Auk

An auk is a bird of the family Alcidae in the order Charadriiformes. Auks are superficially similar to penguins due to their black-and-white colours, their 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.

Unlike penguins, modern auks can fly (except for the recently extinct Great Auk). They are good swimmers and divers, but their walking appears clumsy. Due to their short wings, auks have to flap their wings very quickly in order to fly. Extant auks range in size from the Least Auklet, at 85 g (3 oz) and 15 cm (6 in), to the Thick-billed Murre, at 1 kg (2¼ lb) and 45 cm (18 in).

Auks live on the open sea and only go ashore for breeding, although some species, like the Common Guillemot, spend a great part of the year defending their nesting spot from others.

Several species have different names in Europe and North America. The guillemots of Europe are murres in North America, if they occur in both continents, and the Little Auk becomes the Dovekie.

Some species, such as the Uria guillemots, 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.

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 is the cormorants (which dive powered by their strong feet); in areas where the two groups feed on the same prey, the auks tend to feed further offshore.

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. This reflects the type of prey taken: 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 in the case of Uria guillemots, 40 m for the Cepphus guillemots and 30 m for the auklets.

Evolution and distribution
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). 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. 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. 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.

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. mtDNA cytochrome b sequence and allozyme studies 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.

Compared to other families of seabirds, there are no genera with many species (such as the 47 Larus gulls). 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°C 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 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 unabraided portion of a left humerus (43.7mm long) are tropical or sub-tropical as evidenced by a wealth of warm water shark teeth, palaeophied snake vertebrae and turtles.

Systematics
Family Alcidae (=Pan-Alcidae)
 * Basal and incertae sedis
 * Alcodes (fossil: Late Miocene of Orange County, USA)
 * Hydrotherikornis (fossil: Late Eocene of Oregon, USA) – disputed
 * "Mancalla" diegensis (fossil: Middle Pliocene of California, USA)
 * "Mancalla" milleri (fossil)
 * "Mancalla" emlongi (fossil)
 * Pseudocepphus (fossil: Middle - Late Miocene) – basal Alcinae?
 * Subfamily Petralcinae (fossil) – disputed
 * Petralca (Early ?- Late Oligocene of Austria)
 * Subfamily Mancallinae (fossil)Tordalke01.jpg (Alca torda, rearing up) and Common Guillemots or Murres (Uria aalge) are Alcini]]
 * Praemancalla (Late Miocene - Early Pliocene of Orange County, USA)
 * Mancalla (Late Miocene - Early Pleistocene of W North America)
 * Miomancalla
 * Family Alcidae (crown group)
 * Basal and incertae sedis
 * Miocepphus (fossil: Middle Miocene of CE USA)
 * Subfamily Alcinae
 * Tribe Alcini – typical auks and murresXantus adult.gif (Synthliboramphus hypoleucus) is quite distinct from the brachyramphine murrelets]]
 * Uria
 * Common Guillemot or Common Murre, Uria aalge
 * Brunnich's Guillemot or Thick-billed Murre, Uria lomvia
 * Little Auk or Dovekie, Alle alle
 * Great Auk, Pinguinus impennis (extinct, c.1844)
 * Razorbill, Alca torda
 * Tribe Synthliboramphini – synthliboramphine murreletsBlackguillemot44.jpg (Cepphus grylle, a true guillemot) in summer (front) and winter plumage]]
 * Synthliboramphus
 * Xantus's 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 guillemotsMarbled murrelet breeding plumage.jpg (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 – aukletsPuffin Mrkoww.jpg (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
 * 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. See the genus accounts for prehistoric species.