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{{Short description|Species of bird}}
{{otheruse}}
{{About|the bird|other uses}}

{{speciesbox
{{Taxobox
| image = Falco rusticolus white cropped.jpg
| name = Gyrfalcon or Gerfalcon
| image = Falco rusticolus.jpg
| image2 = Falco rusticolus distr.png
| image_width = 220px
| image_caption = Brown morph
| status = LC
| status = LC
| status_system = iucn3.1
| status_system = IUCN3.1
| status_ref = <ref name="IUCN">{{cite iucn |author=BirdLife International |date=2021 |title=''Falco rusticolus'' |page=e.T22696500A206261845 |doi=10.2305/IUCN.UK.2021-3.RLTS.T22696500A206261845.en |access-date=15 January 2023}}</ref>
| regnum = [[Animal]]ia
| phylum = [[Chordata]]
| status2 = CITES_A1
| status2_system = CITES
| classis = [[Aves]]
| status2_ref = <ref>{{Cite web|title=Appendices {{!}} CITES|url=https://cites.org/eng/app/appendices.php|access-date=2022-01-14|website=cites.org}}</ref>
| ordo = [[Falconiformes]]
| genus = Falco
| familia = [[Falconidae]]
| genus = ''[[Falcon|Falco]]''
| parent = Falco (Hierofalco)
| species = rusticolus
| subgenus = (''[[Hierofalco]]'')
| authority = [[Carl Linnaeus|Linnaeus]], 1758
| species = '''''F. rusticolus'''''
| synonyms = {{collapsible list | bullets = true
| binomial = ''Falco rusticolus''
|''Falco arcticus'' <small>[[Carl Peter Holbøll|Holbøll]],&nbsp;1843</small>
| binomial_authority = [[Carolus Linnaeus|Linnaeus]], [[1758]]
|''Falco candicans'' <small>[[Johann Friedrich Gmelin|Gmelin]],&nbsp;1788</small>
| synonyms =
*''Falco candicans''<br /><small>[[Johann Friedrich Gmelin|Gmelin]], [[1788]]</small>
|''Falco gyrfalco'' [[Carl Linnaeus|Linnaeus,&nbsp;1758]]
*''Falco islandus''<br /><small>[[Morten Thrane Brünnich|Brünnich]], [[1764]]</small>
|''Falco islandus'' <small>[[Morten Thrane Brünnich|Brünnich]],&nbsp;1764</small>
*''Falco obsoletus''<br /><small>Gmelin, 1788</small>
|''Falco obsoletus'' <small>Gmelin,&nbsp;1788</small>
*''Falco rusticolus candicans''<br /><small>Gmelin, 1788</small>
|''Falco rusticolus candicans'' <small>Gmelin,&nbsp;1788</small>
*''Falco rusticolus grebnitzkii''<br /><small>([[Nikolai Alekseevich Severtzov|Severtzov]], [[1885]])</small>
|''Falco rusticolus grebnitzkii'' <small>([[Nikolai Severtzov|Severtzov]],&nbsp;1885)</small>
*''Falco rusticolus intermedius''<br /><small>[[Constantin Wilhelm Lambert Gloger|Gloger]], [[1834]]</small>
|''Falco rusticolus intermedius'' <small>[[Constantin Wilhelm Lambert Gloger|Gloger]],&nbsp;1834</small>
*''Falco rusticolus islandus''<br /><small>[[Morten Thrane Brünnich|Brünnich]], [[1764]]</small> (but see text)
|''Falco rusticolus islandus'' <small>[[Morten Thrane Brünnich|Brünnich]],&nbsp;1764</small> (but see text)
*''Falco rusticolus obsoletus''<br /><small>Gmelin, 1788</small>
|''Falco rusticolus obsoletus'' <small>Gmelin,&nbsp;1788</small>
*''Falco rusticolus rusticolus''<br /><small>Linnaeus, 1758</small>
|''Falco rusticolus rusticolus'' <small>Linnaeus,&nbsp;1758</small>
*''Falco swarthi''<br /><small>[[Loye H. Miller|L.H. Miller]], [[1927]]</small>
|''Falco swarthi'' <small>[[Loye H. Miller|L.H. Miller]],&nbsp;1927</small>
*''Hierofalco grebnitzkii''<br /><small>Severtzov, 1885)</small>
|''Hierofalco grebnitzkii'' <small>Severtzov,&nbsp;1885)</small>
*''Hierofalco islandus''<br /><small>(Brünnich, 1764)</small>
|''Hierofalco islandus'' <small>(Brünnich,&nbsp;1764)</small>
*''Hierofalco rusticolus''<br /><small>(Linnaeus, 1758)</small>
|''Hierofalco rusticolus'' <small>(Linnaeus,&nbsp;1758)</small>
*''Hierofalco rusticolus candicans''<br /><small>(Gmelin, 1788)</small>
|''Hierofalco rusticolus candicans'' <small>(Gmelin,&nbsp;1788)</small>
}}
}}
}}


The '''gyrfalcon''' ({{IPAc-en|ˈ|dʒ|ɜr|ˌ|f|ɔː|(|l|)|k|ən}} or {{IPAc-en|ˈ|dʒ|ɜr|ˌ|f|æ|l|k|ən}})<ref>{{Cite web|title=Definition of gyrfalcon |url=https://www.dictionary.com/browse/gyrfalcon|access-date=2022-02-14|website=www.dictionary.com|language=en}}</ref> ({{Italics correction|'''''Falco rusticolus'''''}}), also abbreviated as '''gyr''', is a [[bird of prey]] from the genus ''Falco'' ([[falcon]]s and [[kestrel]]s) and the largest species of the family [[Falconidae]].<ref name=AUDGYR>{{cite web|url=https://www.audubon.org/field-guide/bird/gyrfalcon|title=Gyrfalcon|publisher=Audubon Guide to North American Birds|access-date=2016-04-04|date=2014-11-13}}</ref> A high-latitude species, the gyrfalcon breeds on the [[Arctic]] coasts and [[tundra]], the islands of northern [[North America]] and the [[Eurosiberian]] region, where it is mainly a resident species. Some gyrfalcons [[bird migration|disperse]] more widely after the [[breeding season]] or in winter, and individual [[vagrancy (biology)|vagrancy]] can take birds for long distances. Its plumage varies with location, with birds being coloured from all-white to dark brown. These colour variations are called [[polymorphism (biology)|morphs]]. Like other falcons, it shows [[sexual dimorphism]], with the female much larger than the male.
The '''Gyrfalcon''' {{IPAEng|ˈdʒɜrˌfɔlkən}} or {{IPAEng|ˈdʒɜrˌfælkən}} ('''''Falco rusticolus'''''), also spelled '''Gyr Falcon''', sometimes '''Gerfalcon''', is the largest of all [[falcon]] species. The Gyrfalcon breeds on [[Arctic]] coasts and islands of North America, Europe and Asia. It is mainly resident, but some Gyrfalcons [[bird migration|disperse]] more widely after the breeding season, or in winter<ref>Individual vagrancy can take birds for long distances. There is a story in the ''Unauthorized Biography of the Spring and Autumn''{{Verify source|date=August 2007}}<!-- Connected to [[Spring and Autumn Annals]]? --> of a 海东青 ('' Hǎidōngqīng'': Gyrfalcon) that succumbed to an arrow wound in the garden of [[Chen Hui Gong]]{{Verify source|date=August 2007}}<!-- depending on who that is, redo distance calculations -->. [[Confucius]] recognized the arrow as one of the [[Sushen]], whose fine stone arrowheads were a famous item of trade and tribute (RAM 2006). Although the Sushen's precise homeland at that time remains unknown, it was in the [[Manchuria]] region, no less than c.600 and perhaps more than 1000 km from the [[Lu (state)|Lu]] capital of [[Qufu]].</ref>.


For centuries, the gyrfalcon has been valued as a [[falconry|hunting bird]]. Typical prey includes the [[ptarmigan]] and [[waterfowl]], which it may attack in flight; and it also hunts [[fish]] and small [[mammal]]s.
The bird's common name comes from [[French language|French]] ''gerfaucon'', and in [[mediaeval Latin]] is rendered as ''gyrofalco''. The first part of the word may come from [[Old High German]] ''gîr'' (cf. modern [[German language|German]] ''Geier''), "[[vulture]]", referring to its size compared to other [[falcon]]s, or the [[Latin]] ''gȳrus'' ("circle", "curved path") from the species' circling as it searches for prey, unlike the other falcons in its range<ref>The Gyrfalcon's names in other [[Germanic language]]s provide few clues to resolve this. In [[Scandinavia]]n languages, it is generally named after its use in [[falconry]], whereas the modern [[Dutch (language)|Dutch]] name ''giervalk'' is peculiarly ambiguous: ''Gier'' means "vulture", whereas ''gieren'' means changing the [[yaw angle]] to circle in the air.</ref>. The male gyrfalcon is called a '''gyrkin''' in [[falconry]].


==Taxonomy and etymology==
Its [[scientific name]] is composed of the Latin terms for a falcon, ''Falco'', and for someone who lives in the countryside, ''rusticolus''.
The gyrfalcon was [[Species description|formally described]] by Swedish naturalist [[Carl Linnaeus]] in 1758 in the [[10th edition of Systema Naturae|tenth edition]] of his ''[[Systema Naturae]]'' under its current [[binomial nomenclature|binomial name]] ''Falco rusticolus''.<ref>{{cite book | last=Linnaeus | first=C. | author-link=Carl Linnaeus | year=1758 | title= Systema Naturæ per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, Volume 1| volume=v.1 | edition=10th | page=88 | publisher=Holmiae:Laurentii Salvii | language=la | url=https://www.biodiversitylibrary.org/page/726995 }}</ref> The genus name is the Late Latin term for a falcon, {{lang|la|Falco}}, from {{lang|la|[[wikt:falx#Latin|falx]]}} a [[sickle]], referencing the talons of the bird.<ref name=SOED>{{cite book|title=Shorter Oxford English dictionary | year=2007 |publisher=Oxford University Press| location=Oxford| isbn=978-0199206872| pages=3804}}</ref> The species name is from the Latin {{lang|la|rusticolus}}, a countryside-dweller, from {{lang|la|[[wikt:rus#Latin|rus]]}}, "country" and {{lang|la|colere}}, "to dwell".<ref name= job>{{cite book | last= Jobling | first= James A | year= 2010| title= The Helm Dictionary of Scientific Bird Names | url= https://archive.org/details/Helm_Dictionary_of_Scientific_Bird_Names_by_James_A._Jobling | publisher = Christopher Helm | location = London | isbn = 978-1-4081-2501-4 |pages =[https://archive.org/details/Helm_Dictionary_of_Scientific_Bird_Names_by_James_A._Jobling/page/n266 266], 344}}</ref> The bird's common name comes from [[French language|French]] {{lang|fr|gerfaucon}}; in [[Medieval Latin]], it is {{lang|la|gyrofalco}}. The first part of the word may come from [[Old High German]] {{lang|goh|gîr}} ({{abbr|cf.|compare with}} modern [[German language|German]] {{lang|de|Geier}}; ultimately from [[Proto-Germanic]] {{lang|gem-x-proto|*girį̄}} ("greed")) for "[[vulture]]", referring to its size in comparison with other [[falcon]]s; or from the [[Latin]] {{lang|la|gȳrus}} for "circle" or "curved path", in turn from the [[Ancient Greek]] {{lang|grc|γῦρος}}, {{transl|grc|gûros}}, meaning "circle" – from the species' circling as it searches for prey, distinct from the hunting of other falcons in its range.<ref group="nb">In [[Scandinavia]]n languages, it is generally named after its use in [[falconry]], whereas the modern [[Dutch (language)|Dutch]] name {{lang|nl|giervalk}} is peculiarly ambiguous: {{lang|nl|gier}} means "vulture", whereas {{lang|nl|gieren}} means changing the [[yaw, pitch, and roll|yaw angle]] to circle in the air.</ref> The male gyrfalcon is called a ''gyrkin'' in falconry.


==Description==
==Description==
The gyrfalcon is the largest falcon in the world, being about the same size as the largest [[buteo]]s but probably slightly heavier. Males are {{convert|48|to|61|cm|in|abbr=off}} long, weigh {{convert|805|to|1350|g|lboz|frac=2|abbr=off}}, with average weights reported as {{convert|1130|or|1170|g|lboz|frac=2|abbr=on}} and have a wingspan from {{convert|110|to|130|cm|in|abbr=on}}. Females are bulkier and larger, at {{convert|51|to|65|cm|in|frac=2|abbr=on}} long, {{convert|124|to|160|cm|in|frac=2|abbr=on}} wingspan, and of {{convert|1180|to|2100|g|lboz|frac=2|abbr=on}} weight, with average weights of {{convert|1585|or|1752|g|lboz|frac=4|abbr=on}}.<ref name=CRC2>{{cite book |title=CRC Handbook of Avian Body Masses |edition=2nd |editor-first=John B. Jr. |editor-last=Dunning |publisher=CRC Press |year=2008 |isbn=978-1-4200-6444-5}}</ref><ref name= Palmer>Palmer, R. S. (ed.) (1988). ''Handbook of North American birds. Volume 5 Diurnal Raptors (part 2)''. 465 pp.</ref> An outsized female from eastern [[Siberia]] was found to have scaled {{convert|2600|g|lboz|abbr=on}}.<ref name= Palmer/><ref>Dementiev, G. P. 1960. ''Der Gerfalke (Falco gyrfalco L. = Falco rusticolus L)''. Die Neue Brehm-Bücherei, no. 264. A. Ziemen Verlag, Wittenberg, Germany.</ref> Among standard measurements, the [[Wing chord (biology)|wing chord]] is {{convert|34.5|to|41|cm|in|abbr=on}}, the [[tail]] is {{convert|19.5|to|29|cm|in|abbr=on}}, the [[Culmen (bird)|culmen]] is {{convert|2|to|2.8|cm|in|frac=8|abbr=on}} and the [[Tarsus (skeleton)|tarsus]] is {{convert|4.9|to|7.5|cm|in|frac=8|abbr=on}}.<ref name=BNA/> The gyrfalcon is larger, broader-winged and longer-tailed than the [[peregrine falcon]], which it is known to compete with (and occasionally hunt). It differs from the [[buzzard]] in general structure, having pointed wings.
This species is a very large falcon, about the same size as the largest [[buteo]]s. Males are 48 to 61 cm (19 to 24 in) long, weigh 805 to 1350 g (1.8 to 3 lbs) and have a wingspan from 110 to 130 cm (43 to 51 in). Females are rather bulkier and larger at 51 to 65 cm (20 to 26 in) long, a weight of 1180 to 2100 g (2.6 to 4.6 lbs) and have a wingspan ranging from 124 to 160 cm (49 to 64 in). Gyrfalcons are somewhat intermediate between a large [[Peregrine Falcon]] and a [[hawk]] in general structure; they are unmistakably falcons with pointed wings, but are stockier, broader-winged and longer-tailed than the Peregrine.


Plumage is very variable in this highly [[polymorphic]] species: the archetypal morphs are called "white", "silver", "brown" and "black" though coloration spans a continuous spectrum from nearly all-white birds to very dark ones. The brown form of the Gyrfalcon is distinguished from the Peregrine by the cream streaking on the nape and crown and by the absence of a well-defined malar stripe and cap. The black morph has its underside strongly spotted black, not finely barred as in the Peregrine. White form Gyrfalcons are unmistakable, as they are the only predominantly white falcons. Silver birds resemble a light, grey [[Lanner Falcon]] of huge size.
The gyrfalcon is a very [[Polymorphism (biology)|polymorphic]] species, so its plumage varies greatly. The archetypal morphs are called "white", "silver", "brown", and "black", though they can be coloured on a spectrum from all-white to very dark. The brown form of the gyrfalcon is distinguished from the peregrine by the cream streaking on the nape and crown and by the absence of a well-defined malar stripe and cap. The black morph is similar but has a strongly black-spotted underside, rather than finely barred as in the peregrine and the brown-morph gyrfalcon. White form gyrfalcons are the only predominantly white falcons. Silver gyrfalcons resemble a light grey [[lanner falcon]] of larger size. The species shows no sex-based colour differences; juveniles are darker and browner than adults.


The black color seems to be [[sex-linked]] and to occur mostly in females; it proved difficult for breeders to get males darker than the dark side of [[slate grey]]. A color variety that arose in captive breeding is "black chick".<ref name=falconscanada/>
There is no difference in coloration between between males and females; juveniles are darker and browner than corresponding adults on average.

<gallery>
File:Falco rusticolus NAUMANN.jpg|Painting of a Greenland white morph (center), an intermediate (lower left), and black morph (back)
File:2007 cze 10 034.JPG|Light silver-morph
File:White Gyr on Block P1144.jpg|A white gyrfalcon
File:Falco rusticolus NAUMANN 2.jpg|Painting of brown morph adult (center) and juveniles
File:Målning, Isländsk jaktfalk, 1759 - Livrustkammaren - 89009.tif|Icelandic gyrfalcon, 1759
</gallery>


==Systematics and evolution==
==Systematics and evolution==
[[Image:USAF falcon.jpg|thumb|left|A [[Hybrid (biology)|hybrid]] white gyrfalcon × [[saker falcon]].]]
[[File:USAF falcon.jpg|thumb|right|[[Hybrid (biology)|Hybrid]] white gyrfalcon × [[Saker falcon|saker]]]]
The Gyrfalcon is a member of the close-knit [[hierofalcon]] complex. In this group, there is ample evidence for rampant [[Hybrid (biology)|hybridization]] and incomplete [[lineage sorting]] which confounds analyses of [[DNA sequence]] data to a massive extent; molecular studies with small [[sample size]]s can simply not be expected to yield reliable conclusions in the entire hierofalcon group. The [[radiation (biology)|radiation]] of the entire living diversity of hierofalcons seems to have taken place in the [[Eemian interglacial]] at the start of the [[Late Pleistocene]], a mere 130,000-115,000 years ago; the Gyrfalcon seems to represents lineages that expanded into the [[Holarctic]] and [[adaptation (biology)|adapted]] to local conditions, whereas the inland populations further south, towards northeastern [[Africa]] where the radiation probably originated, evolved into the [[Saker Falcon]]. Indeed, gyrfalcons hybridize not infrequently with Sakers in the [[Altay Mountains]], and this [[gene flow]] seems to be the origin of the "[[Altai Falcon]]".<ref>Helbig ''et al.'' (1994), Wink ''et al.'' (1998), Wink ''et al.'' (2004), Nittinger ''et al.'' (2005)</ref>
The gyrfalcon is a member of the [[hierofalcon]] complex. In this group, ample evidence indicates [[Hybrid (biology)|hybridisation]] and [[incomplete lineage sorting]], which confounds analyses of [[Nucleic acid sequence|DNA sequence]] data to a massive extent. The [[radiation (biology)|radiation]] of the entire living diversity of hierofalcons took place around the [[Eemian|Eemian Stage]] at the start of the [[Late Pleistocene]]. It represents lineages that expanded into the [[Holarctic]] and [[adaptation (biology)|adapted]] to local conditions; this is in contrast to less northerly populations of northeastern [[Africa]] (where the radiation probably originated) that evolved into the [[saker falcon]]. Previous beliefs held that gyrfalcons hybridized with sakers in the [[Altai Mountains]], and this [[gene flow]] contributed to the genetic lineage of the [[Altai falcon]].<ref name=Helbig/><ref name=Wink1998/><ref name=Wink2004/><ref name=Nittinger/> However, [https://www.ncbi.nlm.nih.gov/pmc/articles/PMC10584951/ recent genetic research] has not found distinct genetic clusters differentiating Altai falcons from eastern saker falcons (''Falco cherrug milvipes''), nor evidence supporting the hybridization theory. Instead, this research suggests that gyrfalcons may have evolved from eastern saker falcons, explaining their close genetic relationship.


Some correlation exists between locality and colour morph. [[Greenland]] gyrfalcons are lightest, with white plumage flecked with grey on the back and wings being most common. Other subpopulations have varying amounts of the darker morphs: the Icelandic birds tend towards pale, whereas the Eurasian populations are considerably darker and typically incorporate no white birds. Natural separation into regional subspecies is prevented by gyrfalcons' habit of flying long distances whilst exchanging [[allele]]s between subpopulations; thus, the allele distributions for the color polymorphism form [[cline (biology)|cline]]s and in darker birds<ref group="nb">The allele combination producing the white morph seems to be [[dominance (genetics)|recessive]].</ref> of unknown origin, theoretically any allele combination might be present. For instance, a mating of a pair of captive gyrfalcons is documented to have produced a [[clutch (eggs)|clutch]] of four young: one white, one silver, one brown, and one black. Molecular work suggests plumage color is associated with the [[melanocortin 1 receptor]] gene (''MC1R''), where a nonsynonymous point substitution was perfectly associated with the white/melanic polymorphism.<ref name=Johnson2012/>
===Subspecies===
There is some [[correlation]] between locality and the frequency of color morphs. [[Greenland]] Gyrfalcons are lightest, with white plumage flecked with grey on the back and wings being most common. Other [[subpopulation]]s have varying amounts of the darker morphs: the [[Iceland]]ic birds tend towards pale, and [[Eurasia]]n ones are considerably darker and not usually have white birds present. Natural separation into regional subspecies is prevented by Gyrfalcons' habit of flying long distances exchanging [[allele]]s between subpopulations; thus, the allele distributions for the color polymorphism form [[cline (biology)|cline]]s and in darker birds<ref>The allele combination producing the white morph seems to be [[recessive]].</ref> of unknown origin, theoretically any allele combination might be present. For example, a mating of a pair of captive Gyrfalcons is documented to have produced a [[clutch]] of 4 young: one white, one silver, one brown, one black.


[[File:Gyrfalcon at Lake Myvatn by Bruce McAdam.jpg|thumb|left|Adult ''F. r. islandus'' at [[Dimmuborgir]] near [[Mývatn|Lake Mývatn]] (Iceland)]]
In general, geographic variation follows [[Bergmann's Rule]] for size and the demands of [[crypsis]] for plumage coloration. Several [[subspecies]] have been named according to perceived differences between populations<ref>''Falco rusticolus candicans'' fron northern Greenland and adjacent North America; ''F. r. obsoletus'' from the southern Greenland into [[subarctic]] North America which is much darker, often brown or black; and ''F. r. islandus'' (Iceland), ''F. r. rusticolus'' (Scandinavia inluding the species' [[type locality (biology)|type locality]], Sweden), as well as ''F. r. intermedius'' and ''F. r. grebnitzkii'' (Siberia), which all tend towards more or less dark "silver" coloration (Snow ''et al.'' 1998).</ref> but none of these are consistent and thus no living subspecies are accepted today.
In general, geographic variation follows [[Bergmann's rule]] for size and the demands of [[crypsis]] for plumage coloration. Several [[subspecies]] have been named according to perceived differences between populations<ref group="nb">''Falco rusticolus candicans'' from northern Greenland and adjacent North America, which is often very white, ''F. r. obsoletus'' from the southern Greenland into [[subarctic]] North America is much darker, often brown or black, ''F. r. islandus'' (Iceland), ''F. r. rusticolus'' (Scandinavia including the species' [[type locality (biology)|type locality]], Sweden), as well as ''F. r. intermedius'' and ''F. r. grebnitzkii'' (Siberia) all tend towards more or less dark "silver" coloration.</ref><ref name=Snow/><ref name=Johnson2007/> but none of these are consistent and thus no living subspecies are currently accepted. The Icelandic population described as ''F. r. islandus'' is perhaps the most distinct. The predominantly white Arctic forms are [[parapatric speciation|parapatric]] and seamlessly grade into the [[subarctic]] populations. The Icelandic types are presumed to have less [[gene flow]] with their neighbors; they show less variation in plumage colors. Comprehensive [[phylogeography|phylogeographic]] studies to determine the proper status of the Icelandic population have yet to be performed.<ref name=Snow/><ref name=White/>


A population genetic study,<ref name=Johnson2007/> however, identified the Iceland population as genetically unique relative to other sampled populations in both eastern and western Greenland, Canada, Alaska, and Norway. Further, within Greenland, differing levels of gene flow between western and eastern sampling locations were identified, with apparent asymmetric dispersal in western Greenland from north to south. This dispersal bias is in agreement with the distribution of plumage colour variants with white gyrfalcons in much higher proportion in north Greenland.<ref name=Johnson2007/> Although further work is required to determine the ecological factors contributing to these distributions relative to plumage differences, a study using demographic data suggested that plumage color distribution in Greenland may be influenced by nesting chronology with white individuals and pairs laying eggs earlier in the breeding season and producing more offspring.<ref name=Johnson2013/>
Perhaps the [[Iceland]]ic population described as '''''Falco rusticolus islandus''''' is the most distinct. The predominantly white Arctic forms are [[parapatric]] and seamlessly grade into the [[subarctic]] populations, whereas the birds of Iceland have presumably less [[gene flow]] with their neighbors and indeed show less variation in plumage colors and often look quite similar to a large, washed-out Peregrine Falcon (though their [[habitus]] is different). Comprehensive [[phylogeographic]] studies are needed to determine the proper status of the Icelandic population however.<ref>White (1994), Snow ''et al.'' (1998)</ref>


===Swarth's gyrfalcon===
There was, however, a [[paleosubspecies]] '''''Falco rusticolus swarthi''''' during the [[Late Pleistocene]] (125,000 - 13,000 years ago). Fossils found in [[Little Box Elder Cave]] ([[Converse County, Wyoming]]), [[Dark Canyon Cave]] ([[Eddy County, New Mexico]]) and [[McKittrick, California]] were initially described as ''Falco swarthi'' ("Swarth Falcon" or more properly '''Swarth's Gyrfalcon''') on account of their distinct size. They have meanwhile proven to be largely inseparable from those of living gyrfalcons, except for being somewhat larger.<ref>Miller (1927, 1935), Howard (1971), Emslie (1985)</ref>
A [[chronospecies|paleosubspecies]], ''Falco rusticolus swarthi'', existed during the [[Late Pleistocene]] (125,000 to 13,000 years ago). Fossils found in Little Box Elder Cave ([[Converse County, Wyoming]]), Dark Canyon Cave ([[Eddy County, New Mexico]]), and [[McKittrick, California]] were initially described as ''Falco swarthi'' ("Swarth falcon" or more properly "Swarth's gyrfalcon") on account of their distinct size. They have meanwhile proven to be largely inseparable from those of living gyrfalcons, except for being somewhat larger.<ref name=Miller1927/><ref name=Miller1935/><ref name=Howard/><ref name=Emslie/>


Swarth's Gyrfalcon was on the upper end of the present Gyrfalcon's size range, strong females even surpassing it (Miller 1935). It seems to have had some adaptations to the temperate [[semiarid]] climate that predominated in its range during the [[last ice age]]. Ecologically more similar to the Siberian populations of today (which are generally small birds however) or the [[Prairie Falcon]], this population of temperate [[steppe]] habitat must have preyed on landbirds and mammals rather than the water- and seabirds which make up much of American gyrfalcon's diet today.
Swarth's gyrfalcon was on the upper end of the present gyrfalcon's size range, with some stronger females even surpassing it.<ref name=Miller1935/> It seems to have had some adaptations to the temperate [[semi-arid climate|semiarid]] climate that predominated in its range during the [[quaternary glaciation|last ice age]]. Ecologically more similar to current Siberian populations (which are generally composed of smaller birds) or to the [[prairie falcon]], this temperate [[steppe]] population must have preyed on landbirds and mammals rather than the sea and landbirds which make up much of the American gyrfalcon's diet today.


==Ecology==
==Ecology==
=== Dietary biology ===
The Gyrfalcon is a bird of [[tundra]] and mountains, with cliffs or a few patches of trees. It feeds only on birds and [[mammal]]s. Like other hierofalcons, it usually hunts in a horizontal pursuit, rather than the Peregrine's speedy [[stoop]] from a height. Most prey is killed on the ground, whether they are captured there or, if the victim is a flying bird, forced to the ground. The diet is to some extent opportunistic, but a majority of breeding birds mostly rely on ''[[Lagopus]]'' [[grouse]]. Avian prey can range in size from [[redpoll]]s to [[goose|geese]] and can include [[gull]]s, [[corvid]]s, smaller [[passerine]]s, [[wader]]s and other [[Bird of prey|raptor]]s (up to the size of ''Buteos''). Mammalian prey can range in size from [[shrew]]s to [[marmot]]s (sometimes 3 times heavier than the assaulting falcon), and often includes include [[lemming]]s, [[vole]]s, [[ground squirrel]]s and [[hare]]s. They only rarely eat [[carrion]].
[[File:Gyrfalcon (falco rusticolus) in flight.jpg|thumb|right|Gyrfalcon in flight (Hastings, MN)]]
The gyrfalcon was originally thought to be a bird of [[tundra]] and mountains only; however, in June 2011, it was revealed to spend considerable periods during the winter on sea ice far from land.<ref name=Burnham2011/> It feeds only on birds and [[mammal]]s, the latter of which it takes more regularly than many other ''[[Falco (bird)|Falco]]'' species. Like other hierofalcons, it usually hunts in a horizontal pursuit, rather than with the peregrine's speedy stoop from a height. Most prey is killed on the ground, whether they are captured there, or if the victim is a flying bird, forced to the ground. The diet is to some extent opportunistic, but a majority breed and hunt coinciding with [[ptarmigan]] and seabird colonies. Avian prey can range in size from [[redpoll]]s around {{convert|20|g|oz|abbr=on}} to [[goose|geese]] and [[Tetrao|capercaillie]]s up to {{convert|4|kg|lb|abbr=on}} in weight, but [[ptarmigan]]s (''Lagopus mutus'') and [[Willow Grouse]]s (''L. lagopus'') are often chief prey in the tundra.<ref name = world>Booms, T. L., T. J. Cade, and N. J. Clum (2020). Gyrfalcon (Falco rusticolus), version 1.0. In Birds of the World (S. M. Billerman, Editor). Cornell Lab of Ornithology, Ithaca, NY, USA. https://doi.org/10.2173/bow.gyrfal.01</ref><ref>Nielsen, Ólafur K., and Tom J. Cade. "Seasonal changes in food habits of Gyrfalcons in NE-Iceland." Ornis Scandinavica (1990): 202–211.</ref><ref name = Ferguson-Lees>Ferguson-Lees, James, and David A. Christie. Raptors of the world. Houghton Mifflin Harcourt, 2001.</ref><ref name =Potapov>Potapov, E. U. G. E. N. E. "Gyrfalcon diet: spatial and temporal variation." Gyrfalcons and ptarmigan in a changing world. Edited by RT Watson, TJ Cade, M. Fuller, G. Hunt, and E. Potapov. The Peregrine Fund, Boise, Idaho, USA (2011): 55–64.</ref><ref>Booms, Travis L., and Mark R. Fuller. "Gyrfalcon diet in central west Greenland during the nesting period." The Condor 105.3 (2003): 528–537.</ref><ref name = Nielsen>Nielsen, Ólafur K. "Gyrfalcon predation on ptarmigan: numerical and functional responses." Journal of Animal Ecology 68.5 (1999): 1034–1050.</ref> Seabirds such as [[auk]]s, [[gull]]s and [[Mergini|seaduck]]s may predominate in coastal areas, and [[wader]]s and [[duck]]s such as [[mallard]]s (''Anas platyrhynchos'') on wetlands.<ref name = Ferguson-Lees/><ref>Dekker, D., and Court, G. "Gyrfalcon predation on Mallards and the interaction of Bald Eagles wintering in central Alberta." Journal of Raptor Research 37.2 (2003): 161–163.</ref> Other avian prey include [[corvidae|corvid]]s, smaller [[passerine]]s, [[dove]]s, and other [[birds of prey]]s.<ref>GILYAZOV, A. "Population of diurnal raptors (Falconiformes) in the Lapland Nature Reserve and adjacent areas: Dynamics in 1930–2005." Status of Raptor Populations in Eastern Fennoscandia. Proceedings of the Workshop, Kostomuksha, Karelia, Russia. 2005.</ref><ref>Koskimies, P. E. R. T. T. I., and S. E. P. P. O. Sulkava. "Diet of the Gyrfalcon ( Falco rusticolus) in northern Fennoscandia." Gyrfalcons and Ptarmigan in a Changing World. The Peregrine Fund, Boise, Idaho, USA. http://dx {{Webarchive|url=https://web.archive.org/web/20220530053146/http://dx/ |date=2022-05-30 }}. doi. org/10 4080 (2011).</ref>


Mammalian prey can be locally important, mainly [[Arctic ground squirrel]]s (Spermophilus parryii) and [[Arctic hare]]s (Lepus arcticus), and occasionally [[Norway lemming]] (''Lemmus lemmus'') in peak years.<ref name = Ferguson-Lees/> Due to the limit of load that they can carry, gyrfalcons mainly take young hares, but both male and female falcons can take down adult hares up {{convert|4.5|kg|lb|abbr=on}} in weight and bring dismembered pieces to their nest.<ref name = world/><ref name =Muir>Muir, Dalton, and David M. Bird. "Food of gyrfalcons at a nest on Ellesmere Island." The Wilson Bulletin (1984): 464–467.</ref><ref name =Booms>Booms, Travis L., and Mark R. Fuller. "Gyrfalcon diet in central west Greenland during the nesting period." The Condor 105.3 (2003): 528–537.</ref><ref>Poole, Kim Gordon, and D. A. Boag. "Ecology of gyrfalcons, Falco rusticolus, in the central Canadian Arctic: diet and feeding behaviour." Canadian Journal of Zoology 66.2 (1988): 334–344.</ref> Other mammalian prey can include [[Water vole (disambiguation)|water vole]]s, [[muskrat]]s, [[stoat]]s, [[mink]]s, [[Arctic fox]] pups, and rarely also [[bat]]s.<ref name =Potapov/><ref name =Muir/><ref name =Booms/><ref>Heintzelman, Donald S. Hawks and owls of North America: a complete guide to North American birds of prey. Universe Books, 1979.</ref><ref>Mikula, P., Morelli, F., Lučan, R.K., Jones, D.N., & Tryjanowski, P. (2016). Bats as prey of diurnal birds: a global perspective. ''Mammal Review'' [https://www.researchgate.net/publication/282980019_Bats_as_prey_of_diurnal_birds_a_global_perspective DOI: 10.1111/mam.12060].</ref> Prey other than birds and mammals are extremely rare, but [[brown trout]] (''Salmo trutta'') have been recorded as prey.<ref name = Nielsen/>
===Reproduction and life history===
[[Image:Falco rusticolus chick.jpg|thumb|right|A Gyrfalcon chick hatched in captivity]]
The Gyrfalcon almost invariably nests on cliff faces. Sometimes, the breeding pair will build their own nests, but most nests are the abandoned nests of other birds, particularly [[Golden Eagle]]s and [[Common Raven]]s. The clutch can range from 1 to 5 eggs, but is usually 2 to 4. The average size of an egg is 58.46 x 45 mm (2.31 x 1.8 in) and the average weight is 62 g (2.2 oz). The incubation period averages 35 days, until the 52 g (1.8 oz) chicks hatch. The nestlings are brooded usually for 10 to 15 days and leave the nest shortly thereafter. At 4 to 6 weeks of age, the immatures become independent of their parents, though they may associate with their siblings through the following winter.


===Threat from climate change===
The only natural predator of gyrfalcons are [[Golden Eagle]]s and even they rarely engage with these formidable falcons. Gyrfalcons have been recorded as aggressively harassing animals that come near their nests, although [[Common Raven]]s are the only predators known to successful pick off Gyrfalcon eggs and hatchlings. Even [[Brown Bear]]s may be dive-bombed, much to their annoyance. Humans, whether accidentally (automobile collisions or poisoning of carrion to kill [[mammal]]ian scavengers) or intentionally (through hunting), are the leading cause of death for Gyrfalcons. Most Gyrfalcons that make it to adulthood live to about 9 to 12 years of age.
In the early 2000s, it was observed that as possible climate change began to temper the Arctic summers, [[peregrine falcon]]s were expanding their range north to parts of [[Greenland]], and competing with gyrfalcons. Although it is specially adapted for high-Arctic life, and larger than the peregrine, the gyrfalcon is less aggressive and more conflict-averse, and so is less able to compete with peregrines, which can attack and overwhelm the gyrs.<ref>{{cite news|last1=Meyer|first1=Robinson|title=The Battle Over 2,500-Year-Old Shelters Made of Poop|url=https://www.theatlantic.com/science/archive/2017/07/falcon-battle-over-nests-of-bird-poop/534510/|access-date=2018-05-10|work=The Atlantic|date=2017-07-24}}</ref> However, it remains on the [[International_Union_for_Conservation_of_Nature|IUCN's]] [[IUCN_Red_List|Red List]] with a [[Conservation_status|Conservation Status]] of [[Least-concern_species|Least Concern]].


==Breeding==
==Relationship with humans==
The gyrfalcon almost invariably nests on cliff faces. Breeding pairs do not build their own nests, and often use a bare cliff ledge or the abandoned nest of other birds, particularly [[golden eagle]]s and [[common raven]]s. The clutch can range from 1 to 5 eggs, but is usually 2 to 4. The average size of an egg is {{convert|58.46|×|45|mm|in|frac=4|abbr=on}}; the average weight is {{convert|62|g|oz|frac=8|abbr=on}}. The incubation period averages 35 days, with the chicks hatching at a weight of around {{convert|52|g|oz|frac=8|abbr=on}}. The nestlings are brooded usually for 10 to 15 days and leave the nest at 7 to 8 weeks. At 3 to 4 months of age, the immature gyrfalcons become independent of their parents, though they may associate with their siblings through the following winter.
The Gyrfalcon is [[Iceland]]'s ''staðfugl'' ([[national bird]]), as well as [[List of national birds#Official birds of subnational entities|the official bird]] of [[Canada]]'s [[Northwest Territories]].


The only natural predators of gyrfalcons are [[golden eagle]]s, and even they rarely engage with these formidable falcons. Gyrfalcons have been recorded as aggressively harassing animals that come near their nests, although common ravens are the only predators known to successfully pick off gyrfalcon eggs and hatchlings. Even [[brown bear]]s have been reportedly dive-bombed. Humans, whether accidentally (automobile collisions or poisoning of carrion to kill [[mammal]]ian scavengers) or intentionally (through hunting), are the leading cause of death for gyrfalcons. Gyrfalcons that survive into adulthood can live up to 20 years of age.
In medieval times, the Gyrfalcon was considered the [[Monarch|king]]'s bird. Due to its rarity and the difficulties involved in obtaining it, in [[falconry]] the gyrfalcon was generally reserved for kings and nobles. Very seldom was a man of lesser rank seen with a Gyrfalcon on his fist.{{Fact|date=February 2007}}


As ''F. rusticolus'' has such a wide range, it is not considered a threatened species by the [[International Union for Conservation of Nature|IUCN]].<ref name="IUCN" /> It is not much affected by [[habitat destruction]], but pollution, for instance by [[pesticide]]s, depressed its numbers in the mid-20th century, and until 1994 it was considered "[[Near Threatened]]". Improving environmental standards in developed countries have allowed the birds to make a comeback.<ref name="IUCN" />
Gyrfalcons are very expensive to buy, and thus owners and breeders may keep them secret to avoid theft. They tend to fly long distances, and falconers may fit a radio-tracker to aid recovery.{{Fact|date=February 2007}}


Wild Gyrfalcons are not much exposed to disease, and as a result have weak [[immune system]]s. As a result, many gyrfalcons taken from the wild quickly die of disease. Several generations of captive breeding from the survivors causes selection for a stronger immune system and thus better resistance to disease.<!-- (Ref.[http://www.falconscanada.com/index.htm], about a third of the way down the page) [link dead as of 2007-AUG-13] -->

===Gyrfalcons and the fall of the Liao Dynasty===
In the 12th century AD the [[Jurchen]] tribes [[rebellion|rebelled]] against the [[China|Chinese]] [[Liao Dynasty]] which was set up by [[Khitan people]]. The primary cause was that the Khitan [[noble]]s extorted a big [[tax]] of Gyrfalcons (which are called ''Hǎidōngqīng'' (海东青) in Chinese), as [[swan]] hunting was highly fashionable among the nobility. Especially under the last Liao Emperor [[Emperor Tianzuo of Liao|Tiānzuòdì]], tax collectors were even entitled to use force to procure the demanded quantity of gyrfalcons. The rebellion caught on, and the Jurchen under chieftain [[Wányán Āgǔdǎ]] annihilated the Liao empire in [[1125]], establishing the [[Jin Dynasty, 1115–1234|Jīn Dynasty]] in its stead.<ref>Beijing Municipal Administration of Cultural Heritage: [http://english.bjww.gov.cn/2006/7-19//18134.shtml Contest for the Southern Capital between the Liao, Song and Jin Dynasties]. Version of 2006-JUL-19. Retrieved 2007-AUG-13.</ref>

==Images==
<gallery>
<gallery>
File:Faucon gerfaut MHNT.jpg|''Falco rusticolus'' egg
Image:Falco rusticolus NAUMANN.jpg|Painting of a Greenland white morph (center), an entirely intermediate bird (lower left) and black-morph birds (back)
File:Falco rusticolus -Yukon Delta National Wildlife Refuge, Alaska, USA -juvenile-8.jpg|Nestling in Alaska
Image:2007 cze 10 034.JPG|A light silver-morph bird
Image:Aa gyrtiercel 2006 03 18.jpg|Male with a darker "silver" coloration
Image:Falco rusticolus NAUMANN 2.jpg|Brown morph adult (center) and juveniles
</gallery>
</gallery>


== Interaction with humans ==
==Footnotes==
[[File:Falconry sport of kings (1920) Gerfalcon striking heron.png|thumb|A tamed gyrfalcon striking a wild [[grey heron]] (1920), [[Louis Agassiz Fuertes]]]]
<references/>
The gyrfalcon has long associated with humans, primarily for hunting and in the art of [[falconry]]. It is the official bird of Canada's [[Northwest Territories]]. The white falcon in the crest of the [[Iceland|Icelandic Republic]]'s coat of arms is a variety of gyrfalcon. The white phase gyrfalcon is the official mascot of the [[United States Air Force Academy]].<ref name=britannicaGyrfalcon/>


In the medieval era, the gyrfalcon was considered a [[Monarch|royal]] bird. The geographer and historian [[Ibn Sa'id al-Maghribi]] (d. 1286) described certain northern [[Atlantic Ocean|Atlantic]] islands west of [[Ireland]] where these falcons would be brought from, and how the Egyptian [[Sultan]] paid 1,000 [[dinar]]s for each gyrfalcon (or, if it arrived dead, 500 dinars).<ref name=al-Maghribi/> Due to its rarity and the difficulties involved in obtaining it, in European falconry the gyrfalcon was reserved for kings and nobles; very rarely was a man of lesser rank seen with a gyrfalcon on his fist.<ref name=Berners/>
==References==
* {{IUCN2006|assessors={{aut|BirdLife International}}|year=2004|id=49516|title=Falco rusticolus|downloaded=[[12 May]] [[2006]]}} Database entry includes justification for why this species is of least concern.
* {{aut|Emslie, Steven D.}} (1985): The late Pleistocene (Rancholabrean) avifauna of Little Box Elder Cave, Wyoming. ''Rocky Mountain Geology'' '''23'''(2): 63-82. [http://pubs.gg.uwyo.edu/CTGs/CTG_23-2.htm HTML abstract]
* {{aut|Helbig, A.J.; Seibold, I.; Bednarek, W.; Brüning, H.; Gaucher, P.; Ristow, D.; Scharlau, W.; Schmidl, D. & Wink, Michael}} (1994): Phylogenetic relationships among falcon species (genus Falco) according to DNA sequence variation of the cytochrome b gene. ''In:'' {{aut|Meyburg, B.-U. & Chancellor, R.D. (eds.)}}: ''Raptor conservation today'': 593-599. [http://www.uni-heidelberg.de/institute/fak14/ipmb/phazb/pubwink/1994/4.%201994.pdf PDF fulltext]
* {{aut|Howard, Hildegarde}} (1971): Quaternary Avian Remains from Dark Canyon Cave, New Mexico. ''[[Condor (journal)|Condor]]'' '''73'''(2): 237-240. <small>{{doi|10.2307/1365844}}</small> [http://elibrary.unm.edu/sora/Condor/files/issues/v073n02/p0237-p0240.pdf PDF fulltext]
* {{aut|Miller, Loye H.}} (1927): The Falcons of the McKittrick Pleistocene. ''[[Condor (journal)|Condor]]'' '''29'''(3): 150-152. <small>{{doi|10.2307/1363081}}</small> [http://elibrary.unm.edu/sora/Condor/files/issues/v029n03/p0150-p0152.pdf PDF fulltext]
* {{aut|Miller, Loye H.}} (1935): A Second Avifauna from the McKittrick Pleistocene. ''[[Condor (journal)|Condor]]'' '''37'''(2): 72-79. <small>{{doi|10.2307/1363879}}</small> [http://elibrary.unm.edu/sora/Condor/files/issues/v037n02/p0072-p0079.pdf PDF fulltext]
* {{aut|Nittinger, F.; Haring, E.; Pinsker, W.; Wink, Michael & Gamauf, A.}} (2005): Out of Africa? Phylogenetic relationships between ''Falco biarmicus'' and other hierofalcons (Aves Falconidae). ''Journal of Zoological Systematics and Evolutionary Research'' '''43'''(4): 321-331. <small>{{doi|10.1111/j.1439-0469.2005.00326.x}}</small> [http://www.uni-heidelberg.de/institute/fak14/ipmb/phazb/pubwink/2005/28.2005.pdf PDF fulltext]
* {{aut|[[Royal Alberta Museum]]}} (2006): ''[http://www.royalalbertamuseum.ca/vexhibit/dragon/arrows.htm Dragon Bytes: Did you know about time's arrows?]. Version of October 12, 2006. Retrieved 2007-AUG-13.+
* {{aut|Snow, David W.; Perrins, Christopher M.; Doherty, Paul & Cramp, Stanley}} (1998): ''The complete birds of the western Palaearctic on CD-ROM''. Oxford University Press. <small>ISBN 0192685791</small>
* {{aut|White, Clayton M.}} (1994): 58. Gyrfalcon. ''In:'' {{aut|del Hoyo, Josep; Elliott, Andrew & Sargatal, Jordi}} (editors): ''[[Handbook of Birds of the World]], Volume 2 (New World Vultures to Guineafowl)'': 274, plate 28. Lynx Edicions, Barcelona. <small>ISBN 84-87334-15-6</small>
* {{aut|Wink, Michael; Seibold, I.; Lotfikhah, F. & Bednarek, W.}} (1998): Molecular systematics of holarctic raptors (Order Falconiformes). ''In:'' {{aut|Chancellor, R.D., Meyburg, B.-U. & Ferrero, J.J. (eds.)}}: ''Holarctic Birds of Prey'': 29-48. Adenex & WWGBP. [http://www.uni-heidelberg.de/institute/fak14/ipmb/phazb/pubwink/1998/31.%201998.pdf PDF fulltext]
* {{aut|Wink, Michael; Sauer-Gürth, Hedi; Ellis, David & Kenward, Robert}} (2004): Phylogenetic relationships in the Hierofalco complex (Saker-, Gyr-, Lanner-, Laggar Falcon). ''In:'' {{aut|Chancellor, R.D. & Meyburg, B.-U. (eds.)}}: ''Raptors Worldwide'': 499-504. WWGBP, Berlin. [http://www.uni-heidelberg.de/institute/fak14/ipmb/phazb/pubwink/2004/27.2004.pdf PDF fulltext]


In the 12th century AD China, swan-hunting with gyrfalcons (海東青 ''hǎidōngqīng'' in Chinese) obtained from the [[Jurchen people|Jurchen]] tribes became fashionable among the [[Khitan people|Khitan]] nobility. When demand for gyrfalcons exceeded supply, the [[Liao Dynasty|Liao Emperor]] imposed a tax payment-in-kind of gyrfalcons on the Jurchen; under the [[Emperor Tianzuo of Liao|last Liao emperor]], tax collectors were entitled to use force to procure sufficient gyrfalcons. This was one cause of the Jurchen rebellion, whose leader [[Emperor Taizu of Jin|Aguda]] annihilated the Liao empire in 1125, and established the [[Jin dynasty (1115–1234)|Jin dynasty]] in its stead.<ref name=BMACH/>
==External links==
{{commons|Falco rusticolus}}
*http://www.falconryforum.co.uk
*http://www.falconscanada.com/index.htm
*http://www.kentishfalconry.co.uk/
*http://www.gyrfalcons.co.uk/
*http://www.mosquitonet.com/~akfalconer/ Alaska Falcons
*[http://www.sdakotabirds.com/species/gyrfalcon_info.htm Gyrfalcon Information and Photos] - South Dakota Birds and Birding
*[http://www.birds.cornell.edu/AllAboutBirds/BirdGuide/Gyrfalcon.html Gyrfalcon Species Account] - Cornell Lab of Ornithology
*[http://www.bird-stamps.org/cspecies/3205800.htm Stamps] (with RangeMap)
*[http://vireo.acnatsci.org/search.html?Form=Search&SEARCHBY=Common&KEYWORDS=gyrfalcon&showwhat=images&AGE=All&SEX=All&ACT=All&Search=Search&VIEW=All&ORIENTATION=All&RESULTS=24 Gyrfalcon photo gallery] VIREO


Falcons are known to be very susceptible to [[avian influenza]]. Therefore, an experiment was done with hybrid gyr-[[saker falcon|saker]] falcons, which found that five falcons vaccinated with a commercial H5N2 influenza vaccine survived infection with a highly pathogenic H5N1 strain, whereas five unvaccinated falcons died. Thus, both wild and captive gyrfalcons can be protected from bird flu by vaccination.<ref name=Lierz2007/>
[[Category:Falco]]
[[Category:Falcons]]
[[Category:Falconry]]
[[Category:Birds of North America]]
[[Category:Birds of Canada]]
[[Category:Birds of Greenland]]
[[Category:Arctic birds]]
[[Category:Arctic land animals]]
[[Category:Circumpolar species]]


== Notes ==
{{Link FA|de}}
{{Reflist|group="nb"}}


== References ==
[[ar:صقر جير]]
{{Reflist|35em|refs=
[[de:Gerfalke]]
<ref name=al-Maghribi>{{cite web |url=http://sh.rewayat2.com/boldan/Web/4097/001.htm |author=Ibn Sa'id al-Maghribi |title=Geographia |language=ar |website=sh.rewayat2.com |access-date=18 December 2011 |archive-url=https://web.archive.org/web/20120302123907/http://sh.rewayat2.com/boldan/Web/4097/001.htm |archive-date=2 March 2012 |url-status=dead}}</ref>
[[es:Falco rusticolus]]

[[eo:Ĉasfalko]]
<ref name=BMACH>{{cite web |publisher=Beijing Municipal Administration of Cultural Heritage |url=http://english.bjww.gov.cn/2006/7-19//18134.shtml |title=Contest for the Southern Capital between the Liao, Song and Jin Dynasties |date=19 July 2006 |access-date=13 August 2007 |archive-url=https://web.archive.org/web/20070930155539/http://english.bjww.gov.cn/2006/7-19//18134.shtml |archive-date=30 September 2007 |url-status=dead }}</ref>
[[fr:Faucon gerfaut]]

[[gl:Falcón albar]]
<ref name=BNA>{{Cite journal |last1=Booms |first1=Travis L. |first2=Tom J. |last2=Cade |first3=Nancy J. |last3=Clum |year=2008 |title=Gyrfalcon (Falco rusticolus) |journal=The Birds of North America Online |editor-first=A. |editor-last=Poole |url=http://bna.birds.cornell.edu/bna/species/114 |doi=10.2173/bna.114 |access-date=9 March 2013}}</ref>
[[is:Fálki]]

[[it:Falco rusticolus]]
<ref name=Berners>{{Cite book |last=Berners |first=Juliana |year=1486 |title=The Boke of St. Albans |publisher=St. Albans Press |location=London|title-link=Book of St. Albans }}</ref>
[[lt:Medžioklinis sakalas]]

[[hu:Északi sólyom]]
<ref name=britannicaGyrfalcon>{{cite web |url=http://www.britannica.com/EBchecked/topic/250470/gyrfalcon |title=Gyrfalcon |website=Britannica.com |access-date=1 October 2014}}</ref>
[[nl:Giervalk]]

[[ja:シロハヤブサ]]
<!--(this citation is not used in content and was thus giving a cite error)
[[no:Jaktfalk]]
<ref name=britannicaRurik>{{cite web |url=http://www.britannica.com/EBchecked/topic/512996/Rurik |title=Rurik (Norse leader) |website= Britannica.com}}</ref>-->
[[nn:Jaktfalk]]

[[pl:Sokół norweski]]
<ref name=Burnham2011>{{cite journal |last1=Burnham |first1=Kurt K. |last2=Newton |first2=Ian |title=Seasonal movements of Gyrfalcons Falco rusticolus include extensive periods at sea |year=2011 |journal=Ibis |volume=153 |issue=3 |pages=468 |doi=10.1111/j.1474-919X.2011.01141.x|doi-access=}}</ref>
[[pt:Falcão-gerifalte]]

[[se:Rievssatfálli]]
<!--(this citation is not used in content and was thus giving a cite error)
[[sl:Norveški sokol]]
<ref name="chernov-trezin.narod.ru">{{Cite web |first=A. Yu. |last=Chernov |date=July 2008 |url=http://chernov-trezin.narod.ru/GerbRurika.htm |title=Coat of arms of Rurik found in Ladoga |language=ru}}</ref>-->
[[fi:Tunturihaukka]]

[[sv:Jaktfalk]]
<ref name=Emslie>{{cite journal |last=Emslie |first=Steven D. |year=1985 |url=http://rmg.geoscienceworld.org/content/23/2/63.citation |title=The late Pleistocene (Rancholabrean) avifauna of Little Box Elder Cave, Wyoming |journal=Rocky Mountain Geology |volume=23 |issue=2 |pages=63–82 |access-date=2011-12-18 |archive-url=https://archive.today/20130414194904/http://rmg.geoscienceworld.org/content/23/2/63.citation |archive-date=2013-04-14 |url-status=dead}}</ref>
[[tr:Ak sungur]]

[[zh:海东青]]
<ref name=falconscanada>{{Cite web |last1=Lejeune |first1=John |last2=Lejeune |first2=Ginny |last3=Cromarty |first3=Monica |url=http://www.falconscanada.com/Black_Gyrs.html |title=Black Gyrs |website=Falconscanada.com |access-date=18 December 2011 |archive-date=22 October 2015 |archive-url=https://web.archive.org/web/20151022093642/http://www.falconscanada.com/Black_Gyrs.html |url-status=dead }}</ref>

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<ref name=Helbig>{{cite conference |last1=Helbig |first1=A.J. |last2=Seibold |first2=I. |last3=Bednarek |first3=W. |last4=Brüning |first4=H. |last5=Gaucher |first5=P. |last6=Ristow |first6=D. |last7=Scharlau |first7=W. |last8=Schmidl |first8=D. |last9=Wink |first9=Michael |year=1994 |title=Phylogenetic relationships among falcon species (genus Falco) according to DNA sequence variation of the cytochrome b gene |editor1-last=Meyburg |editor1-first=B.-U. |editor2-last=Chancellor |editor2-first=R.D. |work=Raptor conservation today |pages=593–599 |url=http://www.uni-heidelberg.de/institute/fak14/ipmb/phazb/pubwink/1994/4.%201994.pdf}}</ref>

<ref name=Howard>{{cite journal |last=Howard |first=Hildegarde |year=1971 |url=http://sora.unm.edu/sites/default/files/journals/condor/v073n02/p0237-p0240.pdf |title=Quaternary Avian Remains from Dark Canyon Cave, New Mexico |journal=[[Condor (journal)|Condor]] |volume=73 |issue=2 |pages=237–240 |doi=10.2307/1365844|jstor=1365844 }}</ref>

<ref name=Johnson2007>{{cite journal |last1=Johnson |first1=Jeff A. |last2=Burnham |first2=Kurt K. |last3=Burnham |first3=William A. |last4=Mindell |first4=David P. |year=2007 |url=http://www.biol.unt.edu/~jajohnson/pub_pdfs/Johnson_etal_2007_Gyrfalcon.pdf |title=Genetic structure among continental and island populations of Gyrfalcons |pages= 3145–3160 |doi=10.1111/j.1365-294X.2007.03373.x |journal=Molecular Ecology |volume=16 |issue=15 |pmid=17651193|bibcode=2007MolEc..16.3145J |hdl=2027.42/71471 |s2cid=17437176 |hdl-access=free }}</ref>

<ref name=Johnson2012>{{cite journal |last1=Johnson |first1=Jeff A. |last2=Ambers |first2=Angie D. |last3=Burnham |first3=Kurt K. |year=2012 |title=Genetics of plumage color in the Gyrfalcon (Falco rusticolus): analysis of the melanocortin-1 receptor gene |journal=Journal of Heredity |volume=103 |issue=3 |pages=315–321 |doi=10.1093/jhered/ess023 |pmid=22504110|doi-access=free }}</ref>

<ref name=Johnson2013>{{cite journal |last1=Johnson |first1=Jeff A. |last2=Burnham |first2=Kurt K. |year=2013 |title=Timing of breeding and offspring number covary with plumage colour among Gyrfalcons Falco rusticolus |journal=Ibis |volume=155 |issue=1 |pages=177–188 |doi=10.1111/j.1474-919X.2012.01276.x|citeseerx=10.1.1.472.3371 }}</ref>

<ref name=Lierz2007>{{Cite journal |last1=Lierz |first1=Michael |first2=Hafez M. |last2=Hafez |first3=Robert |last3=Klopfleisch |first4=Dörte |last4=Lüschow |first5=Christine |last5=Prusas |first6=Jens P. |last6=Teifke |first7=Miriam |last7=Rudolf |first8=Christian |last8=Grund |first9=Donata |last9=Kalthoff |first10=Thomas |last10=Mettenleiter |first11=Martin |last11=Beer |first12=Timm |last12=Harder |title=Protection and Virus Shedding of Falcons Vaccinated against Highly Pathogenic Avian Influenza A Virus (H5N1) |journal=Emerging Infectious Diseases |volume=13 |issue=11 |pages=1667–1674 |date=November 2007 |pmid=18217549 |doi=10.3201/eid1311.070705 |pmc=3375792}}</ref>

<ref name=Miller1927>{{cite journal |last=Miller |first=Loye H. |year=1927 |url=http://sora.unm.edu/sites/default/files/journals/condor/v029n03/p0150-p0152.pdf |title=The Falcons of the McKittrick Pleistocene |journal=[[Condor (journal)|Condor]] |volume=29 |issue=3 |pages=150–152 |doi=10.2307/1363081|jstor=1363081 }}</ref>

<ref name=Miller1935>{{cite journal |last=Miller |first=Loye H. |year=1935 |url=http://sora.unm.edu/sites/default/files/journals/condor/v037n02/p0072-p0079.pdf |title=A Second Avifauna from the McKittrick Pleistocene |journal=[[Condor (journal)|Condor]] |volume=37 |issue=2 |pages=72–79 |doi=10.2307/1363879|jstor=1363879 }}</ref>

<ref name=Nittinger>{{cite journal |last1=Nittinger |first1=F. |last2=Haring |first2=E. |last3=Pinsker |first3=W. |last4=Wink |first4=Michael |last5=Gamauf |first5=A. |year=2005 |title=Out of Africa? Phylogenetic relationships between ''Falco biarmicus'' and other hierofalcons (Aves Falconidae) |journal=Journal of Zoological Systematics and Evolutionary Research |volume=43 |issue=4 |pages=321–331 |doi=10.1111/j.1439-0469.2005.00326.x |url=http://www.uni-heidelberg.de/institute/fak14/ipmb/phazb/pubwink/2005/28.2005.pdf}}</ref>

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<ref name=White>{{cite book |last=White |first=Clayton M. |year=1994 |number=58 |editor1-last=del Hoyo |editor1-first=Josep |editor2-last=Elliott |editor2-first=Andrew |editor3-last=Sargatal |editor3-first=Jordi |series=[[Handbook of Birds of the World]] |title=2: New World Vultures to Guineafowl |pages=[https://archive.org/details/handbookofbirdso0001unse/page/274 274, plate 28] |publisher=Lynx Edicions |location=Barcelona |isbn=978-84-87334-15-3 |url=https://archive.org/details/handbookofbirdso0001unse/page/274 }}</ref>

<ref name=Wink1998>{{cite journal |last1=Wink |first1=Michael |last2=Seibold |first2=I. |last3=Lotfikhah |first3=F. |last4=Bednarek |first4=W. |year=1998 |title=Molecular systematics of holarctic raptors (Order Falconiformes) |editor1-last=Chancellor |editor1-first=R.D. |editor2-last=Meyburg |editor2-first=B.-U. |editor3-last=Ferrero |editor3-first=J.J. |journal=Holarctic Birds of Prey |pages=29–48 |publisher=Adenex & WWGBP |url=http://www.uni-heidelberg.de/institute/fak14/ipmb/phazb/pubwink/1998/31.%201998.pdf}}</ref>

<ref name=Wink2004>{{Cite journal |last1=Wink |first1=Michael |last2=Sauer-Gürth |first2=Hedi |last3=Ellis |first3=David |last4=Kenward |first4=Robert |year=2004 |url=http://www.uni-heidelberg.de/institute/fak14/ipmb/phazb/pubwink/2004/27.2004.pdf |title=Phylogenetic relationships in the Hierofalco complex (Saker-, Gyr-, Lanner-, Laggar Falcon) |editor1-last=Chancellor |editor1-first=R.D. |editor2-last=Meyburg |editor2-first=B.-U. |journal=Raptors Worldwide |pages=499–504 |publisher=WWGBP |location=Berlin}}</ref>
}}

== Further reading ==
* Audubon, John James. "[https://www.biodiversitylibrary.org/page/33239914 The Iceland or Jer Falcon]", ''Ornithological Biography'' volume 2 (1834). [https://www.biodiversitylibrary.org/page/33238620 Another specimen] described in volume 4 (1838). [https://www.biodiversitylibrary.org/page/40383093 Illustration] from ''Birds of America'' octavo edition, 1840.
* {{cite encyclopedia |url=https://www.biodiversitylibrary.org/page/5886212 |title=A manual of the ornithology of the United States and of Canada : Gyrfalcon |first=Thomas |last=Nuttall |volume=1, ''The Land Birds'' |year=1832|publisher=Hilliard and Brown}}

==External links==
{{Wiktionary}}
{{Commons category|Falco rusticolus}}
{{Wikispecies|Falco rusticolus}}
* [http://www.mosquitonet.com/~akfalconer/ Alaska Falcons – White and Silver Gyrfalcons]
* [http://www.birds.cornell.edu/AllAboutBirds/BirdGuide/Gyrfalcon.html Gyrfalcon Species Account] – Cornell Lab of Ornithology
* {{usurped|1=[https://web.archive.org/web/20001026141038/http://www.bird-stamps.org/cspecies/3205800.htm Gyrfalcon stamps]}} at {{usurped|1=[https://web.archive.org/web/19990428065259/http://www.bird-stamps.org/ bird-stamps.org]}}
* {{InternetBirdCollection|gyrfalcon-falco-rusticolus}}
* {{BirdLife|22696500|Falco rusticolus}}
* {{Avibase|name=Falco rusticolus}}
* {{VIREO|Gyrfalcon}}
* {{IUCN_Map|22696500|Falco rusticolus}}
* {{Xeno-canto species|Falco|rusticolus|Gyrfalcon}}

{{Taxonbar|from=Q177688}}
{{Authority control}}

[[Category:Birds of the Arctic]]
[[Category:Arctic land animals]]
[[Category:Birds described in 1758]]
[[Category:Taxa named by Carl Linnaeus]]
[[Category:Holarctic birds]]
[[Category:Falco (genus)]]
[[Category:Falconry]]
[[Category:Territorial symbols of the Northwest Territories]]

Latest revision as of 20:03, 25 December 2024

Gyrfalcon
CITES Appendix I (CITES)[2]
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Falconiformes
Family: Falconidae
Genus: Falco
Subgenus: Hierofalco
Species:
F. rusticolus
Binomial name
Falco rusticolus
Linnaeus, 1758
Synonyms
List
  • Falco arcticus Holbøll, 1843
  • Falco candicans Gmelin, 1788
  • Falco gyrfalco Linnaeus, 1758
  • Falco islandus Brünnich, 1764
  • Falco obsoletus Gmelin, 1788
  • Falco rusticolus candicans Gmelin, 1788
  • Falco rusticolus grebnitzkii (Severtzov, 1885)
  • Falco rusticolus intermedius Gloger, 1834
  • Falco rusticolus islandus Brünnich, 1764 (but see text)
  • Falco rusticolus obsoletus Gmelin, 1788
  • Falco rusticolus rusticolus Linnaeus, 1758
  • Falco swarthi L.H. Miller, 1927
  • Hierofalco grebnitzkii Severtzov, 1885)
  • Hierofalco islandus (Brünnich, 1764)
  • Hierofalco rusticolus (Linnaeus, 1758)
  • Hierofalco rusticolus candicans (Gmelin, 1788)

The gyrfalcon (/ˈɜːrˌfɔː(l)kən/ or /ˈɜːrˌfælkən/)[3] (Falco rusticolus), also abbreviated as gyr, is a bird of prey from the genus Falco (falcons and kestrels) and the largest species of the family Falconidae.[4] A high-latitude species, the gyrfalcon breeds on the Arctic coasts and tundra, the islands of northern North America and the Eurosiberian region, where it is mainly a resident species. Some gyrfalcons disperse more widely after the breeding season or in winter, and individual vagrancy can take birds for long distances. Its plumage varies with location, with birds being coloured from all-white to dark brown. These colour variations are called morphs. Like other falcons, it shows sexual dimorphism, with the female much larger than the male.

For centuries, the gyrfalcon has been valued as a hunting bird. Typical prey includes the ptarmigan and waterfowl, which it may attack in flight; and it also hunts fish and small mammals.

Taxonomy and etymology

[edit]

The gyrfalcon was formally described by Swedish naturalist Carl Linnaeus in 1758 in the tenth edition of his Systema Naturae under its current binomial name Falco rusticolus.[5] The genus name is the Late Latin term for a falcon, Falco, from falx a sickle, referencing the talons of the bird.[6] The species name is from the Latin rusticolus, a countryside-dweller, from rus, "country" and colere, "to dwell".[7] The bird's common name comes from French gerfaucon; in Medieval Latin, it is gyrofalco. The first part of the word may come from Old High German gîr (cf. modern German Geier; ultimately from Proto-Germanic *girį̄ ("greed")) for "vulture", referring to its size in comparison with other falcons; or from the Latin gȳrus for "circle" or "curved path", in turn from the Ancient Greek γῦρος, gûros, meaning "circle" – from the species' circling as it searches for prey, distinct from the hunting of other falcons in its range.[nb 1] The male gyrfalcon is called a gyrkin in falconry.

Description

[edit]

The gyrfalcon is the largest falcon in the world, being about the same size as the largest buteos but probably slightly heavier. Males are 48 to 61 centimetres (19 to 24 inches) long, weigh 805 to 1,350 grams (1 pound 12+12 ounces to 2 pounds 15+12 ounces), with average weights reported as 1,130 or 1,170 g (2 lb 8 oz or 2 lb 9+12 oz) and have a wingspan from 110 to 130 cm (43 to 51 in). Females are bulkier and larger, at 51 to 65 cm (20 to 25+12 in) long, 124 to 160 cm (49 to 63 in) wingspan, and of 1,180 to 2,100 g (2 lb 9+12 oz to 4 lb 10 oz) weight, with average weights of 1,585 or 1,752 g (3 lb 8 oz or 3 lb 13+34 oz).[8][9] An outsized female from eastern Siberia was found to have scaled 2,600 g (5 lb 12 oz).[9][10] Among standard measurements, the wing chord is 34.5 to 41 cm (13.6 to 16.1 in), the tail is 19.5 to 29 cm (7.7 to 11.4 in), the culmen is 2 to 2.8 cm (34 to 1+18 in) and the tarsus is 4.9 to 7.5 cm (1+78 to 3 in).[11] The gyrfalcon is larger, broader-winged and longer-tailed than the peregrine falcon, which it is known to compete with (and occasionally hunt). It differs from the buzzard in general structure, having pointed wings.

The gyrfalcon is a very polymorphic species, so its plumage varies greatly. The archetypal morphs are called "white", "silver", "brown", and "black", though they can be coloured on a spectrum from all-white to very dark. The brown form of the gyrfalcon is distinguished from the peregrine by the cream streaking on the nape and crown and by the absence of a well-defined malar stripe and cap. The black morph is similar but has a strongly black-spotted underside, rather than finely barred as in the peregrine and the brown-morph gyrfalcon. White form gyrfalcons are the only predominantly white falcons. Silver gyrfalcons resemble a light grey lanner falcon of larger size. The species shows no sex-based colour differences; juveniles are darker and browner than adults.

The black color seems to be sex-linked and to occur mostly in females; it proved difficult for breeders to get males darker than the dark side of slate grey. A color variety that arose in captive breeding is "black chick".[12]

Systematics and evolution

[edit]
Hybrid white gyrfalcon × saker

The gyrfalcon is a member of the hierofalcon complex. In this group, ample evidence indicates hybridisation and incomplete lineage sorting, which confounds analyses of DNA sequence data to a massive extent. The radiation of the entire living diversity of hierofalcons took place around the Eemian Stage at the start of the Late Pleistocene. It represents lineages that expanded into the Holarctic and adapted to local conditions; this is in contrast to less northerly populations of northeastern Africa (where the radiation probably originated) that evolved into the saker falcon. Previous beliefs held that gyrfalcons hybridized with sakers in the Altai Mountains, and this gene flow contributed to the genetic lineage of the Altai falcon.[13][14][15][16] However, recent genetic research has not found distinct genetic clusters differentiating Altai falcons from eastern saker falcons (Falco cherrug milvipes), nor evidence supporting the hybridization theory. Instead, this research suggests that gyrfalcons may have evolved from eastern saker falcons, explaining their close genetic relationship.

Some correlation exists between locality and colour morph. Greenland gyrfalcons are lightest, with white plumage flecked with grey on the back and wings being most common. Other subpopulations have varying amounts of the darker morphs: the Icelandic birds tend towards pale, whereas the Eurasian populations are considerably darker and typically incorporate no white birds. Natural separation into regional subspecies is prevented by gyrfalcons' habit of flying long distances whilst exchanging alleles between subpopulations; thus, the allele distributions for the color polymorphism form clines and in darker birds[nb 2] of unknown origin, theoretically any allele combination might be present. For instance, a mating of a pair of captive gyrfalcons is documented to have produced a clutch of four young: one white, one silver, one brown, and one black. Molecular work suggests plumage color is associated with the melanocortin 1 receptor gene (MC1R), where a nonsynonymous point substitution was perfectly associated with the white/melanic polymorphism.[17]

Adult F. r. islandus at Dimmuborgir near Lake Mývatn (Iceland)

In general, geographic variation follows Bergmann's rule for size and the demands of crypsis for plumage coloration. Several subspecies have been named according to perceived differences between populations[nb 3][18][19] but none of these are consistent and thus no living subspecies are currently accepted. The Icelandic population described as F. r. islandus is perhaps the most distinct. The predominantly white Arctic forms are parapatric and seamlessly grade into the subarctic populations. The Icelandic types are presumed to have less gene flow with their neighbors; they show less variation in plumage colors. Comprehensive phylogeographic studies to determine the proper status of the Icelandic population have yet to be performed.[18][20]

A population genetic study,[19] however, identified the Iceland population as genetically unique relative to other sampled populations in both eastern and western Greenland, Canada, Alaska, and Norway. Further, within Greenland, differing levels of gene flow between western and eastern sampling locations were identified, with apparent asymmetric dispersal in western Greenland from north to south. This dispersal bias is in agreement with the distribution of plumage colour variants with white gyrfalcons in much higher proportion in north Greenland.[19] Although further work is required to determine the ecological factors contributing to these distributions relative to plumage differences, a study using demographic data suggested that plumage color distribution in Greenland may be influenced by nesting chronology with white individuals and pairs laying eggs earlier in the breeding season and producing more offspring.[21]

Swarth's gyrfalcon

[edit]

A paleosubspecies, Falco rusticolus swarthi, existed during the Late Pleistocene (125,000 to 13,000 years ago). Fossils found in Little Box Elder Cave (Converse County, Wyoming), Dark Canyon Cave (Eddy County, New Mexico), and McKittrick, California were initially described as Falco swarthi ("Swarth falcon" or more properly "Swarth's gyrfalcon") on account of their distinct size. They have meanwhile proven to be largely inseparable from those of living gyrfalcons, except for being somewhat larger.[22][23][24][25]

Swarth's gyrfalcon was on the upper end of the present gyrfalcon's size range, with some stronger females even surpassing it.[23] It seems to have had some adaptations to the temperate semiarid climate that predominated in its range during the last ice age. Ecologically more similar to current Siberian populations (which are generally composed of smaller birds) or to the prairie falcon, this temperate steppe population must have preyed on landbirds and mammals rather than the sea and landbirds which make up much of the American gyrfalcon's diet today.

Ecology

[edit]

Dietary biology

[edit]
Gyrfalcon in flight (Hastings, MN)

The gyrfalcon was originally thought to be a bird of tundra and mountains only; however, in June 2011, it was revealed to spend considerable periods during the winter on sea ice far from land.[26] It feeds only on birds and mammals, the latter of which it takes more regularly than many other Falco species. Like other hierofalcons, it usually hunts in a horizontal pursuit, rather than with the peregrine's speedy stoop from a height. Most prey is killed on the ground, whether they are captured there, or if the victim is a flying bird, forced to the ground. The diet is to some extent opportunistic, but a majority breed and hunt coinciding with ptarmigan and seabird colonies. Avian prey can range in size from redpolls around 20 g (0.71 oz) to geese and capercaillies up to 4 kg (8.8 lb) in weight, but ptarmigans (Lagopus mutus) and Willow Grouses (L. lagopus) are often chief prey in the tundra.[27][28][29][30][31][32] Seabirds such as auks, gulls and seaducks may predominate in coastal areas, and waders and ducks such as mallards (Anas platyrhynchos) on wetlands.[29][33] Other avian prey include corvids, smaller passerines, doves, and other birds of preys.[34][35]

Mammalian prey can be locally important, mainly Arctic ground squirrels (Spermophilus parryii) and Arctic hares (Lepus arcticus), and occasionally Norway lemming (Lemmus lemmus) in peak years.[29] Due to the limit of load that they can carry, gyrfalcons mainly take young hares, but both male and female falcons can take down adult hares up 4.5 kg (9.9 lb) in weight and bring dismembered pieces to their nest.[27][36][37][38] Other mammalian prey can include water voles, muskrats, stoats, minks, Arctic fox pups, and rarely also bats.[30][36][37][39][40] Prey other than birds and mammals are extremely rare, but brown trout (Salmo trutta) have been recorded as prey.[32]

Threat from climate change

[edit]

In the early 2000s, it was observed that as possible climate change began to temper the Arctic summers, peregrine falcons were expanding their range north to parts of Greenland, and competing with gyrfalcons. Although it is specially adapted for high-Arctic life, and larger than the peregrine, the gyrfalcon is less aggressive and more conflict-averse, and so is less able to compete with peregrines, which can attack and overwhelm the gyrs.[41] However, it remains on the IUCN's Red List with a Conservation Status of Least Concern.

Breeding

[edit]

The gyrfalcon almost invariably nests on cliff faces. Breeding pairs do not build their own nests, and often use a bare cliff ledge or the abandoned nest of other birds, particularly golden eagles and common ravens. The clutch can range from 1 to 5 eggs, but is usually 2 to 4. The average size of an egg is 58.46 mm × 45 mm (2+14 in × 1+34 in); the average weight is 62 g (2+18 oz). The incubation period averages 35 days, with the chicks hatching at a weight of around 52 g (1+78 oz). The nestlings are brooded usually for 10 to 15 days and leave the nest at 7 to 8 weeks. At 3 to 4 months of age, the immature gyrfalcons become independent of their parents, though they may associate with their siblings through the following winter.

The only natural predators of gyrfalcons are golden eagles, and even they rarely engage with these formidable falcons. Gyrfalcons have been recorded as aggressively harassing animals that come near their nests, although common ravens are the only predators known to successfully pick off gyrfalcon eggs and hatchlings. Even brown bears have been reportedly dive-bombed. Humans, whether accidentally (automobile collisions or poisoning of carrion to kill mammalian scavengers) or intentionally (through hunting), are the leading cause of death for gyrfalcons. Gyrfalcons that survive into adulthood can live up to 20 years of age.

As F. rusticolus has such a wide range, it is not considered a threatened species by the IUCN.[1] It is not much affected by habitat destruction, but pollution, for instance by pesticides, depressed its numbers in the mid-20th century, and until 1994 it was considered "Near Threatened". Improving environmental standards in developed countries have allowed the birds to make a comeback.[1]

Interaction with humans

[edit]
A tamed gyrfalcon striking a wild grey heron (1920), Louis Agassiz Fuertes

The gyrfalcon has long associated with humans, primarily for hunting and in the art of falconry. It is the official bird of Canada's Northwest Territories. The white falcon in the crest of the Icelandic Republic's coat of arms is a variety of gyrfalcon. The white phase gyrfalcon is the official mascot of the United States Air Force Academy.[42]

In the medieval era, the gyrfalcon was considered a royal bird. The geographer and historian Ibn Sa'id al-Maghribi (d. 1286) described certain northern Atlantic islands west of Ireland where these falcons would be brought from, and how the Egyptian Sultan paid 1,000 dinars for each gyrfalcon (or, if it arrived dead, 500 dinars).[43] Due to its rarity and the difficulties involved in obtaining it, in European falconry the gyrfalcon was reserved for kings and nobles; very rarely was a man of lesser rank seen with a gyrfalcon on his fist.[44]

In the 12th century AD China, swan-hunting with gyrfalcons (海東青 hǎidōngqīng in Chinese) obtained from the Jurchen tribes became fashionable among the Khitan nobility. When demand for gyrfalcons exceeded supply, the Liao Emperor imposed a tax payment-in-kind of gyrfalcons on the Jurchen; under the last Liao emperor, tax collectors were entitled to use force to procure sufficient gyrfalcons. This was one cause of the Jurchen rebellion, whose leader Aguda annihilated the Liao empire in 1125, and established the Jin dynasty in its stead.[45]

Falcons are known to be very susceptible to avian influenza. Therefore, an experiment was done with hybrid gyr-saker falcons, which found that five falcons vaccinated with a commercial H5N2 influenza vaccine survived infection with a highly pathogenic H5N1 strain, whereas five unvaccinated falcons died. Thus, both wild and captive gyrfalcons can be protected from bird flu by vaccination.[46]

Notes

[edit]
  1. ^ In Scandinavian languages, it is generally named after its use in falconry, whereas the modern Dutch name giervalk is peculiarly ambiguous: gier means "vulture", whereas gieren means changing the yaw angle to circle in the air.
  2. ^ The allele combination producing the white morph seems to be recessive.
  3. ^ Falco rusticolus candicans from northern Greenland and adjacent North America, which is often very white, F. r. obsoletus from the southern Greenland into subarctic North America is much darker, often brown or black, F. r. islandus (Iceland), F. r. rusticolus (Scandinavia including the species' type locality, Sweden), as well as F. r. intermedius and F. r. grebnitzkii (Siberia) all tend towards more or less dark "silver" coloration.

References

[edit]
  1. ^ a b c BirdLife International (2021). "Falco rusticolus". IUCN Red List of Threatened Species. 2021: e.T22696500A206261845. doi:10.2305/IUCN.UK.2021-3.RLTS.T22696500A206261845.en. Retrieved 15 January 2023.
  2. ^ "Appendices | CITES". cites.org. Retrieved 2022-01-14.
  3. ^ "Definition of gyrfalcon". www.dictionary.com. Retrieved 2022-02-14.
  4. ^ "Gyrfalcon". Audubon Guide to North American Birds. 2014-11-13. Retrieved 2016-04-04.
  5. ^ Linnaeus, C. (1758). Systema Naturæ per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, Volume 1 (in Latin). Vol. v.1 (10th ed.). Holmiae:Laurentii Salvii. p. 88.
  6. ^ Shorter Oxford English dictionary. Oxford: Oxford University Press. 2007. p. 3804. ISBN 978-0199206872.
  7. ^ Jobling, James A (2010). The Helm Dictionary of Scientific Bird Names. London: Christopher Helm. pp. 266, 344. ISBN 978-1-4081-2501-4.
  8. ^ Dunning, John B. Jr., ed. (2008). CRC Handbook of Avian Body Masses (2nd ed.). CRC Press. ISBN 978-1-4200-6444-5.
  9. ^ a b Palmer, R. S. (ed.) (1988). Handbook of North American birds. Volume 5 Diurnal Raptors (part 2). 465 pp.
  10. ^ Dementiev, G. P. 1960. Der Gerfalke (Falco gyrfalco L. = Falco rusticolus L). Die Neue Brehm-Bücherei, no. 264. A. Ziemen Verlag, Wittenberg, Germany.
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Further reading

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