Alcidae

Life-forms

Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Cladus: Craniata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Archosauromorpha
Cladus: Crurotarsi
Divisio: Archosauria
Subsectio: Ornithodira
Subtaxon: Dinosauromorpha
Cladus: Dinosauria
Ordo: Saurischia
Cladus: Theropoda
Cladus: Neotheropoda
Infraclassis: Aves
Cladus: Euavialae
Cladus: Avebrevicauda
Cladus: Pygostylia
Cladus: Ornithothoraces
Cladus: Euornithes
Cladus: Ornithuromorpha
Cladus: Ornithurae
Cladus: Carinatae
Parvclassis: Neornithes
Cohors: Neognathae
Ordo: Charadriiformes
Subordo: Alcae

Familia: Alcidae
Genera: Aethia - Alca - Alle - Brachyramphus - Cepphus - Cerorhinca - Fratercula – Ptychoramphus – Synthliboramphus - Uria

Genera extincta: †Pinguinus
Name

Alcidae Leach, 1820
Typus

Alca Linnaeus, 1758

References

Synopsis contents Brit. Mus. (ed. 17): 68
Smith, N. A. 2011. Taxonomic revision and phylogenetic analysis of the flightless Mancallinae (Aves, Pan-Alcidae). ZooKeys 91: 1–116. DOI: 10.3897/zookeys.91.709

Vernacular names
dansk: Alkefugl
Deutsch: Alkenvögel
English: Auks
Nordfriisk: Alken
Frysk: Alkfûgels
norsk: Alkefugler
Türkçe: Dalıcımartıgiller

An auk or alcid is a bird of the family Alcidae in the order Charadriiformes. The alcid family includes the murres, guillemots, auklets, puffins, and murrelets. The word auk /ɔːk/ is derived from Icelandic álka, from Old Norse alka (“auk”), from Proto-Germanic *alkǭ (“sea-bird, auk”).[1][2]

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.

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

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 in order 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, 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

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 feet) 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

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 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 philopatric (they use the nesting site year after year).

Some species, such as the Uria guillemots (murres), nest in large colonies on cliff edges; others, like 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
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. However, 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).[3][4][5] 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.[6] 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 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.[7] 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.[8]

Razorbills are true auks only 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.[9] mtDNA cytochrome b sequence and allozyme studies[3][4] 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.[9]

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 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 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 (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.

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 mm long) are tropical or sub-tropical as evidenced by a wealth of warm water shark teeth, palaeophied snake vertebrae and turtles.
Systematics

Alcidae

Rhinoceros auklet

Tufted puffin

	Horned puffin

	Atlantic puffin

Cassin's auklet

Least auklet

Parakeet auklet

	Whiskered auklet

	Crested auklet

	Guadalupe murrelet

	Scripps's murrelet

Craveri's murrelet

	Japanese murrelet

	Ancient murrelet

Long-billed murrelet

	Marbled murrelet

	Kittlitz's murrelet

Black guillemot

	Spectacled guillemot

	Pigeon guillemot

	Thick-billed murre

	Common murre

Little auk

	†Great auk

	Razorbill

 
 
 
Cladogram of the Alcidae family[6]

Basal and incertae sedis
Miocepphus (fossil: Middle Miocene of CE USA)
Miocepphus mcclungi Wetmore, 1940
Miocepphus bohaskai Wijnker and Olson, 2009
Miocepphus blowi Wijnker and Olson, 2009
Miocepphus mergulellus Wijnker and Olson, 2009
Subfamily Alcinae
Tribe Alcini – typical auks and murres

The synthliboramphine Xantus's murrelet (Synthliboramphus hypoleucus) is quite distinct from the brachyramphine murrelets.

Uria
Common murre or common guillemot, Uria aalge
Thick-billed murre or, Brünnich's guillemot, Uria lomvia
Alle
Little auk or dovekie, Alle alle
Pinguinus
Great auk, Pinguinus impennis (extinct, c.1844)
Alca
Razorbill, Alca torda

Tribe Synthliboramphini – synthliboramphine murrelets
Black guillemot (Cepphus grylle, a true guillemot) in summer (front) and winter plumage

Synthliboramphus
Scripps's murrelet, Synthliboramphus scrippsi – formerly in S. hypoleucus ("Xantus's murrelet")
Guadalupe murrelet, Synthliboramphus hypoleucus – sometimes separated in Endomychura
Craveri's murrelet, Synthliboramphus craveri – sometimes separated in Endomychura
Ancient murrelet, Synthliboramphus antiquus
Japanese murrelet, Synthliboramphus wumizusume

Tribe Cepphini – true guillemots

Marbled murrelet (Brachyramphus marmoratus, a brachyramphine murrelet) in breeding plumage
Cepphus
Black guillemot or tystie, Cepphus grylle
Pigeon guillemot, Cepphus columba
Kurile guillemot, Cepphus columba snowi
Spectacled guillemot, Cepphus carbo
Tribe Brachyramphini – brachyramphine murrelets
Brachyramphus
Marbled murrelet, Brachyramphus marmoratus
Long-billed murrelet, Brachyramphus perdix
Kittlitz's murrelet, Brachyramphus brevirostris

Subfamily Fraterculinae

Tribe Aethiini – auklets

Tufted puffin (Fratercula cirrhata)
Ptychoramphus
Cassin's auklet, Ptychoramphus aleuticus
Aethia
Parakeet auklet, Aethia psittacula
Crested auklet, Aethia cristatella
Whiskered auklet, Aethia pygmaea
Least auklet, Aethia pusilla
Tribe Fraterculini – puffins
Cerorhinca
Rhinoceros auklet, Cerorhinca monocerata
Fratercula
Atlantic puffin, Fratercula arctica
Horned puffin, Fratercula corniculata
Tufted puffin, Fratercula cirrhata

Biodiversity of auks seems to have been markedly higher during the Pliocene.[7] See the genus accounts for prehistoric species.
See also

Kiviak, a traditional Inuit food from Greenland that is made of auks preserved in seal skin
Tradeoffs for locomotion in air and water

References

"Definition of AUK". www.merriam-webster.com.
"Definition of auk | Dictionary.com". www.dictionary.com.
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.
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.
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
Smith, N.A. (2011). "Taxonomic revision and phylogenetic analysis of the flightless Mancallinae (Aves, Pan-Alcidae)". ZooKeys (91): 1–116. doi:10.3897/zookeys.91.709. PMC 3084493. PMID 21594108.
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. doi:10.1023/A:1020457508769.
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.

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.