FOOTNOTES:
[41] Common eider, 2 to 7 h.
[42] No data given.
[43] Indirect evidence that young are brooded this long.
[44] Data from Lack (1968) and Ashmole (1971) unless otherwise noted.
[45] Cullen (1957).
[46] Birkhead (1974).
[47] Speich and Manuwal (1974).
[Zoogeography and Taxonomic Relationships of Seabirds in Northern North America]
by
M. D. F. Udvardy
California State University
Sacramento, California 95819
Abstract
The zoogeography and taxonomic relationships among 42 living and 1 extinct species of marine birds from the northern and northwestern coasts of North America are described. Seventeen species are circumpolar in distribution; 17 are endemic to Beringia, and 8 have origins in the North Pacific.
This discussion concerns the northern and western coasts of the continent, from about the Mackenzie Delta westward and southward to the mouth of the Columbia River. Besides bona fide seabirds, I include marine birds that predominantly breed and feed on or around the marine littoral, but exclude two groups: shorebirds, jaegers, and phalaropes, which breed inland and move out from the Arctic after an undetermined postbreeding period; and Anseriformes which become "marine birds" in their southern winter quarters. What remains is 42 living species (Table 1).
The Procellariiformes, or tube-nosed seabirds, have a predominantly southern hemispheric, Gondwanan distribution. The North Pacific basin is an important feeding ground of several shearwaters (Puffinus spp.) that breed in the South Pacific and subantarctic. Only three species breed in the area under consideration: the fulmar (Fulmarus glacialis) and two storm-petrels (Oceanodroma spp.), all of which are still relatively widespread.
Of the Pelecaniformes, the very successful, worldwide cormorants (Phalacrocorax spp.)—inland water as well as coastal and "amphibious" species are on every continent—are ancient Pacific dwellers, with a high grade of endemism here: Of the two subarctic species, one (P. perspicillatus) became extinct long ago, and the other, the red-faced cormorant (P. urile), is very restricted, and deserves our greatest attention. The pelagic cormorant (P. pelagicus), Brandt's cormorant (P. penicillatus), and the double-crested cormorant (P. auritus) are widespread and successful, extending south of the area here considered; double-crested cormorants also breed inland and across toward the North Atlantic coast. As fish-eaters they are often persecuted where coastal fishermen possess firearms, and thus are sensitive to increasing human influence on the coasts.
Two species of arctic geese need special attention. The emperor goose (Philacte canagica) is a Beringean endemic and lives in a very restricted area of both sides of this sea; its status (endangered?) is unknown to me. Since the black brant (Branta bernicla) is a long-range migrant, it is hunted as a game bird at its winter grounds, and subject to management measures. Whereas the emperor goose is a unique offshoot of the genus Anser, the Pacific brant is considered a subspecies; its general distribution is circumpolar.
Five arctic ducks, and one other, constitute the sea ducks of the area. The common eider (Somateria mollissima), king eider (S. spectabilis), and the oldsquaw (Clangula hyemalis) are widespread, and circumpolar or nearly so; hunting and down-robbing in other parts of the Arctic may provide clues as to their relative tolerance of primitive or advanced civilization. The spectacled eider (S. fischeri) and Steller's eider (Polysticta stelleri) are restricted to the Bering Sea coasts and neighboring High Arctic coasts, respectively; their status is precarious.
| Species | Distribution | ||||||
|---|---|---|---|---|---|---|---|
| Circumpolar | Widespread in North Pacific | North coast of Alaska | Beringia[48] | Aleutian Islands | South coast of Alaska[49] | Temperate northeast Pacific coast[50] | |
| Fulmarus glacialis | x | w | x | x | x | w | |
| Oceanodroma furcata | x | x | x | x | |||
| O. leucorhoa | x | x | x | x | x | ||
| Phalacrocorax auritus | x | x | x | ||||
| P. penicillatus | (x) | x | |||||
| P. pelagicus | x | x | x | x | x | ||
| P. urile | x | x | x | ||||
| Branta bernicla | x | x | x | (w) | (w) | w | |
| Anser canagicus | x | w | w | ||||
| Clangula hyemalis | x | w | x | x | w | w | w |
| Histrionicus histrionicus | x | w | w | w | w | w | |
| Polysticta stelleri | x | x | (w) | ||||
| Somateria mollissima | x | x | x | x | x | ||
| S. spectabilis | x | x | x | w | (w) | ||
| S. fischeri | x | x | |||||
| Larus hyperboreus | x | w | x | x | w | w | w |
| L. glaucescens | x | x | x | x | |||
| L. occidentalis | (x) | ||||||
| L. argentatus | x | w | x | (x)w | |||
| L. thayeri | w | x | w | w | w | w | |
| L. canus | x | w | x | x | (x)w | ||
| Rissa tridactyla | x | w | x | x | x | x | w |
| R. brevirostris | x | x | (x) | ||||
| Xema sabini | x | x | x | w | |||
| Sterna paradisaea | x | w | x | x | x | w | |
| S. aleutica | x | x | |||||
| Uria aalge | x | x | (x) | x | x | x | x |
| U. lomvia | x | x | x | x | x | x | |
| Alle alle | x | * | |||||
| Cepphus grylle | x | x | w | ||||
| C. columba | x | x | x | x | x | ||
| Brachyramphus marmoratus | x | (x) | x | x | |||
| B. brevirostris | x | x | x | ||||
| Synthliboramphus antiquus | x | x | x | x | x | ||
| Ptychoramphus aleuticus | x | x | x | ||||
| Cyclorrhynchus psittacula | x | x | |||||
| Aethia cristatella | x | x | |||||
| A. pusilla | x | x | |||||
| A. pygmaea | x | ||||||
| Cerorhinca monocerata | x | x | x | ||||
| Fratercula corniculata | x | x | x | x | x | ||
| Lunda cirrhata | x | x | x | x | x | ||
| Total number of nesting species | 17 | 11 | 15 | 27 | 25 | 24 | 17 |
| Total number of wintering species | 9 | 4 | 7 | 9 | 9 | ||
| Grand total | 17 | 20 | 15 | 31 | 32 | 33 | 26 |
The harlequin duck (Histrionicus histrionicus) stands alone without close relatives. It often breeds far from the sea, but spends the shortest time—only a few weeks—away from the rocky coast. There is a year-round population of yearlings in the sea. The drakes of the nearest breeding pairs at lower latitudes are back to the sea, abandoning their mates at the breeding stream when the alpine stream-dwellers are still at sea awaiting the thawing of their breeding grounds. Harlequin ducks live in large parts of Siberia, from arctic Alaska to central California and Colorado, and also in the eastern Arctic. They do not seem to me to be in immediate danger globally, though perhaps they are locally.
Gulls are a highly successful group of seabirds, and of the eight species on our coasts the four more southern ones—the western gull (Larus occidentalis), glaucous-winged gull (L. glaucescens), common gull (L. canus), and herring gull (L. argentatus)—are expanding wherever civilization creates new scavenging opportunities. Nothing is said about the populations of the kittiwake (Rissa tridactyla), black-legged kittiwake (R. brevirostris) and Sabine's gull (Xema sabini), or of the other two high arctic species (Pagophila eburnea, Rhodostethia rosea) which do not nest regularly in the area considered here.
The arctic tern (Sterna paradisaea) is circumpolarly widespread and successful, whereas the Aleutian tern (S. aleutica) is a very restricted Beringean endemic, and its status needs to be exactly known.
Almost one-third of the seabirds in this area are alcids, a family centered in the North Pacific and, more specifically, in the Bering Sea. Most species breed in enormous rookeries. Any impact of civilization is highly detrimental under such circumstances. Of the four circumpolar species the two Uria guillemots (murres) are important. The dovekie (Alle alle) is a sparse pioneer of Bering Strait, as is the black guillemot (Cepphus grylle) on our side of the Arctic Sea. Its congener, the pigeon guillemot (C. columba), is common and successful all the way to coastal central California. Of the remaining 11 species, special attention should be paid to the whiskered auklet (Aethia pygmaea) of the Aleutian chain; the Kittlitz's murrelet (Brachyramphus brevirostris) of the eastern Beringean and southern Alaska coast; and to the widespread, but very sporadic rhinoceros auklet, or puffin (Cerorhinca monocerata).
To sum up, I have tabulated these 42 species, and indicated whether modern life-history and population studies are extant:
| No. species | No. studied | |
|---|---|---|
| Procellariiformes | 3 | 2 |
| Phalacrocorax | 4 | 2 |
| Anseres | 2 | 1 |
| Anates | 6 | — |
| Lari | 9 | 2 |
| Sterni | 2 | — |
| Alcidae | 16 | 7 |
| Total | 42 | 14 |
Thus, 28 species await studies preliminary to, and highly necessary for, conservation measures.
Seventeen species of marine birds are spread either circumpolarly around the northern perimeter or along the north-south coasts of the Laurasian continents. Four of these are of the High Arctic (Branta bernicla, Somateria spectabilis, Xema sabini, Alle alle); another seven penetrate the Bering Sea as well (Fulmarus glacialis, Somateria mollissima, Clangula hyemalis, Larus hyperboreus, Rissa tridactyla, Sterna paradisaea, Uria lomvia); and six are panboreal-subboreal, widespread in their distribution—Oceanodroma leucorhoa (extends far south), Histrionicus histrionicus, Larus argentatus (widespread latitudinally), L. canus (also inland), Uria aalge, and Cepphus grylle.
Seventeen species of marine birds are endemic to Beringia: Anser canagicus, Polysticta stelleri, Somateria fischeri, Rissa brevirostris, and Aethia pusilla (and the extinct Phalacrocorax perspicillatus); P. urile, Sterna aleutica, Aethia pygmaea, A. cristatella, and Cyclorrhynchus extend westward to the Sea of Okhotsk, as do Brachyramphus brevirostris and Larus glaucescens, which also extend eastward; and Phalacrocorax pelagicus, Cepphus columba, Fratercula corniculata, and Lunda cirrhata are amphipacific species in Beringia.
Eight species of marine birds are associated with the North Pacific. Four are found on both sides of the ocean—Oceanodroma furcata, Brachyramphus marmoratus, Synthliboramphus antiquus, and Cerorhinca monocerata (very disjunct). The four others occur on only the North American side—Phalacrocorax auritus (also inland), P. penicillatus, Larus occidentalis (albeit barely), and Ptychoramphus aleuticus.
Finally, one species, Larus thayeri, is endemic at the central Canadian Arctic, extending westward into the area here considered.