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Swans are birds of the family Anatidae within the genus ''Cygnus''. The swans' closest relatives include the geese and ducks. Swans are grouped with the closely related geese in the
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
Anserinae The Anserinae are a subfamily in the waterfowl family Anatidae. It includes the swans and true geese. Under alternative systematical concepts (see e.g., Terres & NAS, 1991), it is split into two subfamilies, the Anserinae contain the geese and ...
where they form the tribe Cygnini. Sometimes, they are considered a distinct subfamily, Cygninae. There are six living and many
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
species of swan; in addition, there is a species known as the coscoroba swan which is no longer considered one of the true swans. Swans usually
mate for life In biology, a pair bond is the strong affinity that develops in some species between a mating pair, often leading to the production and rearing of offspring and potentially a lifelong bond. Pair-bonding is a term coined in the 1940s that is freque ...
, although "divorce" sometimes occurs, particularly following nesting failure, and if a mate dies, the remaining swan will take up with another. The number of eggs in each
clutch A clutch is a mechanical device that engages and disengages power transmission, especially from a drive shaft to a driven shaft. In the simplest application, clutches connect and disconnect two rotating shafts (drive shafts or line shafts). ...
ranges from three to eight.


Etymology and terminology

The English word ''swan'', akin to the
German German(s) may refer to: * Germany (of or related to) **Germania (historical use) * Germans, citizens of Germany, people of German ancestry, or native speakers of the German language ** For citizens of Germany, see also German nationality law **Ger ...
, Dutch and
Swedish Swedish or ' may refer to: Anything from or related to Sweden, a country in Northern Europe. Or, specifically: * Swedish language, a North Germanic language spoken primarily in Sweden and Finland ** Swedish alphabet, the official alphabet used by ...
, is derived from
Indo-European root The roots of the reconstructed Proto-Indo-European language (PIE) are basic parts of words that carry a lexical meaning, so-called morphemes. PIE roots usually have verbal meaning like "to eat" or "to run". Roots never occurred alone in the lang ...
' ('to sound, to sing'). Young swans are known as '' cygnets'' or as '' swanlings''; the former derives via Old French or (diminutive suffix et 'little') from the Latin word , a variant form of 'swan', itself from the Greek , a word of the same meaning. An adult male is a ''cob'', from Middle English (leader of a group); an adult female is a ''pen''.


Description

Swans are the largest extant members of the waterfowl family Anatidae, and are among the largest flying birds. The largest living species, including the
mute swan The mute swan (''Cygnus olor'') is a species of swan and a member of the waterfowl family Anatidae. It is native to much of Eurosiberia, and (as a rare winter visitor) the far north of Africa. It is an introduced species in North America, home ...
, trumpeter swan, and whooper swan, can reach a length of over and weigh over . Their wingspans can be over . Compared to the closely related geese, they are much larger and have proportionally larger feet and necks. Adults also have a patch of unfeathered skin between the eyes and bill. The sexes are alike in
plumage Plumage ( "feather") is a layer of feathers that covers a bird and the pattern, colour, and arrangement of those feathers. The pattern and colours of plumage differ between species and subspecies and may vary with age classes. Within species, ...
, but males are generally bigger and heavier than females. The biggest species of swan ever was the extinct ''
Cygnus falconeri ''Cygnus falconeri'', the giant swan, ( Maltese: ) is an extinct, very large swan known from Middle Pleistocene-aged deposits from Malta and Sicily. Its dimensions are described as exceeding those of the living mute swan by one-third, which woul ...
'', a flightless giant swan known from fossils found on the Mediterranean islands of Malta and Sicily. Its disappearance is thought to have resulted from extreme climate fluctuations or the arrival of superior predators and competitors. The Northern Hemisphere species of swan have pure white plumage, but the Southern Hemisphere species are mixed black and white. The Australian
black swan The black swan (''Cygnus atratus'') is a large waterbird, a species of swan which breeds mainly in the southeast and southwest regions of Australia. Within Australia, the black swan is nomadic, with erratic migration patterns dependent upon c ...
(''Cygnus atratus'') is completely black except for the white flight feathers on its wings; the chicks of black swans are light grey. The South American black-necked swan has a white body with a black neck. Swans' legs are normally a dark blackish grey colour, except for the South American black-necked swan, which has pink legs. Bill colour varies: the four
subarctic The subarctic zone is a region in the Northern Hemisphere immediately south of the true Arctic, north of humid continental regions and covering much of Alaska, Canada, Iceland, the north of Scandinavia, Siberia, and the Cairngorms. Generally, ...
species have black bills with varying amounts of yellow, and all the others are patterned red and black. Although birds do not have teeth, swans, like other Anatidae, have beaks with serrated edges that look like small jagged 'teeth' as part of their beaks used for catching and eating aquatic plants and algae, but also molluscs, small fish, frogs, and worms. In the mute swan and black-necked swan, both sexes have a fleshy lump at the base of their bills on the upper mandible, known as knob, which is larger in males, and is condition dependent, changing seasonally.


Distribution and movements

Swans are generally found in temperate environments, rarely occurring in the tropics. A group of swans is called a bevy or a wedge in flight. Four (or five) species occur in the
Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the Equator. For other planets in the Solar System, north is defined as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...
, one species is found in Australia, one
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
species was found in New Zealand and the Chatham Islands, and one species is distributed in southern South America. They are absent from tropical Asia, Central America, northern South America and the entirety of Africa. One species, the mute swan, has been introduced to North America, Australia and New Zealand. Several species are migratory, either wholly or partly so. The mute swan is a partial migrant, being resident over areas of Western Europe but wholly migratory in Eastern Europe and Asia. The whooper swan and tundra swan are wholly migratory, and the trumpeter swans are almost entirely migratory. There is some evidence that the black-necked swan is migratory over part of its range, but detailed studies have not established whether these movements are long or short range migration.


Behaviour

Swans feed in water and on land. They are almost entirely herbivorous, although they may eat small amounts of aquatic animals. In the water, food is obtained by up-ending or dabbling, and their diet is composed of the roots, tubers, stems and leaves of aquatic and submerged plants. Swans famously mate for life, and typically bond even before they reach sexual maturity. Trumpeter swans, for example, who can live as long as 24 years and only start breeding at the age of 4–7, form monogamous pair bonds as early as 20 months. "Divorce", though rare, does occur; one study of mute swans showing a 3% rate for pairs that breed successfully and 9% for pairs that do not. The pair bonds are maintained year-round, even in gregarious and migratory species like the tundra swan, which congregate in large flocks in the wintering grounds. Swans' nests are on the ground near water and about a metre across. Unlike many other ducks and geese, the male helps with the nest construction, and will also take turns incubating the eggs. Alongside the
whistling duck The whistling ducks or tree ducks are a subfamily, Dendrocygninae, of the duck, goose and swan family of birds, Anatidae. In other taxonomic schemes, they are considered a separate family, Dendrocygnidae. Some taxonomists list only one genu ...
s, swans are the only anatids that will do this. Average egg size (for the mute swan) is 113×74 mm, weighing 340 g, in a clutch size of 4 to 7, and an incubation period of 34–45 days."Mute Swan"
British Trust for Ornithology
Swans are highly protective of their nests. They will viciously attack anything that they perceive as a threat to their chicks, including humans. One man was suspected to have drowned in such an attack. Swans' intraspecific aggressive behaviour is shown more frequent than interspecific behaviour for food and shelter. The aggression with other species is shown more in Bewick's swans.


Systematics and evolution

Evidence suggests that the genus ''Cygnus'' evolved in Europe or western Eurasia during the Miocene, spreading all over the Northern Hemisphere until the Pliocene. When the southern species branched off is not known. The mute swan apparently is closest to the Southern Hemisphere ''Cygnus''; its habits of carrying the neck curved (not straight) and the wings fluffed (not flush) as well as its bill color and knob indicate that its closest living relative is the black swan. Given the
biogeography Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, ...
and appearance of the
subgenus In biology, a subgenus (plural: subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between t ...
''Olor'' it seems likely that these are of a more recent origin, as evidence shows by their modern ranges (which were mostly uninhabitable during the last ice age) and great similarity between the taxa.


Phylogeny


Species

Genus ''Cygnus'' The coscoroba swan (''Coscoroba coscoroba'') from South America, the only species in its genus, is apparently not a true swan. Its phylogenetic position is not fully resolved; it is in some aspects more similar to geese and shelducks.


Fossil record

The fossil record of the genus ''Cygnus'' is quite impressive, although allocation to the subgenera is often tentative; as indicated above, at least the early forms probably belong to the ''C. olor'' – Southern Hemisphere lineage, whereas the Pleistocene taxa from North America would be placed in ''Olor''. A number of prehistoric species have been described, mostly from the Northern Hemisphere. In the Mediterranean, the leg bones of the giant swan ('' C. falconeri'') were found on the islands of Malta and Sicily; it may have been over 2 meters from tail to bill, which was taller (though not heavier) than the contemporary local dwarf elephants (''
Palaeoloxodon falconeri ''Palaeoloxodon falconeri'', also known as the pygmy elephant, Maltese pygmy elephant, or Sicilian dwarf elephant, is an extinct Siculo- Maltese species of elephant that was derived from the straight-tusked elephant. It is amongst the smallest of ...
''). * Subgenus ''Chenopis'' **†
New Zealand swan The New Zealand swan ( Moriori: ''poūwa'', ''Cygnus sumnerensis'') is an extinct indigenous swan from the Chatham Islands and the South Island of New Zealand. Discovered as archaeological remains in 1889, it was originally considered a separate ...
, ''Cygnus sumnerensis'', an extinct species related to the black swan of Australia * Other subgenera (see above): **†''Cygnus csakvarensis'' Lambrecht 1933 [''Cygnus csákvárensis'' Lambrecht 1931a nomen nudum; ''Cygnanser csakvarensis'' (Lambrecht 1933) Kretzoi 1957; ''Olor csakvarensis'' (Lambrecht 1933) Mlíkovský 1992b] (Late Miocene of Hungary) **†''Cygnus mariae'' Bickart 1990 (Early Pliocene of Wickieup, U.S.) **†''Cygnus verae'' Boev 2000 (Early Pliocene of Sofia, Bulgaria) **†''Cygnus liskunae'' (Kuročkin 1976) 'Anser liskunae'' Kuročkin 1976(Middle Pliocene of western Mongolia) **†''Cygnus hibbardi'' Brodkorb 1958 (?Early Pleistocene of Idaho, U.S.) **†''Cygnus'' sp. Louchart ''et al''. 1998 (Early Pleistocene of Dursunlu, Turkey) **†
Giant swan ''Cygnus falconeri'', the giant swan, ( Maltese: ) is an extinct, very large swan known from Middle Pleistocene-aged deposits from Malta and Sicily. Its dimensions are described as exceeding those of the living mute swan by one-third, which woul ...
(''Cygnus falconeri'') Parker 1865 sensu Livezey 1997a 'Cygnus melitensis'' Falconer 1868; ''Palaeocygnus falconeri'' (Parker 1865) Oberholser 1908(Middle Pleistocene of Malta and Sicily, Mediterranean) **†''
Cygnus paloregonus ''Cygnus paloregonus'' is a fossil swan. It is an ancestor of, and distantly allied to, the mute swan. It is known from the Pleistocene from Fossil Lake, Oregon, Froman's Ferry, Idaho, and from Arizona. It is referred to by Hildegarde Howard in D ...
'' Cope 1878 'Anser condoni'' Schufeldt 1892; ''Cygnus matthewi'' Schufeldt 1913(Middle Pleistocene of west-central U.S.) **†
Dwarf swan Dwarf or dwarves may refer to: Common uses *Dwarf (folklore), a being from Germanic mythology and folklore * Dwarf, a person or animal with dwarfism Arts, entertainment, and media Fictional entities * Dwarf (''Dungeons & Dragons''), a humanoid ...
(''Cygnus equitum'') Bate 1916 sensu Livezey 1997 'Anser equitum'' (Bate 1916) Brodkorb 1964; ''Cygnus'' (''Olor'') ''equitum'' Bate 1916 sensu Northcote 1988a(Middle – Late Pleistocene of Malta and Sicily, Mediterranean) **†''Cygnus lacustris'' (De Vis 1905) 'Archaeocycnus lacustris'' De Vis 1905(Late Pleistocene of the Lake Eyre region, Australia) **†''Cygnus'' sp. (Pleistocene of Australia) **†''Cygnus atavus'' (Fraas 1870) Mlíkovský 1992 [''Anas atava'' Fraas 1870; ''Anas cygniformis'' Fraas 1870; ''Palaelodus steinheimensis'' Fraas 1870; ''Anser atavus'' (Fraas 1870) Lambrecht 1933; ''Anser cygniformis'' (Fraas 1870) Lambrecht 1933] * Other genera **† ''Annakacygna'' The supposed fossil swans ''"Cygnus" bilinicus'' and ''"Cygnus" herrenthalsi'' were, respectively, a stork and some large bird of unknown affinity (due to the bad state of preservation of the referred material).


In culture


European motifs

Many of the cultural aspects refer to the mute swan of Europe. Perhaps the best known story about a swan is the fable " The Ugly Duckling". Swans are often a symbol of love or fidelity because of their long-lasting, apparently monogamous relationships. See
Wagner's Wilhelm Richard Wagner ( ; ; 22 May 181313 February 1883) was a German composer, theatre director, polemicist, and conductor who is chiefly known for his operas (or, as some of his mature works were later known, "music dramas"). Unlike most op ...
famous swan-related operas '' Lohengrin'' and '' Parsifal''.


As food

Swan meat was regarded as a luxury food in England in the reign of Elizabeth I. A recipe for baked swan survives from that time: "To bake a Swan Scald it and take out the bones, and parboil it, then season it very well with Pepper, Salt and Ginger, then lard it, and put it in a deep Coffin of Rye Paste with store of Butter, close it and bake it very well, and when it is baked, fill up the Vent-hole with melted Butter, and so keep it; serve it in as you do the Beef-Pie." The '' Illustrious Brotherhood of Our Blessed Lady'', a religious confraternity which existed in
's-Hertogenbosch s-Hertogenbosch (), colloquially known as Den Bosch (), is a city and municipality in the Netherlands with a population of 157,486. It is the capital of the province of North Brabant and its fourth largest by population. The city is south of th ...
in the late Middle Ages, had 'sworn members', also called 'swan-brethren' because they used to donate a swan for the yearly banquet.


Heraldics


Ancient Greece and Rome

Swans feature strongly in mythology. In Greek mythology, the story of
Leda and the Swan Leda and the Swan is a story and subject in art from Greek mythology in which the god Zeus, in the form of a swan, seduces or rapes Leda. According to later Greek mythology, Leda bore Helen and Polydeuces, children of Zeus, while at the sa ...
recounts that
Helen of Troy Helen of Troy, Helen, Helena, (Ancient Greek: Ἑλένη ''Helénē'', ) also known as beautiful Helen, Helen of Argos, or Helen of Sparta, was a figure in Greek mythology said to have been the most beautiful woman in the world. She was believe ...
was conceived in a union of Zeus disguised as a swan and
Leda Leda may refer to: Mythology * Leda (mythology), queen of Sparta and mother of Helen of Troy in Greek mythology Places * Leda, Western Australia, a suburb of Perth, Western Australia * Leda makeshift settlement, Bangladesh, a refugee camp ...
, Queen of Sparta. Other references in classical literature include the belief that, upon death, the mute swan would sing beautifully—hence the phrase
swan song The swan song ( grc, κύκνειον ᾆσμα; la, carmen cygni) is a metaphorical phrase for a final gesture, effort, or performance given just before death or retirement. The phrase refers to an ancient belief that swans sing a beautiful so ...
. The mute swan is also one of the sacred birds of Apollo, whose associations stem both from the nature of the bird as a symbol of light, as well as the notion of a "swan song". The god is often depicted riding a chariot pulled by or composed of swans in his ascension from
Delos The island of Delos (; el, Δήλος ; Attic: , Doric: ), near Mykonos, near the centre of the Cyclades archipelago, is one of the most important mythological, historical, and archaeological sites in Greece. The excavations in the island are ...
. In the second century, the Roman poet Juvenal made a sarcastic reference to a good woman being a "rare bird, as rare on earth as a black swan" (black swans being completely unknown in the Northern Hemisphere until Dutch explorers reached Australia in the 1600s), from which comes the Latin phrase ' (rare bird).


‘Black Swan’ event

The Black Swan theory originates from Juvenal’s reference, leading to the black swan as a metaphor for something that could, in theory, exist, but does not. After the "discovery" of actual black swans, this became a metaphor or analogy for something, typically an unexpected event or
outlier In statistics, an outlier is a data point that differs significantly from other observations. An outlier may be due to a variability in the measurement, an indication of novel data, or it may be the result of experimental error; the latter are ...
, that has an unforeseen significance.


Irish lore and poetry

The Irish legend of the
Children of Lir The ''Children of Lir'' ( ga, Oidheadh chloinne Lir) is a legend from Irish mythology. It is a tale from the post-Christianisation period that mixes magical elements such as druidic wands and spells with a Christian message of Christian faith ...
is about a stepmother transforming her children into swans for 900 years. In the legend ''
The Wooing of Etain ''The'' () is a grammatical article in English, denoting persons or things already mentioned, under discussion, implied or otherwise presumed familiar to listeners, readers, or speakers. It is the definite article in English. ''The'' is the m ...
'', the king of the Sidhe (subterranean-dwelling, supernatural beings) transforms himself and the most beautiful woman in Ireland, Etain, into swans to escape from the king of Ireland and Ireland's armies. The swan has recently been depicted on an Irish commemorative coin. Swans are also present in Irish literature in the poetry of
W.B. Yeats William Butler Yeats (13 June 186528 January 1939) was an Irish poet, dramatist, writer and one of the foremost figures of 20th-century literature. He was a driving force behind the Irish Literary Revival and became a pillar of the Irish liter ...
. "The Wild Swans at Coole" has a heavy focus on the mesmerising characteristics of the swan. Yeats also recounts the myth of Leda and the Swan in the poem of the same name.


Nordic lore

In
Norse mythology Norse, Nordic, or Scandinavian mythology is the body of myths belonging to the North Germanic peoples, stemming from Old Norse religion and continuing after the Christianization of Scandinavia, and into the Nordic folklore of the modern period ...
, there are two swans that drink from the sacred
Well of Urd A well is an excavation or structure created in the ground by digging, driving, or drilling to access liquid resources, usually water. The oldest and most common kind of well is a water well, to access groundwater in underground aquifers. The ...
in the realm of Asgard, home of the gods. According to the Prose Edda, the water of this well is so pure and holy that all things that touch it turn white, including this original pair of swans and all others descended from them. The poem '' Volundarkvida'', or the ''Lay of Volund'', part of the Poetic Edda, also features swan maidens. In the Finnish epic '' Kalevala'', a swan lives in the Tuoni river located in Tuonela, the underworld realm of the dead. According to the story, whoever killed a swan would perish as well. Jean Sibelius composed the '' Lemminkäinen Suite'' based on the ''Kalevala'', with the second piece entitled ''
Swan of Tuonela ''The Swan of Tuonela'' (') is an 1895 tone poem by the Finnish composer Jean Sibelius. It is part of the '' (Four Legends from the Kalevala)'', Op. 22, based on the Finnish mythological epic the ''Kalevala''. ''The Swan of Tuonela'' was ori ...
'' ''(Tuonelan joutsen)''. Today, five flying swans are the symbol of the
Nordic countries The Nordic countries (also known as the Nordics or ''Norden''; literal translation, lit. 'the North') are a geographical and cultural region in Northern Europe and the Atlantic Ocean, North Atlantic. It includes the sovereign states of Denmar ...
; the whooper swan (''Cygnus cygnus'') is the national bird of Finland; and the mute swan is the national bird of Denmark.


''Swan Lake'' ballet

The ballet ''
Swan Lake ''Swan Lake'' ( rus, Лебеди́ное о́зеро, r=Lebedínoye ózero, p=lʲɪbʲɪˈdʲinəjə ˈozʲɪrə, link=no ), Op. 20, is a ballet composed by Russian composer Pyotr Ilyich Tchaikovsky in 1875–76. Despite its initial failur ...
'' is among the most canonic of classical ballets. Based on the 1875–76 score by Pyotr Ilyich Tchaikovsky, the most promulgated choreographic version was created by
Marius Petipa Marius Ivanovich Petipa (russian: Мариус Иванович Петипа), born Victor Marius Alphonse Petipa (11 March 1818), was a French ballet dancer, pedagogue and choreographer. Petipa is one of the most influential ballet masters an ...
and Lev Ivanov (1895), the premiere of which was danced by the Imperial Ballet at the Mariinsky Theater in St. Petersburg. The ballet's lead dual roles of Odette (white swan) / Odile (black swan) represent good and evil, and are among the most challenging roles created in Romantic classical ballet. The ballet is in the repertories of ballet companies around the world.


Christianity

A swan is one of the attributes of St. Hugh of Lincoln, based on the story of a swan who was devoted to him.


Spanish language literature

In Latin American literature, the Nicaraguan poet Rubén Darío (1867–1916) consecrated the swan as a symbol of artistic inspiration by drawing attention to the constancy of swan imagery in Western culture, beginning with the rape of Leda and ending with Wagner's ''Lohengrin''. Darío's most famous poem in this regard is ''Blasón – "Coat of Arms"'' (1896), and his use of the swan made it a symbol for the Modernismo poetic movement that dominated Spanish language poetry from the 1880s until the First World War. Such was the dominance of Modernismo in Spanish language poetry that the Mexican poet
Enrique González Martínez Enrique González Martínez (April 13, 1871 in Guadalajara, Jalisco – February 19, 1952 in Mexico City) was a Mexican poet, diplomat, surgeon and obstetrician. His poetry is considered to be primarily Modernist in nature, with elements of Fre ...
attempted to announce the end of Modernismo with a
sonnet A sonnet is a poetic form that originated in the poetry composed at the Court of the Holy Roman Emperor Frederick II in the Sicilian city of Palermo. The 13th-century poet and notary Giacomo da Lentini is credited with the sonnet's invention, ...
provocatively entitled, '' Tuércele el cuello al cisne – "Wring the Swan's Neck"'' (1910).


Hinduism

Swans are revered in Hinduism, and are compared to saintly persons whose chief characteristic is to be in the world without getting attached to it, just as a swan's feather does not get wet although it is in water. The Sanskrit word for swan is ''hamsa'' and the "Raja Hamsam" or the Royal Swan is the vehicle of Devi Saraswati, which symbolises the ''Sattva Guna'' or purity par excellence. The swan, if offered a mixture of milk and water, is said to be able to drink the milk alone. Therefore, Saraswati, the goddess of knowledge, is seen riding the swan because the swan thus symbolizes ''Viveka'', i.e. prudence and discrimination between the good and the bad or between the eternal and the transient. This is seen as a great quality, as shown by this Sanskrit verse: It is mentioned several times in the Vedic literature, and persons who have attained great spiritual capabilities are sometimes called Paramahamsa ("Supreme Swan") on account of their spiritual grace and ability to travel between various spiritual worlds. In the Vedas, swans are said to reside in the summer on Lake Manasarovar and migrate to Indian lakes for the winter. They are believed to possess some powers, such as the ability to eat pearls.


Indo-European religions

Swans are intimately associated with the divine twins in Indo-European religions, and it is thought that in Proto-Indo-European times, swans were a solar symbol associated with the divine twins and the original Indo-European sun goddess.


See also

*
Royal Swans The Royal Swans are a Flock (birds), flock of swans of two species—the mute swan (''Cygnus olor'') and the black swan (''C. atratus'')—the original six pairs of which were a gift to the city of Ottawa from Queen Elizabeth II in 1967, to commemor ...


References


External links

* * Louchart, Antoine; Mourer-Chauviré, Cécile; Guleç, Erksin; Howell, Francis Clark & White, Tim D. (1998): L'avifaune de Dursunlu, Turquie, Pléistocène inférieur: climat, environnement et biogéographie. '' C. R. Acad. Sci. Paris IIA'' 327(5): 341–346. rench with English abridged version
A History of British Birds
* * {{Authority control Anserinae Extant Miocene first appearances Taxa named by François Alexandre Pierre de Garsault