^ Kennedy et al. (2000), Mayr (2005) ^ See Siegel-Causey (1988), Orta (1992) and Kennedy et al. (2000) for a review of classification schemes. ^ a b c Orta (1992) ^ IUCN
IUCN
(2007) ^ Yarrell, William (1828). "On the xiphoid bone and its muscles in the Corvorant (Pelecanus carbo)". The Zoological Journal. 4: 234–237.  ^ Garrod, A. H. (2009). "1. Notes on the Anatomy of Plotus anhinga". Proceedings of the Zoological Society of London. 44: 335. doi:10.1111/j.1096-3642.1876.tb02572.x.  ^ Burger, A E (2015). "Functional Anatomy of the Feeding Apparatus of Four South African Cormorants". Zoologica Africana. 13: 81. doi:10.1080/00445096.1978.11447608.  ^ Shufeldt, R.W. (1915). "Comparative osteology of Harris's Flightless Cormorant
Cormorant
(Nannopterum harrisi)". Emu. 15 (2): 86. doi:10.1071/MU915086.  ^ van Tets (1976), Siegel-Causey (1988) ^ a b c Kennedy et al. (2000) ^ Based on Orta (1992) and Kennedy et al. (2000). Applicable genus names are from Dorst & Mougin (1979) and Orta (1992) ^ Much of Phalacrocoracidae
Phalacrocoracidae
systematics hinges upon this most enigmatic species. The white neck spots and general colouration are very much unlike that of any other living cormorant, though anatomically it is quite similar to the species composing the punctatus superspecies, which are also the only other members of this family with a grey background colour. No satisfying theory has been proposed to explain this oddity. ^ According to Dorst & Mougin (1979), close to the Cape cormorant and possibly the spotted group. Orta (1992) disagrees – see HBW sequence above – but does not give details. As this species and the cape cormorant occupy ranges that almost entirely overlap, it is not too likely that they are closely related. If they were, their ancestors would probably have at one time been restricted to refugia and afterwards, expanding their ranges again, evolved into a striking case of character displacement. ^ Kurochkin (1995) ^ Hope (2002) ^ Hope (2002) and see Hesperornithes ^ A proximal ulna, Specimen PB 311, Pierce Brodkorb
Pierce Brodkorb
collection. Initially assigned to P. idahensis. However, it is far too large, being from a very big species possibly larger than a great cormorant: Murray (1970) ^ At least part of a coracoid is known. Does not appear to belong to the true cormorants. May have been closer in habitus to North Pacific shags, but not closely related[verification needed] to these: Howard (1932). ^ Cracraft (1971) ^ a b Richard J. King (1 October 2013). The Devil's Cormorant: A Natural History. University of New Hampshire Press. pp. 9–. ISBN 978-1-61168-225-0.  ^ a b " Cormorant
Cormorant
Fishing "UKAI"". May 2001. Retrieved 23 June 2016.  ^ "About Dojran lake". Retrieved 23 June 2016.  ^ a b Arin Murphy-Hiscock (18 January 2012). Birds - A Spiritual Field Guide: Explore the Symbology and Significance of These Divine Winged Messengers. Adams Media. pp. 48–49. ISBN 1-4405-2688-5.  ^ John Gunnell (January 2004). Standard Guide to 1950s American Cars. Krause Publications. p. 192. ISBN 0-87349-868-2. 

References[edit]

Benson, Elizabeth (1972): The Mochica: A Culture of Peru. Praeger Press, New York. Berrin, Katherine & Larco Museum
Larco Museum
(1997) The Spirit of Ancient Peru: Treasures from the Museo Arqueológico Rafael Larco Herrera. Thames and Hudson, New York. Cracraft, Joel (1971). "Systematics and evolution of the Gruiformes (Class Aves). 2. Additional comments on the Bathornithidae, with descriptions of new species" (PDF). American Museum Novitates. 2449: 1–14.  Dorst, J. & Mougin, J.L. (1979): Family Phalacrocoracidae. In: Mayr, Ernst & Cottrell, G.W. (eds.): Check-List of the Birds of the World Vol. 1, 2nd ed. (Struthioniformes, Tinamiformes, Procellariiformes, Sphenisciformes, Gaviiformes, Podicipediformes, Pelecaniformes, Ciconiiformes, Phoenicopteriformes, Falconiformes, Anseriformes): 163–179. Museum of Comparative Zoology, Cambridge. Hope, Sylvia (2002): The Mesozoic radiation of Neornithes. In: Chiappe, Luis M. & Witmer, Lawrence M. (eds.): Mesozoic Birds: Above the Heads of Dinosaurs: 339–388. ISBN 0-520-20094-2 Howard, Hildegarde (1932). "A New Species
Species
of Cormorant
Cormorant
from Pliocene Deposits near Santa Barbara, California" (PDF). Condor. 34 (3): 118–120. doi:10.2307/1363540. JSTOR 1363540.  IUCN
IUCN
(2007): 2007 IUCN
IUCN
Red List of Threatened Species. IUCN, Gland. Kennedy, M.; Gray, R.D.; Spencer H.G. (2000). "The Phylogenetic Relationships of the Shags and Cormorants: Can Sequence Data Resolve a Disagreement between Behavior and Morphology?" (PDF). Molecular Phylogenetics and Evolution. 17 (3): 345–359. doi:10.1006/mpev.2000.0840. PMID 11133189. Archived from the original (PDF) on 2007-04-18.  Kurochkin, Evgeny N. (1995). "Synopsis of Mesozoic birds and early evolution of Class Aves" (PDF). Archaeopteryx. 13: 47–66. Archived from the original (PDF) on 2007-09-27.  Mayr, Gerald (2005). "Tertiary plotopterids (Aves, Plotopteridae) and a novel hypothesis on the phylogenetic relationships of penguins (Spheniscidae)" (PDF). Journal of Zoological Systematics. 43 (1): 67–71. doi:10.1111/j.1439-0469.2004.00291.x.  Murray, Bertram G., Jr. (1970). "A Redescription of Two Pliocene Cormorants" (PDF). Condor. 72 (3): 293–298. doi:10.2307/1366006. JSTOR 1366006. CS1 maint: Multiple names: authors list (link) Orta, Jaume (1992): Family Phalacrocoracidae. In: del Hoyo, Josep; Elliott, Andrew & Sargatal, Jordi (eds.): Handbook of Birds of the World, Volume 1 (Ostrich to Ducks): 326–353, plates 22–23. Lynx Edicions, Barcelona. ISBN 84-87334-10-5 Robertson, Connie (1998): Book of Humorous Quotations. Wordsworth Editions. ISBN 1-85326-759-7 Siegel-Causey, Douglas (1988). "Phylogeny of the Phalacrocoracidae" (PDF). Condor. 90 (4): 885–905. doi:10.2307/1368846. JSTOR 1368846.  Thevet, F. André (1558): About birds of Ascension Island. In: Les singularitez de la France Antarctique, autrement nommee Amerique, & de plusieurs terres & isles decouvertes de nostre temps: 39–40. Maurice de la Porte heirs, Paris. van Tets, G. F. (1976): Australasia
Australasia
and the origin of shags and cormorants, Phalacrocoracidae. Proceedings of the XVI International Ornithological Congress: 121–124.

External links[edit]

Wikimedia Commons has media related to Phalacrocoracidae
Phalacrocoracidae
and Phalacrocorax.

Look up cormorant in Wiktionary, the free dictionary.

Cormorant
Cormorant
videos on the Internet Bird
Bird
Collection "Recovery plan for Chatham Island shag and Pitt Island shag 2001–2011" (PDF). Department of Conservation, Wellington, New Zealand. 2001. Retrieved 2007-09-28.  First video of cormorant deep sea dive, by the Wildlife Conservation Society and the National Research Council of Argentina. WCS press release, 2012-07-31

v t e

Order: Suliformes
Suliformes
(Phalacrocoraciformes)

Frigatebirds (family: Fregatidae · genus: Fregata)

Genus

Species

Fregata

Magnificent frigatebird
Magnificent frigatebird
(or Man O'War) Ascension frigatebird Christmas Island frigatebird Great frigatebird Lesser frigatebird

Gannets and boobies (family: Sulidae)

Genus

Species

Sula

Blue-footed booby Peruvian booby Masked booby Nazca booby Red-footed booby Brown booby Tasman booby

Papasula

Abbott's booby

Morus

Northern gannet Cape gannet Australasian gannet

Darters (family: Anhingidae · genus Anhinga)

Genus

Species

Anhinga

Anhinga Oriental darter African darter Australasian darter

Cormorants (family: Phalacrocoracidae
Phalacrocoracidae
· genus: Phalacrocorax)

Genus

Species

Phalacrocorax

Double-crested cormorant
Double-crested cormorant
(white-crested cormorant) Neotropic cormorant
Neotropic cormorant
(olivaceous cormorant) Little black cormorant Great cormorant White-breasted cormorant Indian cormorant Cape cormorant Socotra cormorant Bank cormorant
Bank cormorant
(Wahlberg's cormorant) Japanese cormorant
Japanese cormorant
(Temminck's cormorant) European shag Brandt's cormorant Spectacled cormorant Pelagic cormorant
Pelagic cormorant
(Baird's cormorant) Red-faced cormorant Rock shag Guanay cormorant Pied cormorant (yellow-faced cormorant) Black-faced cormorant Rough-faced shag
Rough-faced shag
(king shag) Foveaux shag Otago shag Chatham shag Auckland shag Campbell shag Bounty shag Imperial shag
Imperial shag
(blue-eyed shag) Heard Island shag Crozet shag Antarctic shag Kerguelen shag Macquarie shag South Georgia shag Red-legged cormorant Spotted shag Pitt shag
Pitt shag
(Featherstone's shag) Little pied cormorant Long-tailed cormorant Crowned cormorant Little cormorant Pygmy cormorant Flightless cormorant

v t e

Birds in culture

Activities

Aviculture Birdwatching Bird
Bird
conservation Fletching In sport

Cockfighting Falconry Pigeon racing Vinkensport

In science

Model organism Ornithology

In mythology and religion

Augury Sacred ibis Sky burial

In heraldry

Crow/Raven Eagle Martlet Turul

In hunting

Cormorant
Cormorant
fishing Driven grouse shooting Plume hunting Wildfowling

Products

Chicken Down Egg Feather Guano Poultry

In the arts

In art

Bird-and-flower painting Feather
Feather
tights Joan Miró

In poetry

The Conference of the Birds Ode to a Nightingale To a Skylark Crow

In prose

A History of British Birds The Tale of Jemima Puddle-Duck The Ugly Duckling

In theatre and ballet

The Birds Swan Lake The Firebird

In film

The Birds Kes Animated films Chicken
Chicken
films Horror films

in music In fashion

Aigrette Feather
Feather
boa Feather
Feather
cloak

In dance

Cendrawasih

Species

Chicken Cormorant Crow Cuckoo Golden eagle Goldfinch Kingfisher Parrot Partridge Peacock Penguin Pheasant Pigeon/Dove Raven

of the Tower of London

Sparrow Swallow

People

Illustrators

John James Audubon
John James Audubon
(The Birds of America) Thomas Bewick John Gould Lars Jonsson John Gerrard Keulemans Edward Lear Richard Lewington Roger Tory Peterson Henry Constantine Richter Joseph Smit Archibald Thorburn Joseph Wolf

Conservationists

Niels Krabbe Peter Scott

Organisations

BirdLife International Royal Society for the Protection of Birds Wildfowl & Wetlands Trust

Related

Category:Birds and humans Dinosaurs in culture Living things in culture

Fish in culture Insects in culture Mammals in culture Reptiles in culture

Zoomusicology

Taxon identifiers

Wd: Q3901247 ADW: Phalacrocoracidae EoL: 7996 EPPO: 1PHLKF Fauna Europaea: 10733 Fossilworks: 39688 GBIF: 5268 ITIS: 174713 NCBI:

.