Phalacrocoracidae is a family of approximately 40 species of aquatic birds found worldwide and commonly known as cormorants and shags. Several different classifications of the family have been proposed, but in 2021 the International Ornithologists' Union (IOU) adopted a consensus taxonomy of seven genera. The common names "cormorant" and "shag" have been assigned to different species in the family somewhat haphazardly.

Cormorants and shags are medium-to-large birds, with body weight in the range of and wing span of . 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 . Cormorants and shags have relatively short wings due to their need for economical movement underwater, and consequently have among the highest flight costs of any flying bird.

Cormorant colonies nest around seacoasts, in trees, islets or cliffs. They are coastal rather than oceanic birds. Some species have colonised inland waters. Species may be found in all areas of the world except for the central Pacific islands.

Cormorants may descend from a freshwater ancestor in south Asia whose offspring adapted in time to new environments and spread across the Eurasian landmass and the world.

Names

"Cormorant" is a contraction probably derived from Latin corvus marinus, "sea raven"; in the early 19th century, the similarly derived spelling "corvorant" was sometimes used. commentary, and existing reference video). Imperial shags fitted with miniaturized video recorders have been filmed diving to depths of as much as to forage on the sea floor.

thumb|Wing-drying behaviour in a little cormorant

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 state that cormorants have waterproof feathers while others say that they have water-permeable feathers. Still others suggest that the outer plumage absorbs water but does not permit it to penetrate the layer of air next to the skin. The wing drying action is seen even in the flightless cormorant but not in the Antarctic shags or red-legged cormorants. Alternate functions suggested for the spread-wing posture include that it aids thermoregulation or digestion, balances the bird, or indicates presence of fish. A detailed study of the great cormorant concluded there is little doubt that it serves to dry the plumage.

thumb|A Cormorant eating a fish at [[Taudaha Lake]]

Cormorants are colonial nesters, using trees, rocky islets, or cliffs. The eggs are a chalky-blue colour. There is usually one brood a year. Parents regurgitate food to feed their young.

Taxonomy

The genus Phalacrocorax, from which the family name Phalacrocoracidae is derived, is Latinised from Ancient Greek phalakros "bald" and korax "raven". This is thought to refer to the ornamental white head plumes prominent in Mediterranean birds of this species, or the creamy white patch on the cheeks of adult great cormorants, but is certainly not a unifying characteristic of cormorants.

The cormorant family was traditionally placed within the Pelecaniformes or, in the Sibley–Ahlquist taxonomy of the 1990s, the expanded Ciconiiformes. Pelecaniformes in the traditional sense—all waterbird groups with totipalmate foot webbing—are not a monophyletic group, even after the removal of the distantly-related tropicbirds. 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 (gannets and boobies), and perhaps the pelicans or even penguins, than to all other living birds.

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. In 2014, a landmark study proposed a 7 genera treatment, which was adopted by the IUCN Red List and BirdLife International, and later by the IOC in 2021, standardizing it. This bone and the highly developed muscles over it, the M. adductor mandibulae caput nuchale, are unique to the families Phalacrocoracidae and Anhingidae.

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 does have some degree of merit. The resolution provided by the mtDNA 12S rRNA and ATPase subunits six and eight sequence data this is now followed by most authorities, including the IOC World Bird List.

{| class="wikitable"

|-

! Image !! Genus !! Species

|-

|175px || Microcarbo ||

  • Crowned cormorant Microcarbo coronatus
  • Little cormorant Microcarbo niger
  • Little pied cormorant Microcarbo melanoleucos
  • Pygmy cormorant Microcarbo pygmeus
  • Reed cormorant or long-tailed cormorant Microcarbo africanus
  • †Serventys' cormorant Microcarbo serventyorum

Around Indian Ocean, one species extending from Central Asia into Europe. Mostly in freshwater habitat. Small (about 50–60 cm long), nondescript black to dark brown (except for one species with white underparts).

|-

|175px || Poikilocarbo ||

  • Red-legged cormorant Poikilocarbo gaimardi

Subtropical to subantarctic Pacific South America, ranging a bit into the southwestern Atlantic. Maritime. Mid-sized (around 75 cm), grey with scalloped wings and contrasting white/yellow/red neck mark and bare parts. Its high-pitched chirping calls are quite unlike those of other cormorants.

|-

|175px || Urile ||

  • Brandt's cormorant Urile penicillatus
  • Red-faced cormorant Urile urile
  • Pelagic cormorant Urile pelagicus
  • Spectacled cormorant †Urile perspicillatus

Northern Pacific, one species extending into subtropical waters on the American West Coast. Maritime. Smallish to large (65–100 cm), generally black with metallic sheen (usually blue/green), in breeding plumage with bright bare facial skin in the eye region and two crests (crown and nape).

|-

|175px || Phalacrocorax ||

  • Bank cormorant Phalacrocorax neglectus
  • Socotra cormorant Phalacrocorax nigrogularis
  • Pitt shag Phalacrocorax featherstoni
  • Spotted shag Phalacrocorax punctatus
  • Black-faced cormorant Phalacrocorax fuscescens
  • Australian pied cormorant Phalacrocorax varius
  • Little black cormorant Phalacrocorax sulcirostris
  • Indian cormorant Phalacrocorax fuscicollis
  • Cape cormorant Phalacrocorax capensis
  • Japanese cormorant or Temminck's cormorant Phalacrocorax capillatus
  • Great cormorant (including white-breasted cormorant and black shag) Phalacrocorax carbo

Mostly around the Indian Ocean, one species group extending throughout Eurasia and to Atlantic North America. Maritime to freshwater. Size very variable (60–100 cm), blackish with metallic sheen (usually bronze to purple) and/or white cheek and thigh patches or underside at least in breeding plumage; usually a patch of bare yellow skin at the base of the bill.

|-

|175px || Gulosus ||

  • European shag Gulosus aristotelis

Breeds in the European Arctic, winters in Europe and North Africa. Maritime. Mid-sized (70–80 cm), glossy black, in breeding plumage with a forehead crest curled to the front.

|-

|175px || Nannopterum ||

  • Flightless cormorant Nannopterum harrisi
  • Neotropic cormorant Nannopterum brasilianum
  • Double-crested cormorant Nannopterum auritum

Throughout the Americas. Mostly freshwater. Smallish to large (65–100 cm), nondescript brownish-black. One species with white tufts on sides of head in breeding plumage.

|-

|175px || Leucocarbo ||

  • Rock shag or Magellanic cormorant Leucocarbo magellanicus
  • Guanay cormorant Leucocarbo bougainvillii
  • Bounty shag Leucocarbo ranfurlyi
  • New Zealand king shag or rough-faced shag Leucocarbo carunculatus
  • Chatham shag Leucocarbo onslowi
  • Stewart Island shag (including Foveaux shag L. stewarti) Leucocarbo chalconotus
  • Auckland shag Leucocarbo colensoi
  • Campbell shag Leucocarbo campbelli
  • Imperial shag or blue-eyed shag Leucocarbo atriceps
  • South Georgia shag Leucocarbo georgianus
  • Crozet shag Leucocarbo melanogenis
  • Antarctic shag Leucocarbo bransfieldensis
  • Kerguelen shag Leucocarbo verrucosus
  • Heard Island shag Leucocarbo nivalis
  • Macquarie shag Leucocarbo purpurascens

Generally Subantarctic, but extending farther north in South America; many oceanic-island endemics. Maritime. Smallish to largish (65–80 cm), typically black above and white below, and with bare yellow or red skin in the facial region. A southern circumpolar group of several species (the blue-eyed shag complex) is characterised by bright blue orbital skin.

|-

|}

Prior to 2021, the IOU (or formerly the IOC) classified all these species in just three genera: Microcarbo, Leucocarbo, and a broad Phalacrocorax containing all remaining species; however, this treatment rendered Phalacrocorax deeply paraphyletic with respect to Leucocarbo. Other authorities, such as the Clements Checklist, formerly recognised only Microcarbo as a separate genus from Phalacrocorax.

Evolution and fossil record

The details of the evolution of the cormorants 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 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:<br />

A scapula from the Campanian-Maastrichtian boundary, about 70 mya (million years ago), was found in the Nemegt Formation in Mongolia; it is now in the PIN collection. It is from a bird roughly the size of a spectacled cormorant, and quite similar to the corresponding bone in Phalacrocorax. A Maastrichtian (Late Cretaceous, c. 66 mya) right femur, AMNH FR 25272 from the 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. However, cormorants likely originated much later, and these are likely misidentifications.

As the Early Oligocene "Sula" ronzoni cannot be assigned to any of the sulid families—cormorants and shags, darters, and gannets and boobies—with certainty, the best interpretation is that the Phalacrocoracidae diverged from their closest ancestors in the Early Oligocene, perhaps some 30 million years ago, and that the Cretaceous fossils represent ancestral sulids, "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.

Phylogenetic evidence indicates that the cormorants diverged from their closest relatives, the darters, during the Late Oligocene, indicating that most of the claims of Cretaceous or early Paleogene cormorant occurrences are likely misidentifications.

  • Phalacrocorax femoralis (Modelo Late Miocene/Early Pliocene of WC North America) &ndash; formerly Miocorax; Nannopterum?<!-- Auk52:75; Condor31:167; Condor58:367 -->
  • Phalacrocorax sp. (Late Miocene/Early Pliocene of Lee Creek Mine, US) &ndash; Nannopterum or Phalacrocorax?
  • Phalacrocorax sp. 1 (Late Miocene/Early Pliocene of WC South America) &ndash; probably Leucocarbo<!-- "Phalacrocorax aff.bougainvillii" -->
  • Phalacrocorax sp. 2 (Pisco Late Miocene/Early Pliocene of SW Peru) &ndash; Poikilocarbo or Leucocarbo?
  • Phalacrocorax longipes (Late Miocene – Early Pliocene of Ukraine) &ndash; formerly Pliocarbo; Microcarbo, Phalacrocorax or Gulosus?<!-- AnnNHMusWien104A:237 -->
  • Phalacrocorax goletensis (Early Pliocene – Early Pleistocene of Mexico) &ndash; Urile or Nannopterum, perhaps Poikilocarbo or Leuocarbo<!-- ContribMusPaleontolUnivMich24:205 -->
  • Phalacrocorax wetmorei (Bone Valley Early Pliocene of Florida) &ndash; Nannopterum or Phalacrocorax?<!-- Condor58:367; Condor72:293 -->
  • Phalacrocorax sp. (Bone Valley Early Pliocene of Polk County, Florida, US) &ndash; Nannopterum or Phalacrocorax?
  • Phalacrocorax leptopus (Juntura Early/Middle Pliocene of Juntura, Malheur County, Oregon, US) &ndash; Nannopterum?
  • Phalacrocorax reliquus (Middle Pliocene of Mongolia) &ndash; Microcarbo, Phalacrocorax or Gulosus?
  • Phalacrocorax idahensis (Middle Pliocene – Pleistocene of Idaho, US, and possibly Florida) &ndash; Nannopterum?<!-- Condor58:367; Condor72:293; ContribMusPaleontolUnivMich24:205 -->
  • Phalacrocorax destefanii<!-- destefani? --> (Late Pliocene of Italy) &ndash; formerly Paracorax; Microcarbo, Phalacrocorax or Gulosus?<!-- Condor58:367 -->
  • Phalacrocorax filyawi (Pinecrest Late Pliocene of Florida, US) &ndash; may be P. idahensis; Nannopterum or Phalacrocorax, perhaps Urile?
  • Phalacrocorax kennelli (San Diego Late Pliocene of California, US) &ndash; Urile or Nannopterum?<!-- Condor72:293 -->
  • Phalacrocorax kumeyaay (San Diego Late Pliocene of California, US) &ndash; Urile or Nannopterum?<!-- OccasionalPapersOfTheUniversityOfKansasMuseumOfNaturalHistory140:1. -->
  • Phalacrocorax macer (Late Pliocene of Idaho, US) &ndash; Nannopterum?<!-- Condor72:293 -->
  • Phalacrocorax mongoliensis (Late Pliocene of W Mongolia) &ndash; Microcarbo, Phalacrocorax or Gulosus?
  • Phalacrocorax sp. (La Portada Late Pliocene of N Chile) &ndash; may be same as Late Miocene/Early Pliocene "Phalacrocorax sp. 2"; Poikilocarbo or Leucocarbo?
  • Phalacrocorax rogersi (Late Pliocene – Early Pleistocene of California, US) &ndash; Urile or Nannopterum?<!-- Condor33:31; Condor34:118; Condor58:367; ContribMusPaleontolUnivMich24:205 -->
  • Phalacrocorax chapalensis (Late Pliocene/Early Pleistocene of Jalisco, Mexico) &ndash; Urile or Nannopterum, perhaps Poikilocarbo or Leucocarbo?<!-- *ContribMusPaleontolUnivMich24:205 -->
  • Phalacrocorax gregorii (Late Pleistocene of Australia) &ndash; possibly not a valid species; Microcarbo, Phalacrocorax or Leucocarbo?<!-- Condor58:367 -->
  • Phalacrocorax vetustus (Late Pleistocene of Australia) &ndash; formerly Australocorax, possibly not a valid species; Microcarbo, Phalacrocorax or Leucocarbo?<!-- Condor58:367 -->
  • Phalacrocorax sp. (Sarasota County, Florida, US) &ndash; may be P. filawyi/idahensis; Nannopterum or Phalacrocorax?<!-- Plio- or Plestocene: Forsberg Shell Pit, Leisey Shell Pit, Macasphalt Shell Pit or Richardson Road Shell Pit? -->

The former "Phalacrocorax" (or "Oligocorax") mediterraneus is now considered to belong to the bathornithid Paracrax antiqua. "P." subvolans was actually a darter (Anhinga).

In human culture

Cormorant culling

Cormorant fishing

thumb|right|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 practised 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. In Japan, cormorant fishing is called and is performed by a fisherman known as an usho. Traditional forms of ukai can be seen on the Nagara River in the city of Gifu, Gifu Prefecture, where cormorant fishing has continued uninterrupted for 1300 years, or in the city of Inuyama, Aichi. In Guilin, Guangxi, cormorants are famous for fishing on the shallow Li River. In Gifu, the Japanese cormorant (P. capillatus) is used; Chinese fishermen often employ great cormorants (P. carbo). In Europe, a similar practice was also used on Doiran Lake in the region of Macedonia. James VI and I appointed a keeper of cormorants, John Wood, and built ponds at Westminster to train the birds to fish.

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 practised as a cultural tradition.

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. McCullagh suggests that the cormorant, associated with cypress trees, might act in part as a symbol of death and decay in the text.

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. Juana Maria's story was fictionalized in the children's novel Island of the Blue Dolphins.

The bird has inspired numerous writers, including Amy Clampitt, who wrote a poem called "The 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&nbsp;... vermilion-strapped" and "big black feet" that she mentions.

A cormorant representing Blanche Ingram appears in the first of the fictional paintings by Jane in Charlotte Brontë's novel Jane Eyre:

<blockquote>One gleam of light lifted into relief a half-submerged mast, on which sat a cormorant, dark and large, with wings flecked with foam; its beak held a gold bracelet, set with gems, that I had touched with as brilliant tints as my palette could yield, and as glittering distinctness as my pencil could impart.</blockquote>

In the Sherlock Holmes story "The Adventure of the Veiled Lodger", Dr. Watson warns that if there are further attempts to get at and destroy his private notes regarding his time with Sherlock Holmes, "the whole story concerning the politician, the lighthouse, and the trained cormorant will be given to the public. There is at least one reader who will understand."

A cormorant is humorously mentioned as having had linseed oil rubbed into it by a wayward pupil during the "Growth and Learning" segment of the 1983 Monty Python film Monty Python's The Meaning of Life.

The cormorant served as the hood ornament for the Packard automobile brand.

Cormorants (and books about them written by a fictional ornithologist) are a recurring fascination of the protagonist in Jesse Ball's 2018 novel Census.

The Pokémon Cramorant, introduced in Pokémon Sword and Shield, closely resembles a cormorant in both design and name.

The cormorant was chosen as the emblem for the Ministry of Defence Joint Services Command and Staff College at Shrivenham. A bird famed for flight, sea fishing and land nesting was felt to be particularly appropriate for a college that unified leadership training and development for the Army, Navy and Royal Air Force.

After a member produced a mock magazine cover from a photograph of roosting cormorants, the bird became the unofficial mascot of the Pentax Discuss Mailing List with many posts dedicated to discussion of the photography of the species.

Stephen Gregory’s 1987 horror novel The Cormorant concerns a man who inherits both a cottage and a cormorant from his late uncle. The 1993 film adaptation stars Ralph Fiennes and is directed by Peter Markham.

The titular protagonist of Seth Dickinson’s hard fantasy series The Masquerade is The Traitor Baru Cormorant, who is named after the bird.

See also

  • Anhinga
  • Cormorant culling
  • Liver bird

References

Sources

  • Benson, Elizabeth (1972): The Mochica: A Culture of Peru. Praeger Press, New York.
  • Berrin, Katherine & Larco Museum (1997) The Spirit of Ancient Peru: Treasures from the Museo Arqueológico Rafael Larco Herrera. Thames and Hudson, New York.
  • 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.
  • IUCN (2007): 2007 IUCN Red List of Threatened Species . IUCN, Gland.
  • 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.
  • Robertson, Connie (1998): Book of Humorous Quotations. Wordsworth Editions.
  • 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.
  • Cormorant videos on the Internet Bird Collection
  • 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