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{{Automatic taxobox
{{Automatic taxobox
| name = Auks
| name = Auks
| fossil_range = [[Late Eocene]] – Recent {{fossil_range|35|0}}
| fossil_range = [[Late Eocene]] – recent {{fossil_range|35|0}}
| image = Parakeetauklets2.jpg
| image = Parakeetauklets2.jpg
| image_caption = [[Parakeet auklet]]s (''Aethia psittacula'')
| image_caption = [[Parakeet auklet]]s (''Aethia psittacula'')
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}}
}}


An '''auk''' or '''alcid''' is a [[bird]] of the family '''Alcidae''' in the order [[Charadriiformes]]. The alcid family includes the [[Uria|murres]], [[guillemot]]s, [[Aethia|auklets]], [[puffin]]s, and [[Brachyramphus|murrelets]]. The word ''auk'' {{IPAc-en|ɔː|k}} is derived from [[Icelandic language|Icelandic]] ''álka'', from [[Old Norse]] ''alka'' (“auk”), from [[Proto-Germanic]] *''alkǭ'' (“sea-bird, auk”).<ref>{{Cite web|url=https://www.merriam-webster.com/dictionary/auk|title=Definition of AUK|website=www.merriam-webster.com}}</ref><ref>{{Cite web|url=https://www.dictionary.com/browse/auk|title=Definition of auk &#124; Dictionary.com|website=www.dictionary.com}}</ref>
'''Auks''' or '''alcids''' are [[bird]]s of the family '''Alcidae''' in the order [[Charadriiformes]].<ref name="ADW">{{cite web |url=https://animaldiversity.org/accounts/Alcidae/classification/#Alcidae |title=Alcidae |date=2022 |last1=Myers |first1=P. |last2=Espinosa |first2=R. |last3=Parr |first3=C. S. |last4=Jones |first4=T. |last5=Hammond |first5=G. S. |last6=Dewey |first6=T. A. |website=Animal Diversity Web |publisher=University of Michigan Museum of Zoology |access-date=31 December 2022}}</ref> The alcid family includes the [[Uria|murres]], [[guillemot]]s, [[Aethia|auklets]], [[puffin]]s, and [[Brachyramphus|murrelets]]. The family contains 25 extant or recently extinct species that are divided into 11 genera. Auks are found throughout the [[Northern Hemisphere]].<ref name="ADW"/><ref name="ioc"/>


Apart from the extinct [[great auk]], all auks can "fly" under water as well as in the air. Although they are excellent swimmers and divers, their walking appears clumsy.
Apart from the extinct [[great auk]], all auks can fly, and are excellent swimmers and divers (appearing to "fly" in water), but their walking appears clumsy.


== Names ==
Several species have different English names in Europe and North America. The two species known as murres in North America are called guillemots in Europe, and the species called little auk in Europe is referred to as dovekie in North America.
Several species have different English names in Europe and North America. The two species known as "murres" in North America are called "guillemots" in Europe, and the species called [[little auk]] in Europe is referred to as dovekie in North America.


==Description==
=== Etymology ===
The word "auk" {{IPAc-en|ɔː|k}} is derived from [[Icelandic language|Icelandic]] ''álka'' and [[Norwegian language|Norwegian]] ''alka'' or ''alke'' from [[Old Norse]] ''ālka'' from [[Proto-Germanic]] *''alkǭ'' (sea-bird, auk).<ref>{{Cite web|url=https://www.merriam-webster.com/dictionary/auk|title=Definition of Auk |website=Merriam-Webster}}</ref><ref>{{Cite web|url=https://www.dictionary.com/browse/auk |title=Definition of auk |website=Dictionary.com}}</ref>
⚫
Auks are superficially similar to [[penguin]]s having black-and-white colours, upright posture and some of their habits. Nevertheless, they are not closely related to penguins, but rather are believed to be an example of moderate [[convergent evolution]]. Auks are [[Sexual dimorphism|monomorphic]] (males and females are similar in appearance).


=== Taxonomy ===
⚫
Extant auks range in size from the [[least auklet]], at 85 g (3&nbsp;oz) and {{convert|15|cm|in|abbr=on}}, to the [[thick-billed murre]], at {{convert|1|kg|lb|abbr=on}} and {{convert|45|cm|in|abbr=on}}.
The [[family (biology)|family]] name Alcidae comes from the genus ''[[Alca (bird)|Alca]]'' given by [[Carl Linnaeus]] in 1758 for the [[razorbill]] (''Alca torda'') from the Norwegian word ''alke''.<ref>{{cite dictionary |last=Jobling |first=James A. |title= Alca |dictionary=The Helm Dictionary of Scientific Bird Names |year=2010 |url=http://www.avesdecostarica.org/uploads/7/0/1/0/70104897/scientific-bird-names.pdf |location=London, United Kingdom |publisher=Christopher Helm |page=40 |isbn=978-1-4081-2501-4}}</ref>
Due to their short wings, auks have to flap their wings very quickly in order to fly.


== Description ==
⚫
Although not to the extent of penguins, auks have largely sacrificed flight, and also mobility on land, in exchange for swimming ability; their wings are a compromise between the best possible design for diving and the bare minimum needed for flying. This varies by subfamily, the ''Uria'' guillemots (including the [[razorbill]]) and murrelets being the most efficient under the water, whereas the puffins and auklets are better adapted for flying and walking.
⚫
Auks are superficially similar to [[penguin]]s, having black-and-white colours, upright posture, and some of their habits. Nevertheless, they are not closely related to penguins, but rather are believed to be an example of moderate [[convergent evolution]]. Auks are [[Sexual dimorphism|monomorphic]] (males and females are similar in appearance).

⚫
Extant auks range in size from the [[least auklet]], at 85 g (3&nbsp;oz) and {{convert|15|cm|in|abbr=on}}, to the [[thick-billed murre]], at {{convert|1|kg|lb|abbr=on}} and {{convert|45|cm|in|abbr=on}}. Due to their short wings, auks have to flap their wings very quickly to fly.

⚫
Although not to the extent of penguins, auks have largely sacrificed flight, and also mobility on land, in exchange for swimming ability; their wings are a compromise between the best possible design for diving and the bare minimum needed for flying. This varies by subfamily, with the ''Uria'' guillemots (including the [[razorbill]]) and murrelets being the most efficient under the water, whereas the puffins and auklets are better adapted for flying and walking.


==Feeding and ecology==
==Feeding and ecology==
The feeding behaviour of auks is often compared to that of [[penguin]]s; both groups are [[wing]]-propelled pursuit divers. In the region where auks live, their only seabird competition are [[cormorant]]s (which are dive-powered by their strong feet). In areas where the two groups feed on the same prey, the auks tend to feed further offshore. Strong-swimming murres hunt faster schooling fish, whereas auklets take slower-moving krill. Time depth recorders on auks have shown that they can dive as deep as {{Convert|100|m|ft|abbr=in}} in the case of ''Uria'' guillemots, {{Convert|40|m|ft|abbr=on}} for the ''Cepphus'' guillemots and {{Convert|30|m|ft|abbr=on}} for the auklets.
The feeding behaviour of auks is often compared to that of penguins; both groups are [[wing]]-propelled, pursuit divers. In the region where auks live, their only seabird competition are [[cormorant]]s (which are dive-powered by their strong feet). In areas where the two groups feed on the same prey, the auks tend to feed further offshore. Strong-swimming murres hunt faster, schooling fish, whereas auklets take slower-moving krill. Time depth recorders on auks have shown that they can dive as deep as {{Convert|100|m|ft|abbr=on}} in the case of ''Uria'' guillemots, {{Convert|40|m|ft|abbr=on}} for the ''Cepphus'' guillemots and {{Convert|30|m|ft|abbr=on}} for the auklets.


==Breeding and colonies==
==Breeding and colonies==
Auks are pelagic birds, spending the majority of their adult life on the open sea and going ashore only for breeding, although some species—like the [[Common murre|common guillemot]]—spend a great part of the year defending their nesting spot from others.
Auks are [[pelagic zone|pelagic]] birds, spending the majority of their adult lives on the open sea and going ashore only for breeding, although some species, such as the [[Common murre|common guillemot]], spend a great part of the year defending their nesting spot from others.


Auks are monogamous, and tend to form lifelong pairs. They typically lay a single egg, and they are [[Philopatry|philopatric]] (they use the nesting site year after year).
Auks are monogamous, and tend to form lifelong pairs. They typically lay a single egg, and they [[Philopatry|use the nesting site]] year after year.


Some species, such as the ''[[Uria]]'' guillemots (murres), nest in large [[bird colony|colonies]] on cliff edges; others, like the ''[[Cepphus]]'' guillemots, breed in small groups on rocky coasts; and the [[puffin]]s, auklets, and some murrelets nest in burrows. All species except the ''[[Brachyramphus]]'' murrelets are [[Bird colony|colonial]].
Some species, such as the ''Uria'' guillemots (murres), nest in large [[bird colony|colonies]] on cliff edges; others, such as the ''[[Cepphus]]'' guillemots, breed in small groups on rocky coasts; and the [[puffin]]s, auklets, and some murrelets nest in burrows. All species except the ''[[Brachyramphus]]'' murrelets are colonial.


==Evolution and distribution==
==Evolution and distribution==
{{See also|List of Charadriiformes by population}}
{{See also|List of Charadriiformes by population}}
[[File:Auks.jpg|thumb|upright|Auks as painted by [[Archibald Thorburn]]]]
[[File:Auks.jpg|thumb|upright|Auks as painted by [[Archibald Thorburn]]]]
Traditionally, the auks were believed to be one of the earliest distinct charadriiform lineages due to their characteristic [[morphology (biology)|morphology]]. However, genetic analyses have demonstrated that these peculiarities are the product of strong natural selection instead: as opposed to, for example, [[plover]]s (a much older charadriiform lineage), auks radically changed from a wading [[wader|shorebird]] to a diving seabird lifestyle. Thus, today, the auks are no longer separated in their own suborder ("'''Alcae'''"), but are considered part of the [[Lari]] suborder which otherwise contains gulls and similar birds. Judging from genetic data, their closest living relatives appear to be the [[skua]]s, with these two lineages separating about 30 [[year|million years ago]] (mya).<ref name=Friesen/><ref name=Moum/><ref name=Thomas/> Alternatively, auks may have split off far earlier from the rest of the Lari and undergone strong morphological, but slow genetic evolution, which would require a very high [[evolutionary pressure]], coupled with a long lifespan and slow reproduction.
Traditionally, the auks were believed to be one of the earliest distinct charadriiform lineages due to their characteristic [[morphology (biology)|morphology]], but genetic analyses have demonstrated that these peculiarities are the product of strong natural selection, instead; as opposed to, for example, [[plover]]s (a much older charadriiform lineage), auks radically changed from a wading [[wader|shorebird]] to a diving seabird lifestyle. Thus today, the auks are no longer separated in their own suborder (Alcae), but are considered part of the [[Lari (bird)|Lari]] suborder, which otherwise contains gulls and similar birds. Judging from genetic data, their closest living relatives appear to be the [[skua]]s, with these two lineages separating about 30 [[year|million years ago]] (Mya).<ref name=Friesen/><ref name=Moum/><ref name=Thomas/> Alternatively, auks may have split off far earlier from the rest of the Lari and undergone strong morphological, but slow genetic evolution, which would require a very high [[evolutionary pressure]], coupled with a long lifespan and slow reproduction.


The earliest unequivocal [[fossil]]s of auks are from the late [[Eocene]], some 35 mya.<ref name=smith2011/> The [[genus]] ''[[Miocepphus]]'', (from the [[Miocene]], 15 mya) is the earliest known from good specimens. Two very fragmentary fossils are often assigned to the Alcidae, although this may not be correct: ''[[Hydrotherikornis]]'' (late [[Eocene]]) and ''[[Petralca]]'' (Late [[Oligocene]]). Most extant genera are known to exist since the Late Miocene or Early [[Pliocene]] (c. 5 mya). Miocene fossils have been found in both [[California]] and [[Maryland]], but the greater diversity of fossils and tribes in the Pacific leads most scientists to conclude that it was there they first evolved, and it is in the Miocene Pacific that the first fossils of extant [[genus|genera]] are found. Early movement between the Pacific and the Atlantic probably happened to the south (since there was no northern opening to the Atlantic), later movements across the Arctic Ocean.<ref name = konyukhov2002/> The flightless subfamily [[Mancallinae]], which was apparently restricted to the Pacific coast of southern North America and became extinct in the Early [[Pleistocene]], is sometimes includes in the family Alcidae under some definitions. One species, ''[[Miomancalla howardae]]'', is the largest charadriiform of all time.<ref>{{cite journal |last1=Smith |first1=N. A. |title=Evolution of body mass in the Pan-Alcidae (Aves, Charadriiformes): the effects of combining neontological and paleontological data |journal=[[Paleobiology (journal)|Paleobiology]] |date=2015 |volume=42 |issue=1 |pages=8–26 |doi=10.1017/pab.2015.24}}</ref>
The earliest unequivocal [[fossil]]s of auks are from the late [[Eocene]], some 35 Mya.<ref name=smith2011/> The [[genus]] ''[[Miocepphus]]'', (from the [[Miocene]], 15 Mya) is the earliest known from good specimens. Two very fragmentary fossils are often assigned to the Alcidae, although this may not be correct: ''[[Hydrotherikornis]]'' (Late [[Eocene]]) and ''[[Petralca]]'' (Late [[Oligocene]]). Most extant genera are known to exist since the Late Miocene or Early [[Pliocene]] (about 5 Mya). Miocene fossils have been found in both [[California]] and [[Maryland]], but the greater diversity of fossils and tribes in the Pacific leads most scientists to conclude they first evolved there, and in the Miocene Pacific, the first fossils of extant [[genus|genera]] are found. Early movement between the Pacific and the Atlantic probably happened to the south (since no northern opening to the Atlantic existed), with later movements across the Arctic Ocean.<ref name = konyukhov2002/> The flightless subfamily [[Mancallinae]], which was apparently restricted to the Pacific Coast of southern North America and became extinct in the Early [[Pleistocene]], is sometimes included in the family Alcidae under some definitions. One species, ''[[Miomancalla howardae]]'', is the largest charadriiform of all time.<ref>{{cite journal |last1=Smith |first1=N. A. |title=Evolution of body mass in the Pan-Alcidae (Aves, Charadriiformes): the effects of combining neontological and paleontological data |journal=[[Paleobiology (journal)|Paleobiology]] |date=2015 |volume=42 |issue=1 |pages=8–26 |doi=10.1017/pab.2015.24|s2cid=83934750 }}</ref>


[[File:Alca torda Roest 2003.jpg|thumb|left|[[Razorbill]]s are true auks only found in the Atlantic Ocean]]
[[File:Alca torda Roest 2003.jpg|thumb|left|[[Razorbill]]s are an auk found in the Atlantic Ocean.]]
The extant auks (subfamily Alcinae) are broken up into 2 main groups: the usually high-billed puffins (tribe Fraterculini) and auklets (tribe Aethiini), as opposed to the more slender-billed murres and true auks (tribe Alcini), and the murrelets and guillemots (tribes Brachyramphini and Cepphini). The tribal arrangement was originally based on analyses of morphology and [[ecology]].<ref name = strauch1985/> [[mitochondrial DNA|mtDNA]] [[cytochrome b|cytochrome ''b'']] [[Nucleic acid sequence|sequence]] and [[allozyme]] studies<ref name=Friesen/><ref name=Moum/> confirm these findings except that the ''[[Synthliboramphus]]'' murrelets should be split into a distinct tribe, as they appear more closely related to the Alcini – in any case, assumption of a closer relationship between the former and the true guillemots was only weakly supported by earlier studies.<ref name = strauch1985 />
The family contains 25 extant or recently extinct species that are divided into 11 genera.<ref name=ioc>{{cite web| editor1-last=Gill | editor1-first=Frank | editor1-link=Frank Gill (ornithologist) | editor2-last=Donsker | editor2-first=David | editor3-last=Rasmussen | editor3-first=Pamela | editor3-link=Pamela Rasmussen | date=August 2022 | title=Noddies, gulls, terns, skimmers, skuas, auks | work=IOC World Bird List Version 12.2 | url=https://www.worldbirdnames.org/bow/gulls/ | publisher=International Ornithologists' Union | access-date=10 November 2022 }}</ref> The extant auks (subfamily Alcinae) are broken up into two main groups - the usually high-billed puffins (tribe Fraterculini) and auklets (tribe Aethiini), as opposed to the more slender-billed murres and true auks (tribe Alcini), and the murrelets and guillemots (tribes Brachyramphini and Cepphini). The tribal arrangement was originally based on analyses of morphology and [[ecology]].<ref name = strauch1985/> [[mitochondrial DNA|mtDNA]] [[cytochrome b|cytochrome ''b'']] [[Nucleic acid sequence|sequence]]s, and [[allozyme]] studies<ref name=Friesen/><ref name=Moum/> confirm these findings except that the ''[[Synthliboramphus]]'' murrelets should be split into a distinct tribe, as they appear more closely related to the Alcini; in any case, assumption of a closer relationship between the former and the true guillemots was only weakly supported by earlier studies.<ref name = strauch1985 />


Of the genera there are only a few species in each. This is probably a product of the rather small geographic range of the family (the most limited of any seabird family), and the periods of [[glacier|glacial]] advance and retreat that have kept the populations on the move in a narrow band of subarctic ocean.
Of the genera, only a few species are placed in each. This is probably a product of the rather small geographic range of the family (the most limited of any seabird family), and the periods of [[glacier|glacial]] advance and retreat that have kept the populations on the move in a narrow band of subarctic ocean.


Today, as in the past, the auks are restricted to cooler northern waters. Their ability to spread further south is restricted as their prey hunting method, pursuit diving, becomes less efficient in warmer waters. The speed at which small fish (which along with [[krill]] are the auk's principal food items) can swim doubles as the temperature increases from {{convert|5|to|15|C}}, with no corresponding increase in speed for the bird. The southernmost auks, in California and Mexico, can survive there because of cold [[upwelling]]s. The current paucity of auks in the Atlantic (6 species), compared to the Pacific (19–20 species) is considered to be because of extinctions to the Atlantic auks; the fossil record shows there were many more species in the Atlantic during the [[Pliocene]]. Auks also tend to be restricted to continental shelf waters and breed on few oceanic islands.
Today, as in the past, the auks are restricted to cooler northern waters. Their ability to spread further south is restricted as their prey hunting method, pursuit diving, becomes less efficient in warmer waters. The speed at which small fish (which along with krill are the auk's principal prey) can swim doubles as the temperature increases from {{convert|5|to|15|C}}, with no corresponding increase in speed for the bird. The southernmost auks, in California and Mexico, can survive there because of cold [[upwelling]]s. The current paucity of auks in the Atlantic (six species), compared to the Pacific (19–20 species) is considered to be because of extinctions to the Atlantic auks; the fossil record shows many more species were in the Atlantic during the Pliocene. Auks also tend to be restricted to continental-shelf waters and breed on few oceanic islands.


''Hydotherikornis oregonus'' (Described by Miller in 1931), the oldest purported alcid from the Eocene of California, is actually a petrel (as reviewed by Chandler in 1990) and is reassigned to the tubenoses (Procellariiformes). A 2003 paper entitled "The Earliest North American Record of Auk (Aves: Alcidae) From the Late Eocene of Central Georgia" by Robert M. Chandler and Dennis Parmley of Georgia College and State University reports a Late Eocene, wing-propelled diving, auk from the Priabonain Stage of the Late Eocene. These sediments have been dated through Chandronian NALMA {North American Land Mammal Age}, at an estimate of 34.5 to 35.5 million years on the Eocene time scale for fossil bearing sediments of the Clinchfield Formation, Gordon, Wilkinson County, Georgia. Furthermore, the sediments containing this unabraded portion of a left humerus (43.7&nbsp;mm long) are tropical or sub-tropical as evidenced by a wealth of warm water shark teeth, palaeophied snake vertebrae and turtles.
''Hydotherikornis oregonus'' (Described by Miller in 1931), the oldest purported alcid from the Eocene of California, is actually a petrel (as reviewed by Chandler in 1990) and is reassigned to the tubenoses (Procellariiformes). A 2003 paper, "The Earliest North American Record of Auk (Aves: Alcidae) From the Late Eocene of Central Georgia", <!-- by Robert M. Chandler and Dennis Parmley of Georgia College and State University --> reports a Late Eocene, wing-propelled, diving auk from the Priabonain stage of the Late Eocene. These sediments have been dated through Chandronian NALMA {North American Land Mammal Age}, at an estimate of 34.5 to 35.5 million years on the Eocene time scale for fossil-bearing sediments of the Clinchfield Formation, Gordon, Wilkinson County, Georgia. Furthermore, the sediments containing this unabraded portion of a left humerus (43.7&nbsp;mm long) are tropical or subtropical as evidenced by a wealth of warm-water shark teeth, palaeophied snake vertebrae, and turtles.


==Systematics==
==Systematics==
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|2=[[Crested auklet]] }} }} }} }} }}
|2=[[Crested auklet]] }} }} }} }} }}
|2={{clade
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|1={{clade
|1={{clade
|1={{clade
|1={{clade
|1=[[Ancient murrelet]]
|1=[[Guadalupe murrelet]]
|2=[[Japanese murrelet]]
|2=[[Scripps's murrelet]] }}
}}
|2=[[Craveri's murrelet]] }}
|2={{clade
|2={{clade
|1=[[Craveri's murrelet]]
|1=[[Japanese murrelet]]
|2={{clade
|2=[[Ancient murrelet]] }} }}
|1=[[Guadalupe murrelet]]
|2=[[Scripps's murrelet]]
}}
}}
}}
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|1={{clade
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**** [[Tufted puffin]], ''Fratercula cirrhata''
**** [[Tufted puffin]], ''Fratercula cirrhata''


[[Biodiversity]] of auks seems to have been markedly higher during the [[Pliocene]].<ref name = konyukhov2002 /> See the genus accounts for prehistoric species.
[[Biodiversity]] of auks seems to have been markedly higher during the Pliocene.<ref name = konyukhov2002 /> See the genus accounts for prehistoric species.


==See also==
==See also==
* [[Kiviak]], a traditional Inuit food from Greenland that is made of auks preserved in seal skin
* ''[[Kiviak]]'', a traditional Inuit food from Greenland that is made of auks preserved in seal skin
* [[Tradeoffs for locomotion in air and water]]
* [[Tradeoffs for locomotion in air and water]]


Line 171: Line 181:
<ref name =Friesen>{{cite journal |author1=Friesen, V.L. |author2=Baker, A.J. |author3=Piatt, J.F. |name-list-style=amp |year=1996 |title= Phylogenetic Relationships Within the Alcidae (Charadriiformes: Aves) Inferred from Total Molecular Evidence |journal=[[Molecular Biology and Evolution (journal)|Molecular Biology and Evolution]] |volume=13 |issue=2 |pages= 359–367 |doi=10.1093/oxfordjournals.molbev.a025595 |doi-access=free |pmid=8587501}}</ref>
<ref name =Friesen>{{cite journal |author1=Friesen, V.L. |author2=Baker, A.J. |author3=Piatt, J.F. |name-list-style=amp |year=1996 |title= Phylogenetic Relationships Within the Alcidae (Charadriiformes: Aves) Inferred from Total Molecular Evidence |journal=[[Molecular Biology and Evolution (journal)|Molecular Biology and Evolution]] |volume=13 |issue=2 |pages= 359–367 |doi=10.1093/oxfordjournals.molbev.a025595 |doi-access=free |pmid=8587501}}</ref>


<ref name = konyukhov2002>{{cite journal |author=Konyukhov, N.B. |year=2002 |title= Possible Ways of Spreading and Evolution of Alcids |journal=[[Biology Bulletin of the Russian Academy of Sciences]] |volume=29 |issue=5 |pages=447–454 |language=ru |doi=10.1023/A:1020457508769}}</ref>
<ref name = konyukhov2002>{{cite journal |author=Konyukhov, N.B. |year=2002 |title= Possible Ways of Spreading and Evolution of Alcids |journal=[[Biology Bulletin of the Russian Academy of Sciences]] |volume=29 |issue=5 |pages=447–454 |language=ru |doi=10.1023/A:1020457508769|bibcode=2002BioBu..29..447K |s2cid=36133751 }}</ref>


<ref name =Moum>{{cite journal |author1=Moum, Truls |author2=Arnason, Ulfur |author3=Árnason, Einar |name-list-style=amp |year=2002 |title= Mitochondrial DNA Sequence Evolution and Phylogeny of the Atlantic Alcidae, Including the Extinct Great Auk (''Pinguinus impennis'') |journal=[[Molecular Biology and Evolution]] |volume=19 |issue=9 |pages= 1434–1439 |doi=10.1093/oxfordjournals.molbev.a004206 |doi-access=free |pmid=12200471}}</ref>
<ref name =Moum>{{cite journal |author1=Moum, Truls |author2=Arnason, Ulfur |author3=Árnason, Einar |name-list-style=amp |year=2002 |title= Mitochondrial DNA Sequence Evolution and Phylogeny of the Atlantic Alcidae, Including the Extinct Great Auk (''Pinguinus impennis'') |journal=[[Molecular Biology and Evolution]] |volume=19 |issue=9 |pages= 1434–1439 |doi=10.1093/oxfordjournals.molbev.a004206 |doi-access=free |pmid=12200471}}</ref>
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<ref name = strauch1985>{{cite journal |author=Strauch J.G. Jr. |year=1985 |title= The phylogeny of the Alcidae |journal=[[The Auk]] |volume=102 |issue=3 |pages= 520–539 |url=http://sora.unm.edu/sites/default/files/journals/auk/v102n03/p0520-p0539.pdf | jstor = 4086647 |doi=10.1093/auk/102.3.520}}</ref>
<ref name = strauch1985>{{cite journal |author=Strauch J.G. Jr. |year=1985 |title= The phylogeny of the Alcidae |journal=[[The Auk]] |volume=102 |issue=3 |pages= 520–539 |url=http://sora.unm.edu/sites/default/files/journals/auk/v102n03/p0520-p0539.pdf | jstor = 4086647 |doi=10.1093/auk/102.3.520}}</ref>


<ref name=smith2011>{{cite journal |author=Smith, N.A. |year=2011 |title=Taxonomic revision and phylogenetic analysis of the flightless Mancallinae (Aves, Pan-Alcidae) |doi=10.3897/zookeys.91.709 |journal=[[ZooKeys]] |issue=91 |pmid=21594108 |pmc=3084493 |pages=1–116}}</ref>
<ref name=smith2011>{{cite journal |author=Smith, N.A. |year=2011 |title=Taxonomic revision and phylogenetic analysis of the flightless Mancallinae (Aves, Pan-Alcidae) |doi=10.3897/zookeys.91.709 |journal=[[ZooKeys]] |issue=91 |pmid=21594108 |pmc=3084493 |pages=1–116|bibcode=2011ZooK...91....1S |doi-access=free }}</ref>


<ref name=Thomas>{{cite journal |author1=Thomas, Gavin H. |author2=Wills, Matthew A. |author3=Székely, Tamás |name-list-style=amp |doi=10.1186/1471-2148-4-28 |pmid= 15329156 |pmc=515296 |year=2004 |journal=[[BMC Evolutionary Biology]] |volume=4 |pages=28 |title=A supertree approach to shorebird phylogeny |issue=1}} [https://www.ncbi.nlm.nih.gov/pmc/articles/PMC515296/#__sec15title Supplementary Material]</ref>
<ref name=Thomas>{{cite journal |author1=Thomas, Gavin H. |author2=Wills, Matthew A. |author3=Székely, Tamás |name-list-style=amp |doi=10.1186/1471-2148-4-28 |pmid= 15329156 |pmc=515296 |year=2004 |journal=[[BMC Evolutionary Biology]] |volume=4 |pages=28 |title=A supertree approach to shorebird phylogeny |issue=1 |doi-access=free }} [https://www.ncbi.nlm.nih.gov/pmc/articles/PMC515296/#__sec15title Supplementary Material]</ref>
}}
}}


Line 185: Line 195:
* {{cite journal | last1 = Collinson | first1 = Martin | year = 2006 | title = Splitting headaches? Recent taxonomic changes affecting the British and Western Palaearctic lists | url = http://www.britishbirds.co.uk/search?id=9283 | journal = [[British Birds (magazine)|British Birds]] | volume = 99 | issue = 6 | pages = 306–323 }}
* {{cite journal | last1 = Collinson | first1 = Martin | year = 2006 | title = Splitting headaches? Recent taxonomic changes affecting the British and Western Palaearctic lists | url = http://www.britishbirds.co.uk/search?id=9283 | journal = [[British Birds (magazine)|British Birds]] | volume = 99 | issue = 6 | pages = 306–323 }}
* {{cite book |last1=Gaston |first1=Anthony J. |last2=Jones |first2=Ian L. |title=The Auks : Alcidae |date=1998 |publisher=Oxford University Press |location=Oxford |isbn=0-19-854032-9}}
* {{cite book |last1=Gaston |first1=Anthony J. |last2=Jones |first2=Ian L. |title=The Auks : Alcidae |date=1998 |publisher=Oxford University Press |location=Oxford |isbn=0-19-854032-9}}
* {{cite journal | last1 = Paton | first1 = T.A. | last2 = Baker | first2 = A.J. | last3 = Groth | first3 = J.G. | last4 = Barrowclough | first4 = G.F. | year = 2003 | title = RAG-1 sequences resolve phylogenetic relationships within charadriiform birds | journal = [[Molecular Phylogenetics and Evolution]] | volume = 29 | issue = 2 | pages = 268–278 | doi = 10.1016/S1055-7903(03)00098-8 | pmid = 13678682 }}
* {{cite journal | last1 = Paton | first1 = T.A. | last2 = Baker | first2 = A.J. | last3 = Groth | first3 = J.G. | last4 = Barrowclough | first4 = G.F. | year = 2003 | title = RAG-1 sequences resolve phylogenetic relationships within charadriiform birds | journal = [[Molecular Phylogenetics and Evolution]] | volume = 29 | issue = 2 | pages = 268–278 | doi = 10.1016/S1055-7903(03)00098-8 | pmid = 13678682 | bibcode = 2003MolPE..29..268P }}
* [http://digitalcommons.unl.edu/bioscidivingbirds/ ''Diving Birds of North America''] by [[Paul Johnsgard]]
* [http://digitalcommons.unl.edu/bioscidivingbirds/ ''Diving Birds of North America''] by [[Paul Johnsgard]]


== External links ==
== External links ==
* {{Commons category-inline |Alcidae}}
* {{Commons category-inline |Alcidae}}
* {{cite EB9 |wstitle= Auk |volume= III |page=85 |short=1}}


{{Taxonbar|from=Q28294}}
{{Taxonbar|from=Q28294}}

Latest revision as of 07:33, 28 December 2024

This article is about a family of birds. For the ornithological journal, see The Auk. For other meanings, see AUK.
"Auklet" redirects here. For the fishery patrol vessel, see USFS Auklet.

Auks
Temporal range: Late Eocene – recent 35–0 Ma
Parakeet auklets (Aethia psittacula)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Charadriiformes
Suborder: Lari
Family: Alcidae
Leach, 1820
Type species
Alca torda
Subfamilies

Auks or alcids are birds of the family Alcidae in the order Charadriiformes.[1] The alcid family includes the murres, guillemots, auklets, puffins, and murrelets. The family contains 25 extant or recently extinct species that are divided into 11 genera. Auks are found throughout the Northern Hemisphere.[1][2]

Apart from the extinct great auk, all auks can fly, and are excellent swimmers and divers (appearing to "fly" in water), but their walking appears clumsy.

Names

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Several species have different English names in Europe and North America. The two species known as "murres" in North America are called "guillemots" in Europe, and the species called little auk in Europe is referred to as dovekie in North America.

Etymology

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The word "auk" /ɔːk/ is derived from Icelandic álka and Norwegian alka or alke from Old Norse ālka from Proto-Germanic *alkǭ (sea-bird, auk).[3][4]

Taxonomy

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The family name Alcidae comes from the genus Alca given by Carl Linnaeus in 1758 for the razorbill (Alca torda) from the Norwegian word alke.[5]

Description

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Auks are superficially similar to penguins, having black-and-white colours, upright posture, and some of their habits. Nevertheless, they are not closely related to penguins, but rather are believed to be an example of moderate convergent evolution. Auks are monomorphic (males and females are similar in appearance).

Extant auks range in size from the least auklet, at 85 g (3 oz) and 15 cm (5.9 in), to the thick-billed murre, at 1 kg (2.2 lb) and 45 cm (18 in). Due to their short wings, auks have to flap their wings very quickly to fly.

Although not to the extent of penguins, auks have largely sacrificed flight, and also mobility on land, in exchange for swimming ability; their wings are a compromise between the best possible design for diving and the bare minimum needed for flying. This varies by subfamily, with the Uria guillemots (including the razorbill) and murrelets being the most efficient under the water, whereas the puffins and auklets are better adapted for flying and walking.

Feeding and ecology

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The feeding behaviour of auks is often compared to that of penguins; both groups are wing-propelled, pursuit divers. In the region where auks live, their only seabird competition are cormorants (which are dive-powered by their strong feet). In areas where the two groups feed on the same prey, the auks tend to feed further offshore. Strong-swimming murres hunt faster, schooling fish, whereas auklets take slower-moving krill. Time depth recorders on auks have shown that they can dive as deep as 100 m (330 ft) in the case of Uria guillemots, 40 m (130 ft) for the Cepphus guillemots and 30 m (98 ft) for the auklets.

Breeding and colonies

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Auks are pelagic birds, spending the majority of their adult lives on the open sea and going ashore only for breeding, although some species, such as the common guillemot, spend a great part of the year defending their nesting spot from others.

Auks are monogamous, and tend to form lifelong pairs. They typically lay a single egg, and they use the nesting site year after year.

Some species, such as the Uria guillemots (murres), nest in large colonies on cliff edges; others, such as the Cepphus guillemots, breed in small groups on rocky coasts; and the puffins, auklets, and some murrelets nest in burrows. All species except the Brachyramphus murrelets are colonial.

Evolution and distribution

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See also: List of Charadriiformes by population
Auks as painted by Archibald Thorburn

Traditionally, the auks were believed to be one of the earliest distinct charadriiform lineages due to their characteristic morphology, but genetic analyses have demonstrated that these peculiarities are the product of strong natural selection, instead; as opposed to, for example, plovers (a much older charadriiform lineage), auks radically changed from a wading shorebird to a diving seabird lifestyle. Thus today, the auks are no longer separated in their own suborder (Alcae), but are considered part of the Lari suborder, which otherwise contains gulls and similar birds. Judging from genetic data, their closest living relatives appear to be the skuas, with these two lineages separating about 30 million years ago (Mya).[6][7][8] Alternatively, auks may have split off far earlier from the rest of the Lari and undergone strong morphological, but slow genetic evolution, which would require a very high evolutionary pressure, coupled with a long lifespan and slow reproduction.

The earliest unequivocal fossils of auks are from the late Eocene, some 35 Mya.[9] The genus Miocepphus, (from the Miocene, 15 Mya) is the earliest known from good specimens. Two very fragmentary fossils are often assigned to the Alcidae, although this may not be correct: Hydrotherikornis (Late Eocene) and Petralca (Late Oligocene). Most extant genera are known to exist since the Late Miocene or Early Pliocene (about 5 Mya). Miocene fossils have been found in both California and Maryland, but the greater diversity of fossils and tribes in the Pacific leads most scientists to conclude they first evolved there, and in the Miocene Pacific, the first fossils of extant genera are found. Early movement between the Pacific and the Atlantic probably happened to the south (since no northern opening to the Atlantic existed), with later movements across the Arctic Ocean.[10] The flightless subfamily Mancallinae, which was apparently restricted to the Pacific Coast of southern North America and became extinct in the Early Pleistocene, is sometimes included in the family Alcidae under some definitions. One species, Miomancalla howardae, is the largest charadriiform of all time.[11]

Razorbills are an auk found in the Atlantic Ocean.

The family contains 25 extant or recently extinct species that are divided into 11 genera.[2] The extant auks (subfamily Alcinae) are broken up into two main groups - the usually high-billed puffins (tribe Fraterculini) and auklets (tribe Aethiini), as opposed to the more slender-billed murres and true auks (tribe Alcini), and the murrelets and guillemots (tribes Brachyramphini and Cepphini). The tribal arrangement was originally based on analyses of morphology and ecology.[12] mtDNA cytochrome b sequences, and allozyme studies[6][7] confirm these findings except that the Synthliboramphus murrelets should be split into a distinct tribe, as they appear more closely related to the Alcini; in any case, assumption of a closer relationship between the former and the true guillemots was only weakly supported by earlier studies.[12]

Of the genera, only a few species are placed in each. This is probably a product of the rather small geographic range of the family (the most limited of any seabird family), and the periods of glacial advance and retreat that have kept the populations on the move in a narrow band of subarctic ocean.

Today, as in the past, the auks are restricted to cooler northern waters. Their ability to spread further south is restricted as their prey hunting method, pursuit diving, becomes less efficient in warmer waters. The speed at which small fish (which along with krill are the auk's principal prey) can swim doubles as the temperature increases from 5 to 15 °C (41 to 59 °F), with no corresponding increase in speed for the bird. The southernmost auks, in California and Mexico, can survive there because of cold upwellings. The current paucity of auks in the Atlantic (six species), compared to the Pacific (19–20 species) is considered to be because of extinctions to the Atlantic auks; the fossil record shows many more species were in the Atlantic during the Pliocene. Auks also tend to be restricted to continental-shelf waters and breed on few oceanic islands.

Hydotherikornis oregonus (Described by Miller in 1931), the oldest purported alcid from the Eocene of California, is actually a petrel (as reviewed by Chandler in 1990) and is reassigned to the tubenoses (Procellariiformes). A 2003 paper, "The Earliest North American Record of Auk (Aves: Alcidae) From the Late Eocene of Central Georgia", reports a Late Eocene, wing-propelled, diving auk from the Priabonain stage of the Late Eocene. These sediments have been dated through Chandronian NALMA {North American Land Mammal Age}, at an estimate of 34.5 to 35.5 million years on the Eocene time scale for fossil-bearing sediments of the Clinchfield Formation, Gordon, Wilkinson County, Georgia. Furthermore, the sediments containing this unabraded portion of a left humerus (43.7 mm long) are tropical or subtropical as evidenced by a wealth of warm-water shark teeth, palaeophied snake vertebrae, and turtles.

Systematics

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Alcidae
Cladogram of the Alcidae family[9]

Biodiversity of auks seems to have been markedly higher during the Pliocene.[10] See the genus accounts for prehistoric species.

See also

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References

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  1. ^ a b Myers, P.; Espinosa, R.; Parr, C. S.; Jones, T.; Hammond, G. S.; Dewey, T. A. (2022). "Alcidae". Animal Diversity Web. University of Michigan Museum of Zoology. Retrieved 31 December 2022.
  2. ^ a b Gill, Frank; Donsker, David; Rasmussen, Pamela, eds. (August 2022). "Noddies, gulls, terns, skimmers, skuas, auks". IOC World Bird List Version 12.2. International Ornithologists' Union. Retrieved 10 November 2022.
  3. ^ "Definition of Auk". Merriam-Webster.
  4. ^ "Definition of auk". Dictionary.com.
  5. ^ Jobling, James A. (2010). "Alca" (PDF). The Helm Dictionary of Scientific Bird Names. London, United Kingdom: Christopher Helm. p. 40. ISBN 978-1-4081-2501-4.
  6. ^ a b Friesen, V.L.; Baker, A.J. & Piatt, J.F. (1996). "Phylogenetic Relationships Within the Alcidae (Charadriiformes: Aves) Inferred from Total Molecular Evidence". Molecular Biology and Evolution. 13 (2): 359–367. doi:10.1093/oxfordjournals.molbev.a025595. PMID 8587501.
  7. ^ a b Moum, Truls; Arnason, Ulfur & Árnason, Einar (2002). "Mitochondrial DNA Sequence Evolution and Phylogeny of the Atlantic Alcidae, Including the Extinct Great Auk (Pinguinus impennis)". Molecular Biology and Evolution. 19 (9): 1434–1439. doi:10.1093/oxfordjournals.molbev.a004206. PMID 12200471.
  8. ^ Thomas, Gavin H.; Wills, Matthew A. & Székely, Tamás (2004). "A supertree approach to shorebird phylogeny". BMC Evolutionary Biology. 4 (1): 28. doi:10.1186/1471-2148-4-28. PMC 515296. PMID 15329156. Supplementary Material
  9. ^ a b Smith, N.A. (2011). "Taxonomic revision and phylogenetic analysis of the flightless Mancallinae (Aves, Pan-Alcidae)". ZooKeys (91): 1–116. Bibcode:2011ZooK...91....1S. doi:10.3897/zookeys.91.709. PMC 3084493. PMID 21594108.
  10. ^ a b Konyukhov, N.B. (2002). "Possible Ways of Spreading and Evolution of Alcids". Biology Bulletin of the Russian Academy of Sciences (in Russian). 29 (5): 447–454. Bibcode:2002BioBu..29..447K. doi:10.1023/A:1020457508769. S2CID 36133751.
  11. ^ Smith, N. A. (2015). "Evolution of body mass in the Pan-Alcidae (Aves, Charadriiformes): the effects of combining neontological and paleontological data". Paleobiology. 42 (1): 8–26. doi:10.1017/pab.2015.24. S2CID 83934750.
  12. ^ a b Strauch J.G. Jr. (1985). "The phylogeny of the Alcidae" (PDF). The Auk. 102 (3): 520–539. doi:10.1093/auk/102.3.520. JSTOR 4086647.

Further reading

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