In 1868 Elliott Coues published A Monograph of the Alcidae which surveyed the literature on the Auk family from the mid 19th century. 141 His work is a taxonomic treatise valuable for tracing the names given to the various alcids as they were discovered during exploration of northern seas. In 1919, A.C. Bent compiled the literature up to that date in his volume Life Histories of North American Diving Birds. 64

Birds of the Soviet Union by Dement'ev and Gladkov published in 1951, is an excellent reference book on auks and has been translated. 154 In 1957, E.V. Kozlova also summarized what was known of the auk family.339 This work, originally published in Russian, has been translated into English and is a basic reference on this family of birds. Since then much has been learned and in 1985, David Nettleship and Tim Birkhead edited a most important volume entitled The Atlantic Alcidae, the first comprehensive book on the biology and ecology of the species of the northern Atlantic.436 Ronald Freethy summarized information especially on the Atlantic species in 1987, 195 and in 1990 several Russian specialists on marine birds published a volume that is especially helpful to readers of the Russian language.

To many people a seabird is synonymous with a gull. But the alcids are not gulls and, according to early assumptions and recent biochemical evidence, the family may be more closely related to penguins than to gulls.225. If this is true, then perhaps the alcids should be removed from the order Charadriiformes and placed in a suborder of the Spheniciformes. To date comparative chromosome studies have not influenced any change in present classification.47 However, it is not the purpose of this book to revise taxonomy and we shall not dwell upon this uncertain point. This book is concerned with promoting a general knowledge of the life and ecology of each species that compose the family. Each one is fascinating, curious, beautiful in its own way, and each forms an important fiber in the web of life which is intricately woven, each strand being dependent upon all the others for its existence and support.

What is an auk? What are the characteristics which set the alcids aside as a particular family? Elliott Coues presents what he calls "an expression of a perfect diagnosis" with this description: "Auks are brachypterous, brachyurous, tridactylous natatores with lateral nostrils." 141 The professional will know what this "perfect diagnosis" means, but for the rest of my readers I will translate his definition into simpler terms. Auks are short-winged, short-tailed, three-toed marine birds, with nostrils at the sides of their bill. This definition sets the family Alcidae apart from all other seabirds except perhaps the penguins, which have extra-short wings and hind limbs located more posteriorly than the alcids. But the Great Auk was very penguin-like and, indeed, was formerly called a penguin. Presently, all penguins live in the southern hemisphere and all alcids live in the northern hemisphere. Both groups use their wings literally 'flying' underwater, a habit shared by Diving Petrels, some ducks and, to a lesser extent by shearwaters. The webbed feet are used mainly as rudders for steering and as paddles in surface maneuvers.

Alcids are gregarious birds, especially during breeding when most species establish large colonies. This colonizing is itself apparently a breeding stimulus. There is, however, a range from solitary nesting habits of the murrelets to the compact, dense masses formed by the murres which often nest shoulder-to-shoulder with as many as two pairs to each 30 centimeters square.

Auks breed on rocky prominences, on offshore rocky stacks, and on islands usually remote from man and predator. Some nest on the surface or in cracks between rocks, while other require deep-sodded soils in which to burrow. Still others fly inland to find refuge between mountain snowfields or in hidden nooks on the branches of moss-laden trees. Piles of driftwood sometimes provide nesting places for auklets and guillemots, and puffins have been seen to nest in the recesses of rusting hulls of shipwrecks.

The food habits of auks categorize them into three groups: plankton-feeders, fish-feeders, or feeders on both fish and plankton.54 Dovekies eat a variety of planktonic forms especially copepods and euphausiids. Cassin's Auklets depend mainly on euphausiids and amphipods. Crested, Whiskered and Least Auklets also feed upon these organisms. These foods are also among the main items consumed by the large plankton-straining whales. In fact, collectively, these organisms are often referred to as "whale-krill."

Most of the other auk species are primarily fish feeders, although puffins are known to take in plankton as well as fish, and the Parakeet Auklet feeds upon a variety of organisms from planktonic forms to marine worms and squid. Less is known about feeding habits of the murrelets but small fish and larval forms apparently make up the bulk of their diet. In fact, the dietary habits of auks varies with the season and they tend to prey upon the organisms which are most available to them at the time. The breeding season brings greater demands, and success in rearing young is critically dependent upon high levels of suitable food items.

Variation of size and structure among the alcids are related to the food eaten and are factors that keep the species separate.55 However, it is difficult to separate a number of species of auks on the basis of food eaten and in the structure of the bill as a food-catching device. Furthermore, studies of food habits presently known are fragmentary in scope often covering a small geographical area and season.

Along with other differences in body morphology, seasonal modification of the bill structure and the color of the mouth lining, play an important role as social releasing mechanisms in mating activities. In other words, like humans who use their lips, auks use their bills for sexual and other social communication. Such activity includes gape display, head-flicking, waggling bills together, nebbing neck feathers, and snapping and clicking of the mandibles in various rhythmic patterns. Body size, accessory plumage decorations, color patterns, facial markings and vocalizations are all important factors that tend to keep the various species separate. Twenty-one of the twenty-three known species of auks are listed by the American Ornithological Union's Check List of North American Birds. 8 Two additional species not listed for North America range in Japanese waters.

Following the nomenclature recommended by the A.O.U. (1983)8 and J.G. Strauch (1985)586 these 23 species are divided into seven tribes. Although it is not the intent of this book to distinguish between the various subspecies and since many of these suggested by authorities are difficult to recognize some races are listed for convenience.60

Family Alcidae
Tribe Alcini
Great Auk
(extinct since 1844) Alca impennis Linnaeus

Razorbill Alca torda torda Linnaeus
Alca torda pica Salomonsen
Alca torda islandica Salomonsen

Thin-billed or
Common Murre Uria aalge aalge (Pontoppidan)
Uria aalge californica (Bryant)
Uria aalge inornata Salomonsen

Thick-billed Murre Uria lomvia lomvia (Linnaeus)
Uria lomvia arra (Pallas)

Dovekie Alle alle alle (Linnaeus)
Alle Alle polaris (Stenhouse)


Tribe Cepphini

Black Guillemot Cepphus grylle atlantis Salomonsen
Cepphus grylle arcticus (Brehm)
Cepphus grylle ultimus Salomonsen

Cepphus grylle manditii (Mandt)

Pigeon Guillemot Cepphus columba columba Pallas
Cepphus columba eureka Storer
Cepphus columba kiaurka Portenko
Cepphus columba adianta Storer
Cepphus columba snowi Stejneger


Sooty or
Spectacled Guillemot Cepphus carbo Pallas

Xantus' Murrelet Synthliboramphus hypoleuca hypoleuca (Xantus)
Synthliboramphus hypoleuca scrippsi Green and Arnold

Craveri's Murrelet Synthliboramphus craveri (Salvadori)

Ancient Murrelet Synthliboramphus antiquus

(Gmelin) Japanese Murrelet Synthliboramphus wumizusume (Temminck)

Tribe Brachyramphini

Marbled Murrelet Brachyramphus marmoratus (Gmelin)

Kittlitz's Murrelet Brachyramphus brevirostris (Vigors)

Tribe Aethiini

Cassin's Auklet Ptychoramphus aleuticus aleuticus (Pallas)
Ptychoramphus aleuticus australis van Rossem

Parakeet Auklet Aethia psittacula (Pallas)

Crested Auklet Aethia cristatella (Pallas)

Least Auklet Aethia pusilla (Pallas)

Whiskered Auklet Aethia pygmaea (Gmelin)

Tribe Fraterculini

Rhinoceros Auklet Cerorhinca monocerata (Pallas)

Atlantic Puffin Fratercula arctica arctica (Linnaeus)
Fratercula arctica naumanni (Norton)
Fratercula arctica grabae (Brehm)

Horned Puffin Fratercula corniculata (Naumann)

Tufted Puffin Lunda cirrhata (Pallas)