part I

have not found an indication of a major perturbation over the past 40 years. Before 1936, the record is too fragmentary to allow an assessment.

One of the interesting aspects of kittiwake ecology in the Gulf of Alaska is the common occurrence of breeding failure. David Snarski (December 1943 Quarterly Progress Report, Alaska Cooperative Wildlife Research Unit, University of Alaska) observed breeding failure on colonies in the Tuxedni National Wildlife Refuge in 1970 and 1971 and obtained circumstantial evidence of another failure in 1972. In 1973 all of the breeding cliffs were occupied and nesting was successful. Whatever the cause of these periodic failures, they do not yet appear to have had a permanent impact that we are able to measure.

Red-legged Kittiwake _(Rissa brevirostris)_

Red-legged kittiwakes are not now known to breed in the western Gulf of Alaska. Turner (1886) stated that he saw a few at Sanak in 1878. We also have Stejneger's (1885) statement, that "red-legged" kittiwakes nest from Bering Island to Sanak. Friedmann (1937) reported two humeri from Kodiak Island middens. During the summer of 1976, two birds were observed off Kodiak Island by Irving M. Warner (personal communication), and one at 158°W and 54°30'-54°20'N south and east of the Shumagin Islands (Patrick J. Gould, personal communication).

Turner (1885) listed the red-legged kittiwake as abundant and breeding in the Near Islands. Turner (1886) also stated that he had seen quite a number about a cliff back of the village on Akutan Island in 1878. He added that to the westward this kittiwake was more abundant than the black-legged kittiwake. Murie (1959) expressed the opinion that Turner had confused the short-billed gull with the "short-billed" kittiwake. Clark (1911) also reported that he had seen the red-legged kittiwake in small numbers near Unalaska and that they became progressively more common west to the Near Islands. Nelson (1887) reported seeing large numbers of red-legged kittiwakes at Unalaska. Murie (1959) and Gabrielson (1940, 1944) did not see any red-legged kittiwakes in the Aleutian Islands. The species has recently been discovered breeding at Buldir and Bogoslof islands (G. Vernon Byrd, personal communication).

Is it possible that we have here another species which is exhibiting a response to some unknown long-term perturbation? The suggestion that such an event has occurred is faint, but it is there. Do we have in the red-legged and black-legged kittiwakes an example of yet another congener pair that has been affected by some perturbation in which one was affected positively and the other negatively? Clark (1911) reported small numbers of black-legged kittiwakes to go with large numbers of red-legged kittiwakes in the Near Islands, which is the reverse of the current situation.

Arctic Tern _(Sterna paradisaea)_

Gabrielson and Lincoln (1959) attribute to the Arctic tern the most extensive range of any Alaskan water bird. It is found in suitable habitat everywhere north of Tracy Arm in Southeastern Alaska. Murie (1959) stated that he found it nesting at suitable sites everywhere he went. Isleib and Kessel (1973) considered it to be an abundant breeder in Prince William Sound and along the northern Gulf Coast.

The Arctic tern was observed in FWS aerial surveys in Prince William Sound, and surveys in July and August 1972 provided an estimate of 45,000 terns in the Sound (Isleib and Kessel 1973). On the other hand, tern colonies were located only rarely in the FWS colony surveys before 1975. This is, however, a reflection of the equipment and methods used and not of the abundance of terns.

From the fragmentary data available, it is not possible to detect changes in Arctic tern status at the present time. We have to assume that the widespread introduction of fox had at least local impact. Although this tern uses a wide variety of nesting sites, it tends to nest on flat sites where access by mammalian predators is easy.

Aleutian Tern _(Sterna aleutica)_

No Aleutian tern colonies were discovered in the Gulf of Alaska area during FWS colony surveys in the early 1970's. This is again a reflection of the fact that surveys were not designed to locate tern colonies. Aleutian terns were encountered at least twice, once during late March 1972 in Hawkins Cutoff, Prince William Sound, and again when two birds were noted offshore from the Katmai National Monument on 30 May 1973 (L. W. Sowl, personal observations).

The type specimen of the Aleutian tern and a single egg were collected at Kodiak Island on 12 June 1868 by Bischoff (Coues 1874). Fisher (Gabrielson and Lincoln 1959) collected four more eggs in 1882. The bird was not found breeding there until Howell (1948) found a colony of 50 pairs at Bell's Flats in 1944. Walker (1923) found them nesting on the Situk River, Yakutat, in 1917 and shortly thereafter saw them at the Alsek River Flats. He also reported that D. H. Stevenson of the Bureau of Biological Survey had told him that they nested on the Isanotski Islands at the end of the Alaska Peninsula. This latter report was the only one from the Aleutian Island chain for many years. Isleib and Kessel (1973) considered it an uncommon local breeder in the northern Gulf of Alaska. Isleib estimated its population at a few hundred pairs on the Copper River Delta in May 1973 and 300-500 birds in June 1970. He also reported that they appeared more or less regularly near Controller Bay and off the Situk River.

In recent years Aleutian terns have been seen with increasing frequency in many places in western Alaska and the Aleutian Islands. This is probably partly due to the increasing level of field work. At Amchitka Island the several colonies that have been found in recent years are almost certainly exhibiting a response to the removal of fox from the island.

Although there is no way of determining what the past status of the Aleutian tern has been in the Gulf of Alaska area, it has been there in small numbers since it was first discovered on Kodiak. It has probably not been abundant at any time and may have suffered a long-term decline brought about by the introduction of fox.

Common Murre _(Uria aalge)_

The common murre is resident in the northern and western Gulf of Alaska from Pinnacle Rock, Kayak Island, westward. East of Cook Inlet colonies are located at Wingham Island, the Martin Islands, Middleton Island, Porpoise Rock in Hinchinbrook Entrance, Barwell Island/Cape Resurrection, the Chiswell Islands, the Barren Islands, and Chisik Island.

For some reason, the islands of the Kodiak-Afognak Archipelago do not host any known major murre colonies. There is also a rather large gap between the Chisik Island colony and the next major colony at Oil Creek west of Puale Bay. Directly west of Oil Creek is another colony at Cape Unalishagvak. Both of these latter colonies are Class V and they are the first colonies of this size to be encountered in the Gulf of Alaska. West of these colonies the next large colony is at Atkulik Island. To the south, midway between the last-named colonies, lies the major composite murre colony in the Semidi Islands. These sites make up the only Class VI colony in the Gulf of Alaska. Westward, the next major colony, a Class V, is at Spitz Island south of Mitrofania Island. In the Shumagin Islands one Class V colony is at Karpa Island, and lesser colonies with large murre components are found at the Haystacks, Castle Rock, and Bird Island. Only minor murre colonies are found between the Shumagin Islands and the end of the Alaska Peninsula.

Gabrielson and Lincoln (1959) were aware only of the colonies at Cape Resurrection (which Gabrielson considered to be large), at the Chiswell Islands, and at Chisik Island for the area from Cook Inlet east. Gabrielson visited the Barren Islands on 13 June 1940 and apparently did not notice the present murre colonies, both Class IV, at East Amutuli (an island which he visited) and Nord Island.

Gabrielson (Gabrielson and Lincoln 1959) found a few small colonies at Kodiak, mostly on small offshore islands. Gabrielson found common murres to be abundant in the Semidi Islands and stated that there were no notable colonies in the Shumagins, although on his return to the Shumagins in 1949 he did find a fairly large colony at the Haystacks. That size description would fit the colony that is there now. He obviously did not see the other colonies. Rausch (1958) reported murres from Middleton Island.

There is quite a difference between the distribution of murres as we know it today and the way Gabrielson and Lincoln pictured it. Why does this difference exist? There are two possible answers: either the number of colonies has increased, or the coverage of colony locations has improved. The latter case, at least, is established. I must confess to being puzzled by the way Gabrielson was able to move about close to what are now known to be sizeable colonies without seeing them, those in the Barren Islands and the Shumagin Islands in particular. Perhaps this represents the vague outlines of yet another population change.

The center of abundance for murre distribution in the Gulf of Alaska today is from Paule Bay west to eastern Shumagin Islands. The Semidi Islands are the heartland of this area of maximum abundance. We have no definitive data on species composition of these colonies. Common murres undoubtedly dominate in most of the colonies; the only ones where we know of a sizeable thick-billed murre component are in the Shumagin Islands.

Thick-billed Murre _(Uria lomvia)_

Thick-billed murre population information cannot be separated from that of the common murre on the basis of existing data. A direct assessment of present-day status is not possible. After reviewing what we know about their distribution, I suggest a way to examine the question indirectly.

The thick-billed murre is found in colonies with the common murre from Middleton Island westward; Rausch (1958) noted about 400 murres at Middleton Island and observed that the thick-billed murre outnumbered the common murre by several times. Isleib and Sowl (FWS, unpublished data) saw a thick-billed murre mixed with common murres at Porpoise Rock in July 1972. Isleib and Kessel (1973) expressed the opinion that small numbers of thick-billed murres will be found in most common murre colonies in the northern Gulf of Alaska when it is possible to survey these colonies in detail. Karpa Island had a significant component of thick-billed murres in June 1973, and they constituted 40% of the colony at the Haystacks (L. W. Sowl, unpublished data).

Bent (1963) reported that many thick-billed murre eggs have been taken by collectors at Round Island in the Shumagin Islands. Dall and Bannister (1869) reported a thick-billed murre that was taken at Kodiak in 1867.

The Gulf of Alaska is at the periphery of the breeding range of the thick-billed murre. While it probably occurs in mixed colonies with the common murre throughout this area, the thick-billed murre is much less abundant. Occasionally in the Gulf of Alaska, a colony will be occupied predominantly by the thick-billed murre. Gabrielson and Lincoln (1959) noted that the thick-billed murre outnumbered the common murre in many colonies in the Aleutians and that it became progressively more common at higher latitudes.

We have almost no data relative to the species composition of murre colonies in the Gulf of Alaska. Until we do it will not be possible to fully understand the population status of the thick-billed murre. It appears that changes in the species composition of murre colonies in the Bering Sea may be an indicator of perturbation. The data for the Gulf of Alaska are still too fragmentary to provide any indication of whether or not the same indicator would work there. Close monitoring of the Shumagin Islands colonies over a number of years might produce the answer.

Earlier in this paper I noted the dramatic changes in species composition of murre colonies on Walrus Island. Gabrielson and Lincoln (1959) also commented on this well-documented and anything but static situation. Investigators who visited this island during 1976 reported seeing no murres on the island and only small numbers on offshore rocks. James Bartonek (personal communication) said that this situation has prevailed for several years.

There is an indication that a similar population fluctuation and change in species composition of murre colonies have also occurred on St. Matthew Island. Bent (1963) found mostly common murres and few thick-billed murres at St. Matthew. Hanna (1916) saw only thick-billed murres. Later, Gabrielson (1941) found this to be true in 1940.

Dramatic fluctuation in murre populations may be common and, at least in some cases, the two species may be affected differently. Perhaps this phenomenon has potential for providing us with an indicator of some natural perturbations.

Peterson and Fisher (1955) expressed the opinion that thick-billed murres arrived at the nesting ledges later than the common murre and had to take the sites that were left. Tuck (1960) reported data from the western Atlantic showing that thick-billed murres arrive later than common murres. On the other hand, Belopol'skii (1961) reported data showing that the two species arrive on breeding colonies in East Murman simultaneously. At Cape Thompson, Swartz (1966) found that thick-billed murres arrived about a week before common murres. The date of arrival, while perhaps a contributing factor, is probably not decisive. Interspecific competition of another sort is indicated.

In mixed murre colonies where there are large numbers of common murres, this species occupies the choice nesting sites. Thick-billed murres are usually left with the narrower ledges while the common murres occupy the longer, broader ledges (Belopol'skii 1961). The broader ledges have lower chick and egg mortality (Spring 1971). Spring also noted that thick-billed murres are excluded from the centers of mixed colonies. Johnson (1938) found that this contributes to higher losses of eggs to predators and to the loss of other social benefits of occupying the colony center (Johnson 1941).

Kozlova (1961) said that during the occupation of a colony there is a sharp competitive struggle between the two species. In the end thick-billed murres are pushed out to the periphery of the colonies or left with narrow ledges or other equally unfavorable sites. Spring (1971) studied the functional anatomy of both species and concluded that the common murre is more successful in these encounters because it has a more upright gait and greater agility than the thick-billed murre.

It follows that in a portion of their respective ranges, where the two species overlap and where there is an equal chance that either common murres or thick-billed murres will dominate a given colony, the common murre dominates. I conclude from this that where there are dramatic changes in species composition of murre colonies, such as at Walrus Island, it is probably because the common murre has been greatly reduced in numbers at the colony.

Spring (1971) concluded that the common murre is well adapted to pursuit and capture of pelagic fishes and that the thick-billed murre is better adapted for deep diving and the capture of benthic fishes and pelagic and benthic invertebrates. Having greater latitude for food selection, the thick-billed murre would have a greater tolerance for ecological perturbations affecting the available food supply. The common murre has an advantage when pelagic fishes are available but cannot switch to the other foods as readily as can the thick-billed murre. The low density of pelagic fishes in high arctic areas probably also accounts for the greater success of the thick-billed murre at higher latitudes relative to common murres.

Belopol'skii (1961) presented data from East Murman which indicates that the common murre restricts its diet almost entirely to a small number of fish species. Swartz (1966) found strong indications that there were significant differences in the food preferences of the two species of murres. Thick-billed murres made much greater use of invertebrates. Bédard (1976) asserted that it is well known that the common murre is quite partial to zooplankton. So again the issue is not clear-cut.

The situation is, of course, much more complex than I have portrayed it. Nonetheless, I think that it offers potential for use as a tool in assessing population change and perturbations in the food supply which should be studied quite closely.

Pigeon Guillemot _(Cepphus columba)_

Gabrielson and Lincoln (1959) noted that the pigeon guillemot was one of the most regularly observed birds in Alaskan waters. It is found everywhere throughout the northern and western Gulf of Alaska area, with only a few understandable and relatively small blanks, such as in the silty waters of Upper Cook Inlet. Because it obviously lacks the breeding murres' need for close contact with its nearest neighbors, it is able to exploit the available nesting habitat to the fullest. It seems that literally every bit of suitable nesting habitat is normally occupied.

Because of the dispersed way in which it breeds and because it does much of its feeding in the onshore zone (which is hazardous for boats) the pigeon guillemot is an almost impossible species to inventory by standard methods.

There is no evidence that the pigeon guillemot has been greatly affected by any major perturbation. Because of its choice of nesting habitat, it is probably subject to the attack of only one egg predator, the rat. Because of its loose social structure and the way it selects nesting sites, eggs and young do not sustain loss from panic flights. Its dispersed distribution should insure that man-made impacts such as oil spills will have limited impact.

The population levels of the pigeon guillemot are probably relatively very stable. The widespread introduction of the rat to most of its nesting range undoubtedly had impact, but this impact has gone undocumented. It would be interesting to follow the response of guillemot populations on islands where rats had been totally removed, if that ever becomes more than a dream.

Marbeled Murrelet _(Brachyramphus marmoratus)_

The marbled murrelet apparently breeds throughout most of the northern and western Gulf of Alaska. This apparently is a necessary condition because to date, at least in this part of Alaska, we can only guess where and under what conditions this murrelet breeds.

In some relatively sheltered waters like Prince William Sound, where marbled murrelets were estimated to number about 250,000 in 1972 (Isleib and Kessel 1973), they are the most abundant seabirds. We know from Dixon (1908) and Grinnell (1910) that this has been so in Prince William Sound since the beginning of the century. We know also that the type specimens came from there as well (Stresemann 1949), which is not necessarily an indication of abundance but is suggestive of their abundance relative to species not collected.

Gabrielson (Gabrielson and Lincoln 1959) found marbled murrelets common near Yakutat, in Prince William Sound, in Resurrection Bay, and at Kodiak, and reported seeing them at the Chiswell Islands and at Chignik and Pavlof Bay on the Alaska Peninsula. Cahalane (1943, 1944) found them to be common in Kupreanof Strait, and along the Alaska Peninsula north of Katmai Bay. Murie (1959) found them all along the Alaska Peninsula. My own field notes from 1973 indicate that the only place where they were common along the Alaska Peninsula was at Wide Bay.

We can sample marbled murrelet numbers by using standard transect methodology; however, I have some very serious reservations about our ability to convert these data into a population estimate. This is not an unusual assessment for Alaskan seabirds in general, but I think it is particularly apropos to this species.

We are still able only to guess at where the marbled murrelet nests and we have not a clue as to what sort of nesting strategy they pursue. I am not prepared to accept, on the basis of one North American record (Binford et al. 1975), that tree nesting is its habit throughout its range. What has been proved is that the marbled murrelet nests in trees and not, as these authors would have us believe, that it does not nest on the ground. It has become rather fashionable to ignore the Chichagof Island record (a ground nest), but it has not been discredited. The color of the Chichagof egg differs from that of the Big Basin egg, but does agree with the one taken from an oviduct by Cantwell (Gabrielson and Lincoln 1959). My own experience leads me to believe that tree nesting, if it occurs, is not the common habit of marbled murrelets nesting in the Prince William Sound region.

After many hours of observing marbled murrelets over a period of several years, I am intrigued by a number of things. These birds, as often as not, appear to be clustered in "pairs" as they feed. This occurs even at what should be the height of the breeding season. On several occasions I have noted a very pronounced evening flight of these birds from gathering areas on the water up into the surrounding mountains at sunset. This has moved me to wonder if their nesting strategy includes incubating at night but less than full-time attendance on days when the eggs can be warmed by the sun. We know that periodic egg-neglect is an aspect of storm-petrel behavior (Pefaur 1974). Is this behavior also possible on a more regular basis in an alcid? If so, it would certainly help explain why nests are hard to find.

It is apparent that more needs to be known about the population dynamics and life history of the marbled murrelet before we can make a proper estimate of its abundance. In spite of the fragmentary record, I conclude that the marbled murrelet probably enjoys the same relative abundance and distribution that it did at the beginning of the century.

Kittlitz's Murrelet _(Brachyramphus brevirostris)_

The Kittlitz's murrelet is not as abundant as the marbled murrelet, but locally it is sometimes found in large numbers. FWS surveys conducted during July-August 1972 provide an estimate of 57,000 murrelets of this species in Prince William Sound. Almost a fifth of these were concentrated in Unakwik Inlet above Unakwik Reef. Even more interesting, about 2,500 of these birds were concentrated in one loose flock.

In addition to Unakwik Inlet, Kittlitz's murrelets concentrate in College Fjord in Prince William Sound and in the waters fronting the Bering-Malaspina ice-fields (Isleib and Kessel 1973). Common as they are in these waters, this species is supposed to be even more abundant at Glacier Bay. The common feature of these waters is the amount of ice that can be found below their tributary glaciers.

The Kittlitz's murrelet is apparently distributed from LeConte Bay, east of Petersburg, Alaska, north to Point Barrow and west across the Aleutians to Attu, where Murie collected a pair (Gabrielson and Lincoln 1959). I once flushed a murrelet from an area of tread and riser topography near the top of the highest point on Kiska Island in heavy cloud cover, and although I could not see this bird well, I thought it to be of this species. From the range description in Gabrielson and Lincoln (1959) and Udvardy's (1963) range map, it is apparent that the distribution of this species is rather patchy, but I suspect that for the more mountainous part of its range this is more apparent than accurate. The record is too fragmentary to allow an assessment of any change in status during the historical period.

Ancient Murrelet _(Synthliboramphus antiquus)_

Chase Littlejohn (Bendire 1895) spent the spring and summer of 1894 collecting eggs on islands south of the Alaska Peninsula. He has left us a detailed record of what he saw but not where he saw it. Bent (1963) stated flatly that the site of his collecting was Sanak Island and this has common acceptance. Several things in his account point to a site which was a small island with several peers close by, but this could not have been Sanak. It could have been an island in the Sanak Island group or it could equally well have been somewhere in the Sandman Reefs. Unfortunately, because of this the record is clouded. There has never been anything approaching a survey of the southern half of the Sandman Reefs. We do not know what breeding colonies are there.

At any rate, Littlejohn told of the large numbers of Leach's storm-petrels, fork-tailed storm-petrels, auklets (of which only Cassin's is specifically identified), and ancient murrelets which occupied a large number of small islands. He could not calculate the number of breeding murrelets on his small island, the size of which I interpret to have been of the same order of magnitude as two others which he estimated were about 2 acres. He does say that the murrelets must have numbered several thousand and could, if left alone by the Aleuts, have quickly grown too numerous for the island to accommodate.

Murie (1959) made a brief visit to Sanak in 1937 and learned that there were no longer any large colonies of seabirds. He attributes this to exploitation of the fisheries and to the fox-farming industry. Littlejohn told of the repeated visits of Aleuts to his small islands, where they took hundreds of birds each time and all of the eggs they could find. This kind of activity could not help but disrupt the breeding on these islands.

Littlejohn's description of the ancient murrelet's nest leaves little doubt that the birds could be reached by fox or rats with ease. The birds showed no particular care in selecting a nest site and often worked their way back no more than about a meter into the dead vegetative cover from preceding years, where they scratched out a shallow nest.

There are few records of the ancient murrelet from the northern and western Gulf of Alaska. Friedmann (1935) reported the collection of a series of eggs in 1884 on Kodiak Island. Chase Littlejohn (Bendire 1895) collected eggs from somewhere in the Sanak Group in 1894. In 1908 Dixon (Grinnell 1910) saw a bird in Port Nellie Juan. Several were seen by Jaques (1930) near Belkofski in May 1928. Gabrielson collected one bird at Cordova in September 1941 and another at the Chiswell Islands in July 1945 (Gabrielson and Lincoln 1959). He saw numerous flocks in the Gulf of Alaska on 30 July of an unnamed year. In 1943, he would have been near Cape Spencer on that date. In 1945 he would have been near the Chiswell Islands. In either case, he was probably somewhere in Blying Sound.

The ancient murrelet is relatively uncommon but regularly observed in the inshore waters along the outer coasts of the islands fronting Prince William Sound. FWS surveys in July-August 1972 provided an estimate of almost 1,000 birds, mostly in nonbreeding plumage, along the outer coast of Prince William Sound (Isleib and Kessel 1973). Small numbers were found feeding close to the Wooded Islands on 24 July (my personal observation). Rausch (1958) saw a few off Middleton Island in 1956. Isleib (Isleib and Kessel 1973) saw 400-500 widely distributed at the mouth of Yakutat Bay in July and August 1968. The only large numbers of ancient murrelets encountered on the FWS survey of the Alaskan Peninsula in 1973 were in the Shumagin Islands. They were very common in East Nagai Strait on 9 June and more than half of the 1,300 seabirds per square nautical mile encountered between Little Koniuji and Chernabura Islands on 11 June were ancient murrelets. At Nagai Island an estimated 5,000 ancient murrelets were observed in the west bay at Pirate Shake, and later (on 19 June) several were observed in the vicinity of Midun Island (FWS, Anchorage, Alaska, unpublished data).

On the basis of the observations recounted above, I have to conclude that ancient murrelets are fairly regularly, if patchily, distributed throughout the northern and western Gulf of Alaska. I do not believe that the void in their range shown for the northern Gulf of Alaska by Udvardy (1963) is correct. Several colonies are there, awaiting discovery.

Ancient murrelets are not abundant in the Gulf of Alaska but they are certainly more numerous than we have been able to prove. It is not possible to tell from the existing data whether they were once more abundant than they are now. I suspect, on the basis of the Sanak Island experience, that we can conclude that this species has been reduced in number by various of man's activities.

Cassin's Auklet _(Ptychoramphus aleuticus)_

Cassin's auklet is a very uncommon bird in the northern Gulf of Alaska. In the western Gulf it is more common, particularly from the Shumagins west.

This auklet apparently once bred in great numbers on islands in or near the Sanak Group where Chase Littlejohn (Bendire 1895) found them to be twice as numerous as the ancient murrelets. Murie (1959) did not find them there.

Littlejohn began encountering Cassin's auklets at sea some 290 km southeast of Unga, Shumagin Islands. Murie (1959) encountered them near the Shumagins in May 1937. During the FWS 1973 reconnaissance survey of the Alaska Peninsula, these auklets were not encountered (or at least not identified) until we reached the vicinity of Unga Strait where we saw a few in mixed flocks with other murrelets and auklets. They were most numerous in East Nagai Strait. We encountered them only twice in a situation which indicated they might be breeding--on Hall and Herendeen islands on the north end of Little Koniuji Island.

Murie (1959) considered Cassin's auklet to be no longer common west of Kodiak. In Gabrielson's many voyages through the northern and western Gulf of Alaska he encountered them only twice, once off Cape Spencer and once in the Chiswell Islands.

Thoresen (1964) commented that throughout the northern part of its range the Cassin's auklet has become gradually less frequent. Although there are no data to dispute this, I believe, as do Isleib and Kessel (1973), that they are more numerous than observations would indicate, and I would apply this to the entire area. There are certainly colonies remaining in the Shumagin Islands, and quite probably along the south coast of the Kenai Peninsula. When it is possible to fully explore the Sandman Reefs there is a good probability that they will be found there.

We can only guess at the reasons for their decline. Bendire (1895) and Murie (1959) have described some contributing factors.

Parakeet Auklet _(Cyclorrhyncus psittacula)_

Gabrielson and Lincoln (1959) described the parakeet auklet as the least colonial of any of the Alaskan auklets. They also considered the Aleutian Islands to be its principal nesting grounds. There are old records of breeding parakeet auklets from Kodiak (Friedmann 1935) and Little Koniuji (Bean 1882). Grinnell (1910) reported two that were seen on Green Island, Prince William Sound, and several more that were seen near Knight Island.

Murie (1959) did not see any parakeet auklets near Kodiak and Afognak islands which he considered to be the eastern part of their range. He did not think they were abundant anywhere along the Alaska Peninsula. He found a few near Sutwik Island in May 1936 and then noted that they were fairly common near the Shumagins in May 1937.

Gabrielson found this species to be quite numerous on the north side of Chowiet Island in the Semidi Islands in 1945 (Gabrielson and Lincoln 1959). He also saw numerous individuals in Marmot Strait and saw one in the Chiswell Islands during the same year. David Roseneau (Isleib and Kessel 1973) found hundreds close to East Amatuli Island in the Barren Islands in 1965.

During FWS colony surveys, parakeet auklets have been found in close proximity to six seabird colonies in Prince William Sound. During the July-August 1972 surveys, they were estimated to number about 3,000 in the Sound. They have also been found closely associated with Chisik Island (David Snarski, personal communication), the Chiswell Islands, Nord and Sud islands in the Barrens, Sea Otter Island, and Central and Long islands along the Alaska Peninsula. They were most numerous in the Shumagin Islands, where they were found near Castle Rock, Hall (9,000), Herendeen (3,000), Atkins (more than 5,000), and Little Koniuju islands. They were again encountered south and west of Cold Bay at High, Fawn, Let, Amagat, Umga, and Patton islands. Many of these islands are in the north half of the Sandman Reefs, the only portion where any attempt has been made to survey seabird colonies.

The parakeet auklet may not be abundant anywhere in the Gulf of Alaska but, based on the numbers of places it has been seen in recent years, its population appears to be well dispersed and probably doing very well. This auklet is most abundant from the Shumagin Islands westward. It is almost certainly more numerous than has been thought. Its habits are secretive enough so that it could easily escape notice.

Because the parakeet auklet nests predominantly under boulders, it probably was not much affected by fox. Rats would certainly have reduced its numbers wherever these were introduced into its breeding habitat. We have no data to tell us whether there may have been population fluctuations in the past, but there undoubtedly were at least minor ones locally after rats were introduced.

Crested Auklet _(Aethia pygmaea)_

Udvardy (1963) shows the breeding range of the crested auklet as extending from southern Kodiak Island westward. Within the northern and western Gulf of Alaska, it is certainly most abundant in the eastern Shumagin Islands.

Isleib saw this auklet in Prince William Sound 3 times during the winter of 1972-73. These are the only records he was aware of for that area (Isleib and Kessel 1973). David Roseneau (Isleib and Kessel 1973) saw several in Amatuli Cove, Barren Islands, in June 1965. I observed one in the vicinity of Cape Spencer in August 1973.

Friedmann (1935) listed the crested auklet as a breeding bird at Kodiak, but considered it to be much more abundant as a wintering bird. Townsend (1913) has provided us with a vivid description of the myriads of crested auklets he encountered at Yukon Harbor, Little Koniuji Island. Gabrielson and Lincoln (1959) noted large numbers of crested auklet around Simeonof and Bird islands in the Shumagin Islands in 1946 and stated that the Yukon Harbor colony was still thriving.

Crested auklets were not encountered on the 1973 FWS reconnaissance survey until we reached the Shumagin Islands. They were abundant only at the southeastern end of Little Koniuji, where we encountered perhaps 10,000 in Yukon Harbor and more than 50,000 in a small cove directly south of Yukon Harbor on the opposite side of the island. As numerous as they were, they did not match Townsend's myriads or even come close to his assessment that they "were here more numerous than the 'choochkies' at St. George." St. George Island in the Pribilofs is famous for its least auklets which, in the past, have been estimated to number as high as 36 million (Peterson and Fisher 1955). The numbers there today do not even approach this level and we have no way of knowing how abundant they were when Townsend visited the Pribilofs, but I think it is safe to say that they probably numbered in the millions. There are probably more crested auklets than we observed on Little Koniuji, but there is certainly no longer anything approaching millions of birds. Properly pronounced, Koniuji is the Aleut name for the crested auklet, so we can assume that the original inhabitants were impressed by its numbers.

During the 1973 FWS survey we did not see crested auklets at either Simeonof or Bird islands. On the overgrazed and cattle-trampled Simeonof it does not seem possible that any could still exist.

I suspect that a cattleman's greed has been the undoing of any crested auklets that may have nested on Simeonof Island. This would not account for the loss of any colonies that may have been on Bird Island, but the decaying fox-trapper's cabin on that island undoubtedly tells the story. Churnabura, with its feral cattle, presents much the same problem as Simeonof. As for Little Koniuji, have horned puffins been

## partly responsible for the decrease in crested auklets? The puffin

colony at the south end of Little Koniuji must be exactly where Townsend's millions of crested auklets once nested.

Least Auklet _(Aethia pusilla)_

No least auklets were encountered in FWS surveys in the Gulf of Alaska in the early 1970's. Udvardy (1963) shows their breeding range as starting well west in the Aleutians. Gabrielson and Lincoln (1959) give the eastern limit of their breeding range as the Shumagin Islands. Bent (1963) listed their breeding range as extending east to Kodiak Island, and Friedmann (1935) knew of only a few specimens taken in the winter from Kodiak. Perhaps least auklets nested somewhere in the western Gulf of Alaska, and they may still, but at the moment we have no evidence to prove that they do.

Rhinoceros Auklet _(Cerorhinca monocerata)_

Udvardy (1963) would have us believe that the rhinoceros auklet did not nest between southeastern Alaska and the southern Kurile Islands. Bent (1963_b_), on the other hand, lists their breeding range as extending from Washington to Agattu. Clark (1910) noted this species in small numbers at Atka and Agattu. Because of the lack of proof, Udvardy probably had no options. I believe that Bent was probably closer to describing their original range. I base this assumption on recent observations and on the additional fragments of information reported by Gabrielson and Lincoln (1959). Murie (1959) failed to find this species anywhere in the Aleutians, but his primary reason for being there, the fox-farming industry, may have had a lot to do with his not being able to find any.

The FWS surveys in Prince William Sound in July-August 1972 located small numbers of rhinoceros auklets in breeding plumage at the Wooded Islands and at Stoney Island and Channel Island in Montague Strait. These birds gave every impression of being local breeders. David Roseneau (Isleib and Kessel 1973) encountered two at the Barren Islands in June 1965. Isleib and Kessel (1973) list a few other records from this area.

My own experience leads me to believe that there is a large colony somewhere on Afognak Island, probably on or near Tonki Cape. On 30 May 1973 I noted a lone bird north of Afognak Island. Later, on 8 and 9 August, I saw several in the same area. On 13 August in Marmot Strait I observed a number of rhinoceros auklets, either singly or in groups of up to 12. Some of these had small fish in their beaks. As they flushed, they all flew off toward Tonki Cape. This observation was made just at last light, and I believe that there were many others about that could not be seen in the dying light. We did not encounter this species along the Alaska Peninsula during the FWS survey in 1973 until we reached the end. There I had one quick glimpse of what I was certain was a rhinoceros auklet at Amagat Island.

Horned Puffin _(Fratercula corniculata)_

The horned puffin is one of the most abundant breeding birds in the Gulf of Alaska. There are only a few really large colonies but these birds breed just about anywhere there is a cliff (even a low one) with suitable fractures and crevices. During the Alaska Peninsula surveys in 1973, I estimated that the frequency with which these birds were seen on the water was about half that of the tufted puffin. They have been recorded in so many places that there is nothing to be gained by a reiteration of the record in the literature.

The horned puffins reach their greatest density in the Gulf of Alaska west of Kodiak Island. Murie (1959) estimated that the colony at Amagat Island, Morzhovi Bay, contained 15,000 birds, one of the largest he had seen. It contained at least 50,000 in 1973. Even at that, it was no match for the colony on Little Koniuji Island with its minimum 140,000 horned puffins. Other colonies with large horned puffin components were at High Island (40,000), Castle Rock (20,000), Mitrofani Island (35,000), and Sosbee Bay (15,000).

Earlier in this paper, I commented at length on the great and often rapid fluctuations in populations of tufted puffins. The same phenomenon affects horned puffins. In 1975 there were relatively small numbers of horned puffins at Little Koniuji where they had flourished 2 years earlier (James Bartonek, personal communication). Because they are apparently subject to erratically oscillating populations, it is hard to tell how they have fared over the years.

Tufted Puffin _(Lunda cirrhata)_

The tufted puffin, as previously indicated, is also a bird with widely fluctuating populations. Until we develop an understanding of their population dynamics and can understand the underlying cause of these fluctuations it will not be possible to assess trends in their populations or understand the implications of such trends.

Tufted puffins are abundant throughout the Gulf of Alaska. Small colonies can be located almost anywhere. Along the Alaska Peninsula there are a number of colonies with an estimated breeding population in 1973 of more than 15,000 birds. These are: Ashiiak Island (20,000), Central Island (90,000), the Brother Islands (45,000), The Haystacks (19,000), Castle Rock (85,000), Bird Island (none, but may contain 500,000-1,000,000 at times), Peninsula Islands (35,000), the Twins (18,000), Amagat Island (40,000), and Umga Island (22,000). These colonies correspond to the area where colonies were listed for the horned puffin.

Tufted puffin populations respond readily to some undetermined short-term perturbations. This is clearly demonstrated by their rapid population fluctuations. Because of their numbers and because of the apparent rapidity with which their numbers rebound, it is not so apparent that they have been affected by long-term perturbations, as so many other seabirds apparently have.

There is much unused or underused nesting habitat suitable for this species. In some cases there are very strong clues pointing to why this habitat is vacant. On many islands along the Alaska Peninsula, which have very good-looking tufted puffin nesting habitat and no puffins, there are visible signs of the presence of fox--either fox trails or abandoned trappers' cabins. I also suspect that the brown bear _(Ursa arctos)_ is another possible contributing factor to population declines of burrow nesters along this coast. I have seen brown bears swimming from island to island on foraging expeditions. George J. Divoky (personal communication) has found brown bears visiting Ugaiushak Island, which is 13 km from shore. There are other islands between Ugaiushak and the mainland but the shortest route from shore would require one swim of 7 km. The motivation must be strong.

Tufted puffins may shift from colony to colony. This could be an explanation for apparent local population fluctuation, but if so, I am puzzled by the apparent tenacity with which puffins cling to some sites. Their constant occupancy of sites where the vegetative mat is breakaway tundra (Amundsen 1972) or is underlain by sand results in the destruction of these sites. Tufted puffins often cling to them in spite of the fact that they have been reduced to "slums."

My conclusion is that in spite of their large numbers it appears that tufted puffin populations in the Gulf of Alaska probably have been reduced to a level below that of their undisturbed state.

Conclusions

Seabird numbers in the Gulf of Alaska are not static. Generally, they are probably much less abundant than they were when Bering made his voyage of discovery. There are, nonetheless, considerable numbers of seabirds breeding along the coasts of these waters. Some species show signs of recovery from past insults by man. With enlightened management there is still time to preserve the vast natural heritage that they represent and, in many cases, to improve their status.

In attempting to address a complicated subject in short space and a relatively narrow frame of reference, I have certainly erred a number of times. I would like to see the wealth of new data that will be derived from current work applied to this concept. An understanding of past population fluctuations and the underlying perturbations that they reflect is essential for managers faced with the problem of making good decisions on measures to mitigate the potential adverse impact of development.

References

American Ornithologists' Union. 1931. Check-list of North American birds, 4th ed. American Ornithologists' Union, Baltimore, Md. 530 pp.

American Ornithologists' Union. 1957. Check-list of North American birds, 5th ed. American Ornithologists' Union, Baltimore, Md. 691 pp.

Amundsen, C. C. 1972. Plant ecology of Amchitka Island. Amchitka bio-environmental program. Final Report, Battelle Columbus Lab., Columbus, Ohio. 27 pp.

Bailey, A. M. 1948. Birds of arctic Alaska. Colo. Mus. Nat. Hist. Pop. Publ. 8. 316 pp.

Baird, S. F. 1869. On additions to the bird fauna of North America, made by the scientific corps of the Russo-American Telegraph Expedition. Trans. Chic. Acad. Sci. 1(2):311-325.

Beaglehole, J. C. 1974. The life of Captain James Cook. Stanford Univ. Press, Stanford, Calif. 734 pp.

Bean, T. H. 1882. Notes on birds collected during the summer of 1880 in Alaska and Siberia. Proc. U.S. Natl. Mus. 5:144-173.

Bédard, J. 1976. Coexistence, coevolution and convergent evolution in seabirds: a comment. Ecology 57(1):177-184.

Belopol'skii, L. O. 1961. Ecology of sea colony birds of the Barents Sea. (Transl. from Russian.) Israel Program for Scientific Translations, Jerusalem. 346 pp.

Bendire, C. 1895. Notes on the ancient murrelet (Synthliboramphus antiquus) by Chase Littlejohn with annotations. Auk 12(3):270-278.

Bent, A. C. 1963. Life histories of North American diving birds. Dover Publications, Inc., New York. 239 pp.

Bent, A. C. 1964. Life histories of North American petrels, pelicans and their allies. Dover Publications, Inc., New York. 335 pp.

Binford, L. C., B. G. Elliot, and S. W. Singer. 1975. Discovery of the nest and downy young of the marbled murrelet. Wilson Bull. 87(3):303-319.

Bowles, J. H. 1908. Tapeworm epidemic among Washington seabirds. Condor 11:33.

Cahalane, V. H. 1943. Notes on the Kodiak-Afognak Island group. Auk 60(4):536-541.

Cahalane, V. H. 1944. Birds of the Katmai region, Alaska. Auk 61(3):351-375.

Clark, A. H. 1910. The birds collected and observed during the cruise of the United States Fisheries Steamer "Albatross" in the North Pacific Ocean and in the Bering, Okhotsk, Japan and Eastern seas from April to December 1906. Proc. U.S. Natl. Mus. 38:25-74.

Cody, M. L. 1973. Coexistence, coevolution and convergent evolution in seabirds. Ecology 51(1):31-44.

Coues, E. 1874. Birds of the Northwest. U.S. Geological Survey of the Territories, Misc. Publ. 3. 791 pp.

Dall, W. H. 1873. Notes on the avifauna of the Aleutian Islands, from Unalaska eastward. Proc. Calif. Acad. Sci. 5(1):25-35.

Dall, W. H. 1874. Notes on the avifauna of the Aleutian Islands, especially those west of Unalaska. Proc. Calif. Acad. Sci. 5:270-281.

Dall, W. H., and H. M. Bannister. 1869. List of the birds of Alaska, with biographical notes. Trans. Chic. Acad. Sci. 1(2):267-310.

Dement'ev, G. P., and N. A. Gladkov, eds. 1966. Birds of the Soviet Vol. I. (Transl. from Russian.) Israel Program for Scientific Translations, Jerusalem. 553 pp.

Dixon, J. S. 1908. Field notes from Alaska. Condor 10:139-143.

East, B. 1943. Seabird cities of the Aleutians. Nat. Hist. 51:64-71.

Fisher, J., and R. M. Lockley. 1954. Seabirds. William Collins and Sons, London. 320 pp.

Friedmann, H. 1934. Bird bones from Eskimo ruins on St. Lawrence Island. J. Wash. Acad. Sci. 24:83-96.

Friedmann, H. 1935. The birds of Kodiak Island, Alaska. Bull. Chic. Acad. Sci. 5(3):13-54.

Friedmann, H. 1937. Bird bones from archeological sites in Alaska. J. Wash. Acad. Sci. 27:431-438.

Gabrielson, I. N. 1940. America's greatest bird concentrations,

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Gabrielson, I. N. 1941. America's greatest bird concentrations,

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