Arctic Seabirds: Diversity, Populations, Trends, and Causes

Home , Digges Sound, Lancaster Sound, Spectacled cormorant

ARCTIC SEABIRDS: DIVERSITY, POPULATIONS, TRENDS, AND CAUSES

ANTHONY J. GASTON

Wildlife Research Division, Environment Canada, National Wildlife Research Centre, Ottawa K1A 0H3, Canada. E-mail: [email protected]

ABSTRACT.—Populations and trends of Arctic seabirds have been the subject of substantial research since the 1930s in Europe and Greenland and since the 1950s in North America. The marine waters of the Arctic support 44 species of seabirds comprising 20 genera. There are four endemic monotypic genera and an additional 25 species for which the bulk of the population is confined to Arctic and sub-Arctic regions. Most Arctic seabirds have large populations, with only two species comprising less than 100,000 individuals and many species numbering in the millions. Population trends for several widespread Arctic species have been negative in recent decades. Conversely, some sub-Arctic species are spreading northwards. Climate change with consequent changes in competition and predation, and intensifying development in the north, increasingly threaten Arctic seabirds. Changes in ice conditions are likely to have far-reaching and potentially irreversible results. Received 22 February 2011, accepted 26 May 2011.

GASTON, A. J. 2011. Arctic seabirds: Diversity, populations, trends, and causes. Pages 147–160 in R. T. Watson, T. J. Cade, M. Fuller, G. Hunt, and E. Potapov (Eds.). Gyrfalcons and Ptarmigan in a Changing World, Volume I. The Peregrine Fund, Boise, Idaho, USA. http://dx.doi.org/10.4080/ gpcw.2011.0201

Key words: Seabirds, diversity, population size, population trends, Arctic.

SEABIRDS HAVE PROVIDED A SOURCE OF FOOD for Census of Arctic seabird colonies began in the Arctic peoples throughout their history and 1930s in Greenland (Salomonsen 1950) and most major seabird colonies within the range Russia (Uspenski 1956), the 1950s in eastern of the post-Pleistocene Inuit expansion are Canadian Arctic (Tuck 1961), the 1960s in associated with archaeological sites that attest Spitzbergen (Norderhaug et al. 1977, Mehlum to significant harvest and storage of seabirds and Bakken 1994), and the 1970s in the north- (Freuchen and Salmonsen 1958, Nelson 1983). ern Bering and Chukchi seas (www.seabirds. Some Arctic communities are heavily depend- net/maps/dev/north-pacific.php?v=14). Subse- ent on seabird harvesting (e.g., Ivujivik, Que- quent monitoring was sporadic in most regions bec, Gaston et al. 1985; Siorapaluk, Greenland, until the 1980s, but has become more regular Malaurie 1985). Early European explorations since then (CAFF Seabird Working Group, in Arctic and sub-Arctic waters also made unpubl.). However, many breeding sites are extensive use of seabirds for food (e.g., Henry extremely remote, in places where navigation Hudson’s crew, Prickett 1611). Consequently, is challenging and support for aircraft very dis- these birds have been of interest to people for tant. Consequently, although we have a good a very long time. picture of distributions, and some idea of pop-

147 –GASTON – ulation sizes, our information on population Several taxa have been elevated to species sta- trends is rather fragmentary and localised and tus only recently and were previously consid- our knowledge of causes behind change in ered sub-species. These splits mainly involve populations is even less substantial (Gaston et distinguishing North American and Eurasian al. 2009). This paper attempts to review what populations: Arctic/Pacific Loons (Gavia arc- we do know of seabird population size and sta- tica/pacifica), American/European Herring tus, understanding that there is considerable Gull (Larus smithsonianus/argentatus). The uncertainty especially about population trends. large white-headed gulls of the genus Larus are divided into several poorly differentiated and mostly allopatric species in northern Asia SPECIES RICHNESS AND DISTRIBUTION and on the west coast of North America. Much Forty-four species of seabirds breed within the of their diversity was regarded as infra-specific Arctic (Table 1), 23 in the High Arctic, 41 in until recently (cf. Vaurie 1965, Olsen and Lars- the Low Arctic. They belong to 20 genera, the son 2003). richest in species being Larus (10 spp.), Gavia (5 spp.) and Stercorarius (4 spp.). Six genera The distributions of many species of Arctic are monotypic. The majority are members of marine birds were poorly known until the latter the order Charadriiformes, 34 species, includ- half of the twentieth century. In addition, many ing four endemic genera, all monotypic: Little species are long-lived and conservative in their Auk (Alle alle), Ivory Gull (Pagophila breeding site adherence, making them slow to eburnea), Sabine’s Gull (Xema sabini), and alter their breeding range. Consequently, we Ross’s Gull (Rhodostethia rosea). Fifteen have few data on which to assess trends in species are circumpolar in their distribution, range extent among Arctic seabirds. No strictly occurring in Canada, Alaska and over most of Arctic species has become extinct during his- the Russian Arctic. There are two ‘bi-polar’ toric times, although three sub-Arctic species, genera, found at high latitudes in both hemi- Spectacled Cormorant (Phalacrocorax per- spheres–the fulmars (Fulmarus), and the skuas spicillatus) (Commander Islands), Labrador and jaegers (Stercorarius), the former likely Duck (Camptorhynchus labradorius) originating in the southern hemisphere (Voous (Labrador) and Great Auk (Pinguinus impen- 1949), the latter in the northern hemisphere nis) (Newfoundland and Iceland) were hunted (Furness 1987). All four species of Stercorar- to extinction by Europeans in the 19th Century ius found in the northern hemisphere are (Fuller 2000). Ivory Gull and Ross’s Gull are endemic to the Arctic and sub-Arctic, as is the listed by IUCN/Birdlife International as threat- single fulmarine petrel, all the loons, terns and ened or endangered at a world scale. auks and five species of Larus gulls. There is some evidence for the recent north- Overall diversity is highest in the low-Arctic ward spread of predominantly temperate or of the Pacific Basin (Chukchi and Bering seas low-Arctic species: Ancient Murrelet (Synthli- and adjacent coasts) where 28 species occur in boramphus antiquus) in the Bering Sea (Gas- the Alaskan low-Arctic (including islands ton and Shoji 2010), Horned Puffin south to 60º N) and 26 species on the Asian (Fratercula corniculata) in the Beaufort Sea side. Other biodiversity hotspots occur in West (Moline et al. 2008), Mew (Common) Gull Greenland (24 spp.), eastern Canadian Arctic (Larus canus) in Iceland (Petersen and (Nunavut, northern Quebec and Labrador, 22 Thorstensen 2004), Black-headed Gull spp.), and Iceland (22 spp. excluding the sub- (Chroicocephalus ridibundus) in Labrador Arctic/boreal species found only on the south (Chaulk et al. 2004), Great Black-backed Gull coast). (Larus marinus) and Razorbill (Alca torda) in Hudson Bay (Gaston and Woo 2008). At the

148 –POPULATIONS AND STATUS OF ARCTIC SEABIRDS – same time there is evidence of a retreat for at associated species such as Ross’s and Ivory least one high-Arctic species, with the range of gulls and Thick-billed Murre (Uria lomvia). the Ivory Gull contracting in northern Nunavut, with most colonies on northern Baf- POPULATION SIZES AND DENSITIES fin Island and eastern Devon Island deserted while numbers have remained stable farther Most species have populations numbering in north on central Ellesmere Island (Environ- the hundreds of thousands and only two are ment Canada 2010). Southern colonies are also believed to number less than 100,000 breeding decreasing in Greenland (Gilg et al. 2009). The individuals: Ivory Gull and Thayer’s Gull population trend in Russia is unclear (Gilchrist (Larus thayeri) (Table 1). Among high-Arctic et al. 2008) but some colonies at their western specialists, the Ivory Gull has decreased pre- extremity in the Barents Sea region have been cipitously in Canada (by 80% since the 1980s), deserted (Gavrilo 2010). The population of has decreased in Greenland, and shows range Kittlitz’s Murrelet (Brachyramphus brevi- contraction in the northern Barents Sea. In all rostris), a species associated with tidewater cases the southern parts of the range seem to glaciers in the low- and sub-Arctic of the be more affected than northern parts (Gilchrist North Pacific, is declining in its core breeding and Mallory 2005, Gilchrist et al. 2005, Gilg et range in south central Alaska and perhaps else- al. 2009, Environment Canada 2010). Of the where (Kuletz et al. 2003, Stenhouse et al. other two exclusively high-Arctic species, 2008). Similar changes have been noted by population size for Thayer’s Gull, which is local people: confined to eastern and central parts of the Canadian high-Arctic and northwest Green- “I have started to notice birds which I land, is very poorly known, but certainly num- used to only see on TV, little birds which bers less than 100,000 (AJG, M.L. Mallory have multi-coloured bills, that fly home and H.G. Gilchrist unpubl.). The Little Auk, with multiple cod in their beaks and that although well-distributed in small pockets burrow into the soil. I think these are around the Arctic Ocean, is numerically con- the Atlantic puffins [Fratercula arctica], centrated into a single location in northwest which are located some distance south Greenland (Crimson Cliffs and adjacent coasts migrating north due to the disappear- of Thule District) where the population, ance of the ice cover during the summer although difficult to count, is believed to months” (Pijamin: Elders Conference greatly exceed ten million birds (Renaud et al. on Climate Change 2001). 1982). Censusing such an aggregation is almost impossible and no information is avail- With little evidence for range changes, it is dif- able on trends. Some small colonies in south- ficult to ascribe causes. The spread of Razor- ern Greenland and in Iceland have disappeared bill in Hudson Bay has been linked to an since the 1930s (Nettleship and Evans 1985). increase in sandlance Ammodytes spp., perhaps related to diminishing ice cover (Gaston and Thick-billed and Common Murres (Uria Woo 2008). Reduced ice cover also is likely to aalge)are among the most abundant seabirds in be involved in the arrival of Horned Puffin in the Northern Hemisphere with both species the Beaufort Sea. The association of Kittlitz’s exceeding 10 million adults (Gaston and Jones Murrelet with tidewater glaciers makes it 1998). Both have circumpolar distributions. likely that recent declines are caused by the The more northern species, U. lomvia, occurs retreat of many Alaskan coastal glaciers (Sten- mostly in Arctic waters, where it constitutes a house et al. 2008). In the longer run, changes higher proportion of seabird biomass than any in ice cover must affect the distribution of ice- other species. Both species of murre have shown regional population changes over the

149 –GASTON –

Table 1. Seabird species occurring in the circumpolar Arctic, by region, with global population estimate (orders of magnitude and IUCN status). 1 Alaska NWT Nunavut Quebec Labrador Greenland Iceland Svalbard Europe Russia Siberia pop'n World IUCN Status

Loons Gaviidae Red-throated Loon Gavia stellata XXXXXXXX6LC Black-throated Loon Gavia arctica X XXX6LC Pacific Loon Gavia pacifica XXX X6LC Great Northern Loon Gavia immer XXXXX 6LC Yellow-billed Loon Gavia adamsii XXXX XX5LC

Petrels Procellariidae Northern Fulmar Fulmarus glacialis X XXXXXX8LC

Cormorants Phalacrocoracidae Great Cormorant Phalacrocorax carbo XX 6LC European Shag Phalacrocorax aristotelis XX6LC Pelagic Cormorant Phalacrocorax pelagicus XX6LC

Gannets Sulidae Northern Gannet Morus bassanus XX6LC

Jaegers/Skuas Stercorariidae Great Skua Stercorarius skua XXX 5LC Pomarine Skua Stercorarius pomarinus XXXX XX6LC Parasitic Jaeger Stercorarius parasiticus XXXXXXXX7LC Long-tailed Jaeger Stercorarius longicaudus XXXX XXX7LC

150 –POPULATIONS AND STATUS OF ARCTIC SEABIRDS – 1

Table 1. (continued) Alaska NWT Nunavut Quebec Labrador Greenland Iceland Svalbard Europe Russia Siberia pop'n World IUCN Status

Gulls and Terns Laridae Black-headed Gull Chroicocephalus ridibundus X8LC Mew Gull Larus canus XX X XX7LC G. Black-backed Larus marinus XXXXX 6LC Gull Glaucous Gull Larus hyperboreus XXXXXXXX6LC Iceland Gull Larus glaucoides XX 6LC Thayer's Gull Larus thayeri XX 4LC Amer. Herring Gull Larus smithsonianus XX 6LC Lesser Black-backed Larus fuscus XX XX6LC Gull Herring Gull Larus argentatus XX7LC Vega Gull Larus vegae X6LC Slaty-backed Gull Larus schistisagus XX6 Ivory Gull Pagophila eburnea XX XX 4NT Ross's Gull Rhodostethia rosea XX X5LC Sabine's Gull Xema sabini XXXX X X6LC Black-leg. Kittiwake Rissa tridactyla X XXXXXX8LC Arctic Tern Sterna paradisaea XXXXXXXX7LC Aleutian Tern Onychoprion aleuticus X5LC

Auks Alcidae Little Auk Alle alle X XXXXXX8LC Common Murre Uria aalge X XXXXX7LC Thick-billed Murre Uria lomvia X XXXXXX8LC Razorbill Alca torda XXX X 6LC Black Guillemot Cepphus grylle X XXXXXX7LC Pigeon Guillemot Cepphus columba X6LC Kittlitz's Murrelet Brachyrhamphus brevirostris XX5CE Parakeet Auklet Aethia psittacula XX7LC Crested Auklet Aethia cristatella XX7LC Least Auklet Aethia pusilla XX7LC Atlantic Puffin Fratercula arctica XXXXX 8LC Horned Puffin Fratercula corniculata XX7LC Tufted Puffin Fratercula cirrhata XX7LC

1(orders of magnitude, breeding individuals)

151 –GASTON – past three decades, with trends in the North and 2005–2008, while numbers of Common Pacific and Northwest Atlantic generally pos- Murres decreased abruptly between 1999– itive or stable when trends in the European 2005 after modest increases earlier (Gardars- Arctic were negative and vice versa (Irons et son 2006). Northern Fulmar (Fulmarus al. 2008). By combining population trend data glacialis), Black-legged Kittiwake (Rissa tri- from around the Arctic with information on sea dactyla) and Razorbill also decreased, surface temperature changes (SST) and although some small colonies increased (Gar- decadal-scale climate-ocean oscillations, Irons darsson et al. 2009). et al. (2008) showed that population growth was most often positive where conditions CAUSES OF POPULATION CHANGES remained relatively stable and negative when change, either colder or warmer, was large. The causes of population and range changes This result suggests that not only the direction can rarely be confidently attributed to a single but the magnitude of change may be important source. The decline of Ivory Gulls in the Cana- in determining biological outcomes of climate. dian Arctic illustrates a case where several Trends in different regions switched direction potential contributory causes can be identified: with regime shifts. However, U. lomvia popu- heavy hunting of adults in Greenland (Sten- lations have declined in all regions except the house et al. 2004), high levels of mercury in Canadian eastern Arctic since the 1970s, eggs (Braune et al. 2006) and changes in ice whereas no single global trend can be identi- conditions associated with global warming fied for U. aalge. (Environment Canada 2010). All may have contributed to recent population decline. Only The population of Thick-billed Murres in Cen- where population declines are abrupt and asso- tral West Greenland is much depressed com- ciated with strong environmental signals, can pared to numbers in the early nineteenth causes be confidently assigned. This was the century, as a result of heavy harvesting of case for Common Murre populations in the adults at colonies (Evans and Kampp 1991), as southern Barents Sea in 1985–87 when num- well as drownings in gill-net fisheries (Tull et bers fell by 80% in response to starvation fol- al. 1972) and shows no sign of recovery, with lowing the collapse of the Barents Sea Capelin the population south of Thule District remain- (Mallotus villosus) stock (Anker-Nilssen et al. ing at <20% of historical levels (Kampp et al. 1997). The population subsequently recovered 1994, Merkel et al. 2007, F. Merkel pers. to near its former level (Krasnov et al. 2007). comm. 2010). Numbers in East Greenland, Similarly, an 80% decrease in Lesser Black- although small, have also declined. Similarly, backed Gulls in northern Norway coincided numbers in Novaya Zemlya are considerably with a collapse in the stock of spring spawning lower than in the early twentieth century when Herring (Clupea harengus) (Bustnes et al. the population numbered several million birds. 2010). Currently, there are thought to be in the region of one million breeders (Bakken and Most changes in demography and population Pokrovskaya 2000). In Spitzbergen, numbers status of Arctic seabirds that have been linked of Thick-billed Murres were thought to be sta- with climate changes have, to date, been ble up to the 1990s, but have since decreased, ascribed to causes operating through the food especially in the southern part of the archipel- chain (Harris et al. 2005, Sandvik et al. 2005, ago (CAFF Circumpolar Seabird Working Durant et al. 2004, 2006, Irons et al. 2008). Group, unpubl.) However, a few cases where direct effects have occurred have been documented. Mallory et al. In Iceland, numbers of Thick-billed Murres (2009) reported a wide range of weather- decreased at 7% per year between 1983–1985 related mortalities at Arctic seabird colonies

152 –POPULATIONS AND STATUS OF ARCTIC SEABIRDS – and suggested that some types of mortality, likely. Eventually it may be replaced by U. especially those associated with increases in aalge and other more southern auks. extreme weather events, could create heavier mortality in the future. In northern Hudson Substantial research has been carried out in the Bay in the late 1990s, a combination of warm Barents Sea region and in the Canadian Arctic summer weather and earlier emergence by on concentrations and trends in contaminants, mosquitoes, leading to heavy blood-sucking, especially organohaline compounds and heavy caused the death of some incubating Thick- metals (Braune et al. 2001, Letcher et al. billed Murres through a combination of dehy- 2010). Very high levels of mercury (Braune et dration and hyperthermia. In addition, some al. 2006) and organohaline compounds (Mil- birds left their eggs unattended for periods of jeteig et al. 2009) have been found in the eggs several hours, resulting in many losses to of Ivory Gulls from Canada and Svalbard and predatory gulls (Gaston et al. 2002). These high organohaline concentrations occur also in effects had not been recorded previously in 20 Glaucous Gulls (Larus hyperboreus) from years of observations. Changes in the timing of Svalbard (Bustnes et al. 2003, 2004), perhaps snow-cover and ice-melt affect the availability causing mortality in some cases (Gabrielsen et of breeding sites to crevice, scree and burrow- al. 1995, Sagerup et al. 2009). These species nesting species, such as puffins and Little Auks scavenge marine mammal carcasses, putting (Birkhead and Harris 1985). Such changes in them high up the food chain and hence subject accessibility can result in altered interactions to high biomagnification effects. They may with predators, as observed for Antarctic also frequent garbage dumps around human Petrels (Thalassoica antarctica) by van population centres. Levels of contaminants in Franeker et al. (2001). other species generally do not approach those likely to impact populations (Gabrielsen 2007, Although both species of murre are currently Letcher et al. 2010), except in the case of abundant, many populations have been declin- point-source pollution resulting from industrial ing for several decades (Johnsen et al. 2010). sites (e.g., Kuzyk et al. 2003). Problems facing murres include fisheries interactions, contaminants and oil spills and, in Changes in the timing of seasonal events for some parts of their range, hunting (especially high-latitude marine birds have been identified of U. lomvia). For U. lomvia, changes in the for many southern hemisphere species (Croxall extent and timing of sea-ice cover over the past et al. 2002, Rolland et al. 2010), as well as several decades are leading to changes in phe- some Arctic seabird populations (Gaston et al. nology and reproduction with adverse conse- 2005, Byrd et al. 2008a,b, Moe et al. 2009). quences for nestling growth (Gaston et al. For some Arctic species, reproductive success 2005). These changes seem likely to intensify. is inversely correlated with date of laying, e.g., Levels of some contaminants, especially mer- Little Auks (Moe at al. 2009), but this relation- cury, have increased in murre eggs in the North ship may vary among geographical areas; it is American Arctic since the 1970s, although true for Thick-billed Murres breeding at Prince they remain at sublethal levels (Braune et al. Leopold Island, Nunavut, but not for the same 2001). If climate change leads to increased species breeding in northern Hudson Bay shipping and oil and gas exploitation in Arctic (Gaston et al. 2005). The importance of timing waters, the increased risk of spills would also of breeding in determining the dynamics of pose a potential hazard for murres, which are Arctic seabird populations is supported by a extremely susceptible to mortality from oil correlation found between colony size and the pollution (Wiese and Robertson 2004). In the timing of sea ice withdrawal in adjacent waters long-term, range contraction of U. lomvia in for Thick-billed Murres in Greenland (Laidre response to the retreat of Arctic sea ice appears et al. 2008).

153 –GASTON –

Mismatching of breeding initiation with the 2007, Sheffield Guy 2009). These changes seasonal peak of food availability may be a were associated with a warming of the adja- common phenomenon among seabirds con- cent surface waters and a retreat of winter sea fronted with rapidly changing seasonal timing ice. Similarly, in Iceland, the diet of most (Bertram 2001, Wilhelm et al. 2008, Watanuki seabirds switched from sandlance to other et al. 2009). It has been identified as a likely fishes in the 2000s (Gardarsson 2006), a cause of reduced nestling growth for Thick- change also observed in boreal waters of the billed Murres in northern Hudson Bay (Gas- North Sea (Wanless et al. 2005). This diet ton et al. 2009), as well as accounting for change was contemporary with declines in some of the variation in reproductive success most seabird populations. of Black-legged Kittiwakes and Common Murres in sub-Arctic Alaska (Suryan et al. Many seabirds are very conservative in their 2006, Schultz et al. 2009) and Newfoundland breeding sites, returning faithfully to large (Wilhelm et al. 2008). colonies that, in some cases, have been in exis- tence for millennia (Gaston and Donaldson Changes in seabird diets, both from year to 1996). If climate change alters environmental year and over decades, have been reported conditions around such colonies it is unlikely from many sites. Diet switching is likely a that a mass exodus will take place in search of fairly routine aspect of seabird biology (e.g., new colony locations. There are examples of Montevecchi and Myers 1995, 1997, Barrett large colonies suffering repeated reproductive 2002). At Coats Island, northern Hudson Bay, failure over many years without any substan- Thick-billed Murres switched from feeding tial emigration (e.g., Atlantic Puffins at Rost, their chicks predominantly the ice-associated Norway reared few chicks between 1969– Arctic Cod (Boreogadus saida) to the more 1982, Anker-Nilssen and Rostad 1993). How- sub-Arctic Capelin in the mid-1990s (Gaston ever, parasites and predators may be more et al. 2003). The change was associated with mobile in response to climate change and may an advance in the date of sea-ice clearance in initiate or expand their activities at new sites. the region. Some examples of such expansions have already been observed, with an increase in the Not all prey are equally suitable, especially for incidence of tapeworms in alcids in Labrador rearing nestling birds, and some prey switches and Greenland since the 1960s (Bin Muzaffar can result in reduced productivity among 2009) and the appearance of the parasitic tick seabirds (Litzow et al. 2002, Wanless et al. (Ixodes uriae) on murres in Svalbard after 2005, Gremillet et al. 2008). In the southwest 2000 (Coulson et al. 2009). The implications Barents Sea in recent decades Herring has of these parasite range expansions are not yet come to dominate over Capelin as a forage clear but adverse consequences for the seabird fish. This change has coincided with a decline populations involved are possible. in numbers of breeding Back-legged Kitti- wakes (–8% per year after 1995). Apparently Currently Golden Eagles (Aquila chrysaetos) Herring is not as satisfactory as Capelin as and White-tailed Sea-eagles (Haliaeetus albi- food for kittiwakes (Barrett 2007). At the Pri- cilla), both of which cause disruption to nest- bilof Islands, Sinclair et al. (2008) also ing seabirds, only reach the fringes of the observed a reduction in the proportion of Arctic. Their northward spread could create Capelin in Black-legged Kittiwake and Thick- problems for gulls, murres and other open- billed Murre diets between the 1980s and nesting seabirds. Increasing predation of birds 2000s, while changes in the zooplankton diet and their nests by Polar Bears (Ursus marinus) of Least Auklets (Aethia pusilla) was also has also been observed, probably as a result of observed over the same period (Springer et al. the bears coming ashore earlier in the season

154 –POPULATIONS AND STATUS OF ARCTIC SEABIRDS –

(Rockwell and Gormezano 2009, Smith et al. ACKNOWLEDGMENTS 2010). This could affect especially accessible species such as Little Auks (Stempniewicz Thanks to the many students, contractors, col- 2007). Because of the potential for alternative laborators and volunteers who have assisted prey, it is extremely difficult to predict how with seabird research and monitoring in the seabird populations will respond to changes in Canadian Arctic. Our efforts would not have predator distributions. been successful without the financial and logistic support of Natural Resources Canada (Polar Continental Shelf Program), the CONCLUSIONS Nunavut Research Institute, Indian and North- ern Affairs Canada (Northern Contaminants The Arctic is an important area for marine bird Program), and various branches of Environ- diversity and endemism. Most Arctic seabird ment Canada (Canadian Wildlife Service and populations for which information is available Science and Technology Branch). For informa- over several decades have shown negative tion and advice on literature sources, I thank trends in recent years. These trends are super- Carsten Egevang, Maria Gavrilov, Arnthor imposed on a situation where several impor- Gardarsson, Grant Gilchrist, David Irons, tant populations were substantially depressed Mark Mallory, Freydig Vigfúsdóttir, and mem- by anthropogenic mortality, compared with bers of the C-bird working group of CAFF. numbers in the first half of the twentieth century (especially Thick-billed Murres in Green- land and Novaya Zemlya). LITERATURE CITED

Only a few instances are available where recent trends can be traced to particular causes ANKER-NILSSEN, T., AND O. D. ROSTAD. 1993. but stressors include fisheries activities, pollu- Census and monitoring of puffins Frater- tion, and climate change. The last, especially cula arctica on Rost, North Norway, 1979– as manifested in changes in the timing of the 1988. Ornis Scandinavica 24:1–9. open water season, is affecting the timing of ANKER-NILSSEN, T., R. T. BARRETT, AND J. V. seasonal events in marine ecosystems and this KRASNOV. 1997. Long- and short-term is affecting the optimal timing of breeding, responses of seabirds in the Norwegian and especially in low-Arctic areas. Changes in ice Barents Seas to changes in stocks of prey conditions, especially, are likely to have far- fish. Forage Fishes in Marine Ecosystems– reaching and potentially irreversible conse- Lowell Wakefield Fisheries Symposium quences. These changes are also encouraging Series, no. 14:683–698. the northward expansion of sub-Arctic species, BAKKEN, V., AND I. V. POKROVSKAYA. 2000. although such changes in range are relatively Thick-billed Murre. Pages 119–124 in T. small, as yet. Changes in the distributions of Anker-Nilssen, V. Bakken, H. Strøm, A.N. predators and parasites have also been noted Golovkin, V.V. Bianki, and I.P. Tatarinkova and these may have important consequences (Eds.). The Status of Marine Birds Breed- for Arctic seabirds. Because of the number of ing in the Barents Sea Region. Norsk Polar- Arctic endemic seabird taxa, the decline of institutt, Tromsø, Norway. Arctic marine birds presages a significant loss BARRETT, R. T. 2002. Atlantic Puffin Frater- of global biodiversity. cula arctica and Common Guillemot Uria aalge chick diet and growth as indicators of fish stocks in the Barents Sea. Marine Ecol- ogy Progress Series 230:275–287. BARRETT, R. T. 2007. Food web interactions in the southwestern Barents Sea: Black-

155 –GASTON –

legged Kittiwakes Rissa tridactyla respond of piscivorous seabirds in the Pribilof negatively to an increase in Herring Clupea Islands: A 30-year perspective. Deep Sea harengus. Marine Ecology Progress Series Research Part II—Topical Studies in 349:269–276. Oceanography 55:1846–1855. BERTRAM, D. F., D. L. MACKAS, AND S. M. BYRD, G. V., W. J. SYDEMAN, H. M. RENNER, MCKINNELL. 2001. The seasonal cycle AND S. MINOBE. 2008b. Responses of pis- revisited: Interannual variation and ecosys- civorous seabirds at the Pribilof Islands to tem consequences. Progress in Oceanogra- ocean climate. Deep Sea Research Part II— phy 49:283–307. Topical Studies in Oceanography 55:1856– BIN MUZAFFAR, S. 2009. Helminths of murres 1867. (Alcidae: Uria spp.): Markers of ecological CAFF CIRCUMPOLAR SEABIRD GROUP. 1996. change in the marine environment. Journal Murre conservation in the circumpolar Arc- of Wildlife Diseases 45:672–683. tic. Poster published by Committee on Arc- BIRKHEAD, T. R., AND M. P. HARRIS. 1985. tic Flora and Fauna, Environment Canada, Ecological adaptations for breeding in the Canada. Atlantic Alcidae. Pages 205–261 in D. N. CHAULK, K. G., J. G. ROBERTSON, AND W. A. Nettleship and T. R. Birkhead (Eds.). The MONTEVECCHI. 2004. Breeding range Atlantic Alcidae. Academic Press, London, update for three seabird species in UK. Labrador. Northeastern Naturalist 11:479– BRAUNE, B. M., G. M. DONALDSON, AND K. A. 485. HOBSON. 2001. Contaminant residues in COULSON, S. J., E. LORENTZEN, H. STROM, AND seabird eggs from the Canadian Arctic. Part G. W. GABRIELSEN. 2009. The parasitic tick I. Temporal trends 1975–1998. Environ- Ixodes uriae (Acari: Ixodidae) on seabirds mental Pollution 114:39–54. from Spitsbergen, Svalbard. Polar Research BRAUNE B. M., M. L. MALLORY, AND H. G. 28:399–402. GILCHRIST. 2006. Elevated mercury levels CROXALL, J. P., P. N. TRATHAN, AND E. J. MUR- in a declining population of Ivory Gulls in PHY. 2002. Environmental Change and the Canadian Arctic. Marine Pollution Bul- Antarctic Seabird Populations. Science letin 52:978–982. 297:1510–1514. BUSTNES, J. O., T. ANKER-NILSSEN, AND S. H. DURANT, J. M., T. ANKER-NILSSON, D. O. LORENTSEN. 2010. Local and large-scale HJERMANN, AND N. C. STENSETH. 2004. climatic variables as predictors of the Regime shifts in the breeding of an Atlantic breeding numbers of endangered Lesser Puffin population. Ecology Letters 7:388– Black-backed Gulls on the Norwegian 394. Coast. Journal of Ornithology 151:19–26. DURANT, J. M., T. ANKER-NILSSON, AND N. C. BUSTNES, J. O., K. E. ERIKSTAD, J. U. SKAARE, STENSETH. 2006. Ocean climate prior to V. B AKKEN, AND F. MEHLUM. 2003. Ecolog- breeding affects the duration of the nestling ical effects of organochlorine pollutants in period in the Atlantic Puffin. Biology Let- the Arctic: A study of the Glaucous Gull. ters 2: 628–631. Ecological Applications 13:504–15. ELDERS CONFERENCE ON CLIMATE CHANGE. BUSTNES, J. O., S. A. HANSSEN, I. FOLSTAD, K. 2001. Cambridge Bay, Nunavut. Nunavut E. ERIKSTAD, D. HASSELQUIST, AND J. U. Tunngavic Inc., Iqaluit, Canada. SKAARE. 2004. Immune function and ENVIRONMENT CANADA. 2010. Recovery Strat- organochlorine pollutants in arctic breeding egy for the Ivory Gull (Pagophila eburnea) Glaucous Gull. Archives of Environmental in Canada [Draft]. Species at Risk Act Contamination and Toxicology 47:530–41. Recovery Strategy Series. Environment BYRD, G. V., J. A. SCHMUTZ, AND H. M. REN- Canada, Ottawa, Canada. NER. 2008a. Contrasting population trends

156 –POPULATIONS AND STATUS OF ARCTIC SEABIRDS –

EVANS, P. G. H., AND K. KAMPP. 1991. Recent GASTON, A. J., AND K. WOO. 2008. Razorbills changes in Thick-billed Murre populations Alca torda follow subarctic prey into the in West Greenland. Canadian Wildlife Canadian Arctic: Colonization results from Service Occasional Paper 69:7–14. climate change? Auk 125:939–942. FREUCHEN, P., AND F. SALMONSEN. 1958. The GASTON, A. J., D. F. BERTRAM, A. W. BOYNE, Arctic Year. G. P. Putnam, New York, USA. J. W. CHARDINE, G. DAVOREN, A.W. DIA- FULLER, E. 2000. Extinct Birds. Oxford Uni- MOND, A. HEDD, W. A. MONTEVECCHI, J. M. versity Press, Oxford, UK. HIPFNER, M. J. F. LEMON, M. L. MALLORY, FURNESS, R. W. 1987. The Skuas. T. and A. D. J-F. RAIL, AND G. J. ROBERTSON. 2009. Poyser, Calton, UK. Changes in Canadian seabird populations GABRIELSEN, G. W. 2007. Levels and effects of and ecology since 1970 in relation to persistent organic pollutants in arctic ani- changes in oceanography and food webs. mals. Pages 377–412 in J. B. Orbaek, R. Environmental Reviews 17:267–286. Kallenborn, I. Tombre, E. N. Hegseth, S. GASTON, A. J., D. K. CAIRNS, R. D. ELLIOT, Falk-Petersen, and A. H. Hoel (Eds.). Arc- AND D. G. NOBLE. 1985. A natural history tic-Alpine Ecosystems and People in a of Digges Sound. Canadian Wildlife Serv- Changing Environment. Springer Verlag, ice Report Series No. 46:1–63. Berlin, Germany. GASTON, A. J., H. G. GILCHRIST, AND J. M. GABRIELSEN, G. W., J. U. SKAARE, A. POLDER, HIPFNER. 2005. Climate change, ice condi- AND V. B AKKEN. 1995. Chlorinated hydro- tions and reproduction in an Arctic nesting carbons in Glaucous Gulls (Larus hyper- marine bird: The Thick-billed Murre (Uria boreus) in the southern part of Svalbard. lomvia, L.). Journal of Animal Ecology Science of the Total Environment 74:832–841. 160/161:337–346. GASTON, A. J., J. M. HIPFNER, AND D. CAMP- GARDARSSON, A. 2006. Nýlegar breytingar á BELL. 2002. Heat and mosquitoes cause fjölda íslenskra bjargfugla [Recent changes breeding failures and adult mortality in an in cliff-breeding seabirds in Iceland]. Bliki Arctic-nesting seabird. Ibis 144:185–191. 27:13–22. GASTON, A. J., K. WOO, AND J. M. HIPFNER. GARDARSSON, A., G. A. GUDMUNDSSON, K. 2003. Trends in forage fish populations in LILLIENDAHL, AND F. VIGFÚSDÓTTIR. 2009. northern Hudson Bay since 1981, as deter- Status of cliff-breeding seabirds in Iceland mined from the diet of nestling Thick- in 2005–08. Poster for Seabird group Con- billed Murres Uria lomvia. Arctic ference, March 2009, Brugge, Belgium. 56:227–233. GASTON, A. J., AND G. DONALDSON. 1996. Peat GAVRILO, M. V. 2010. Increasing importance of deposits and Thick-billed Murre colonies in Siberian shelf seas for polar marine top Hudson Strait and northern Hudson Bay: predators under conditions of the modern Clues to post glacial colonization of the warming Arctic. Presentation at Oslo Sci- area by seabirds. Arctic. 48:354–358. ence Conference, 2010, Oslo, Norway. GASTON, A. J., AND I. L. JONES. 1998. The GILCHRIST, H. G., AND M. L. MALLORY. 2005. Auks – Alcidae. Oxford University Press, Declines in the abundance and distribution Oxford, UK. of the Ivory Gull (Pagophila ebernea) GASTON, A. J., AND A. SHOJI. 2010. The breeding in Arctic Canada. Biological Con- Ancient Murrelet Synthliboramphus servation 121:303–309. antiquus. In A. Poole and F. Gill (Eds.). GILCHRIST, H. G., M. L. MALLORY, AND F. The Birds of North America, no. 132 MERKEL. 2005. Can local ecological [revised] Academy of Natural Sciences, knowledge contribute to wildlife manage- Philadelphia, and American Ornithologists' ment? Case studies of migratory birds. Union, Washington, DC, USA. Ecology and Society 10(1):20. [Online.]

157 –GASTON –

Available at http://www.ecologyandsociety Kittiwakes (Rissa tridactyla), Common .org/ vol10/iss1/art20/ Guillemots (Uria aalge) and Brunnich's GILCHRIST, H. G., H. STROM, M. V. GAVRILO, Guillemots (U-lomvia) in Murman, north- AND A. MOSBECH. 2008. International ivory west Russia, and Varanger, north-east Nor- gull conservation strategy and action plan. way. Polar Research 26:113–117. CAFF International Secretariat, Circumpo- KULETZ, K. J., S. W. STEPHENSEN, D. B. IRONS, lar Seabird Group, CAFF Technical Report E. A. LABRUNSKI, AND K. M. BRENNEMAN. no. 18. 2003. Changes in the distribution and abun- GILG, O., D. BOERTMANN, F. MERKEL, A. dance of Kittlitz’s Murrelets Brachyram- AEBISCHER, AND B. SABARD. 2009. Status phus brevirostris relative to glacial of the endangered Ivory Gull, Pagophila recession in Prince William Sound, Alaska. eburnea, in Greenland. Polar Biology Marine Ornithology 31:133–140. 32:1275–1286. KUZYK, Z. Z. A., N. M. BURGESS, J. P. STOW, GREMILLET, D., L. PICHEGRU, G. KUNTZ, A. G. AND G. A. FOX. 2003. Biological effects of WOAKES, S. WILKINSON, R. J. M. CRAW- marine PCB contamination on Black FORD, AND P. G. RYAN. 2008. A junk-food Guillemot nestlings at Saglek, Labrador: hypothesis for gannets feeding on fishery Liver biomarkers. Ecotoxicology 12:183– waste. Proceedings of the Royal Society B 197. 275:1149–1156. LAIDRE, K. L., M. P. HEIDE-JØRGENSEN, J. NYE- HARRIS, M. P., T. ANKER-NILSSEN, R. H. LAND, A. MOSBECH, AND D. BOERTMANN. MCCLEERY, K. E. ERIKSTAD, D. N. SHAW, 2008. Latitudinal gradients in sea ice and AND V. G ROSBOIS. 2005. Effect of wintering primary production determine Arctic area and climate on the survival of adult seabird colony size in Greenland. Proceed- Atlantic Puffins Fratercula arctica in the ings of the Royal Society B 275:2695– eastern Atlantic. Marine Ecology Progress 2702. Series 297:283–296. LETCHER, R. J., J. O. BUSTNES, R. DIETZ, B. M. IRONS, D. B., T. ANKER-NILSSEN, A .J. GASTON, JENSSEN, E. H. JØRGENSEN, C. SONNE, J. G. V. BYRD, K. FALK, H. G. GILCHRIST, M. VERREAULT, M. M. VIJAYAN, AND G. W. HARIO, M. HJERNQUIST, Y. V. K RASNOV, A. GABRIELSEN. 2010. Exposure and effects MOSBECH, B. OLSEN, A. PETERSEN, J. REID, assessment of persistent organohalogen G. J. ROBERTSON, H. STROM, AND K. D. contaminants in Arctic wildlife and fish. WOHL. 2008. Magnitude of climate shift Science of the Total Environment determines direction of circumpolar seabird 408:2995–3043. population trends. Global Change Biology LITZOW, M. A., J. F. PIATT, A. K. PRICHARD, 14:1455–1463. AND D. D. ROBY. 2002. Response of Pigeon JOHNSEN, K. I., B. ALFTHAN, L. HISLOP, AND J. Guillemots to variable abundance of high- F. SKAALVIK (Eds.). 2010. Protecting Arctic lipid and low-lipid prey. Oecologia Biodiversity. United Nations Environment 132:286–295. Programme, GRID-Arendal, Norway. MALAURIE, J. 1985. The Last Kings of Thule. KAMPP, K., D. N. NETTLESHIP, AND P. G. H. University of Chicago Press, Chicago, EVANS. 1994. Thick-billed Murres of USA. Greenland: Status and prospects. Pages MALLORY, M. L., A. J. GASTON, AND H. G. 133–154 in D. N. Nettleship, J. Burger, and GILCHRIST. 2009. Sources of breeding sea- M. Gochfeld (Eds.). Seabirds on Islands: son mortality in Canadian Arctic seabirds. Threats, Case Studies, and Action Plans. Arctic 62:333–341. Birdlife International, Cambridge, UK. MEHLUM, F., AND V. B AKKEN. 1994. Seabirds KRASNOV, Y. V., R. T. BARRETT, AND N. G. in Svalbard (Norway): Status, recent NIKOLAEVA. 2007. Status of Black-legged changes and management. Pages 155–171

158 –POPULATIONS AND STATUS OF ARCTIC SEABIRDS –

in D. N. Nettleship, J. Burger, and M. dae. Pages 55–154 in D. N. Nettleship, and Gochfeld (Eds.). Seabirds on Islands: T. R. Birkhead (Eds.). The Atlantic Alcidae. Threats, Case Studies and Action Plans. Academic Press, London. Birdlife International, Cambridge, UK. NORDERHAUG, M., E. BRUN, AND G. U. MERKEL F. R., A. L. LABANSEN, AND L. WIT- MØLLEN. 1977. Seabird Resources of the TING. 2007. Monitering af lomvier og rider Barents Sea. Norsk Polarinstitutt Med- i Qaanaaq Kommune. 2006. Technical delelser 104. Norsk Polarinstitutt, Oslo, Report no. 69, Greenland Institute of Nat- Norway. ural Resources, Pinngortitaleriffik. OLSEN, K. M., AND H. LARSSON. 2003. Gulls of MILJETEIG, C., H. STROM, M. V. GAVRILO, A. Europe, Asia and North America. Christo- VOLKOV, B. M. JENSSEN, AND G. W. pher Helm, London, UK. GABRIELSEN. 2009. High levels of contam- PETERSEN, A., AND S. THORSTENSEN. 2004. inants in Ivory Gull Pagophila eburnea Vöktun stormmamfssofnsins i Eyjafirdi eggs from the Russian and Norwegian Arc- 1980–2000. Natturufraedingurinn 72:144– tic. Environmental Science and Technology 154. 43:5521–5528. PRICKETT, H. 1611. Description of Digges MOE, B., L. STEMPNIEWICZ, D. JAKUBAS, F. Island. Pages 98–135 in G. M. Asher (Ed.). ANGELIER, O. CHASTEL, F. DINESSEN, G. W. Henry Hudson the Navigator. The Hakluyt GABRIELSEN, F. HANSSEN, N. J. Society, London, 1860. KARNOVSKY, B. RØNNING, J. WELCKER, K. RENAUD, W. E., P. L. MCLAREN, AND S. R. WOJCZULANIS-JAKUBAS, AND C. BECH. JOHNSON. 1982. The Dovekie Alle alle as a 2009. Climate change and phenological spring migrant in Lancaster Sound and responses of two seabird species breeding western Baffin Bay. Arctic 35:118–125. in the high-Arctic. Marine Ecology ROCKWELL, R. F., AND I. J. GORMEZANO. 2009. Progress Series 393:235–246. The early bear gets the goose: climate MOLINE, M. A., N. J. KARNOVSKY, Z. BROWN, change, Polar Bears and Lesser Snow G. J. DIVOKY, T. K. FRAZER, C. A. JACOBY, Geese in western Hudson Bay. Polar Biol- J. J. TORRES, AND W. R. FRASER. 2008. High ogy 32:539–547 latitude changes in ice dynamics and their ROLLAND, V., H. WEIMERSKIRCH, AND C. BAR- impact on polar marine ecosystems. Annals BRAUD. 2010. Relative influence of fish- of the New York Academy of Sciences eries and climate on the demography of 1134:267–319. four albatross species. Global Change Biol- MONTEVECCHI, W. A., AND R .A. MYERS. 1995. ogy 16:1910–1922. Prey harvests of seabirds reflect pelagic SAGERUP, K., L. B. HELGASON, A. POLDER, H. fish and squid abundance on multiple spa- STRØM, T. D. JOSEFSEN, J. U. SKÅRE, AND tial and temporal scales. Marine Ecology G. W. GABRIELSEN. 2009. Persistent organic Progress Series 117:1–9. pollutants and mercury in dead and dying MONTEVECCHI, W. A., AND R. A. MYERS. 1997. Glaucous Gulls (Larus hyperboreus) at Centurial and decadal oceanographic influ- Bjørnøya (Svalbard). Science of the Total ences on changes in gannet populations and Environment 407:6009–6016. diets in the north-west Atlantic: Implica- SALOMONSEN, F. 1950. Grønlands Fugle. tions for climate change. ICES Journal of Munkssgaard, Copenhagen, Denmark. Marine Science 54:608–614. SANDVIK, H., K. E. ERIKSTAD, R. T. BARRETT, NELSON, E. W. 1983. The Eskimo about Bering AND N. G. YOCCOZ. 2005. The effect of cli- Strait. Smithsonian Institution Press, Wash- mate on adult survival in five species of ington, DC, USA. North Atlantic seabirds. Journal of Animal NETTLESHIP, D. N., AND P. G. H. EVANS. 1985. Ecology 74:817–831. Distribution and status of the Atlantic Alci-

159 –GASTON –

SCHULTZ, M. T., J. F. PIATT, A. M. A. HARDING, TUCK, L. M. 1961. The Murres. Canadian A. B. KETTLE, AND T. I. VAN PELT. 2009. Wildlife Service, Ottawa, Canada. Timing of breeding and reproductive per- TULL, C. E., A. W. MAY, AND P. G ERMAIN. formance in murres and kittiwakes reflect 1972. Mortality of Thick-billed Murres in mismatched seasonal prey dynamics West Greenland salmon fishery. Nature Marine Ecology Progress Series 393:249– 237:42–44. 260. USPENSKI, S. M. 1956. “The bird bazaars of SHEFFIELD GUY, L. M., D. D. ROBY, A. E. Novaya Zemlya.” Canadian Wildlife Serv- GALL, D. B. IRONS, AND I. C. ROSE. 2009. ice Translations of Russian Game Reports, The influence of diet and ocean conditions vol. 4, Ottawa, Canada. on productivity of auklets on St. Lawrence VAN FRANEKER, J. A., J. C. S. CREUWELS, W. Island, Alaska. Marine Ornithology VAN DER VEER, S. CLELAND, AND G. 37:227–236. ROBERTSON. 2001. Unexpected effects of SINCLAIR, E. H., L. S. VLIETSTRA, D. S. JOHN- climate change on the predation of Antarc- SON, T. K. ZEPPELIN, G. V. BYRD, A. M. tic Petrels. Antarctic Science 13:430–439. SPRINGER, R. R. REAM, AND G. L. HUNT, JR. VAURIE, C. 1965. The Birds of the Palearctic 2008. Deep Sea Research Part II—Topical Fauna—non-Passeriformes. H. F. and G. Studies in Oceanography 55:1897–1918. Witherby, London, UK. SMITH, P. A., K. H. ELLIOTT, A. J. GASTON, AND VOOUS, K. H. 1949. The morphological, H. G. GILCHRIST. 2010. Early ice clearance anatomical and distributional relationships leads to increased predation on breeding of the Arctic and Antarctic fulmars (Aves, birds by Polar Bears. Polar Biology Procellariidae). Ardea 37:113–122. 33:1149–1153. WANLESS, S., M. P. HARRIS, P. REDMAN, AND J. SPRINGER, A. M., G. V. BYRD, AND S. J. IVER- R. SPEAKMAN. 2005. Low energy values of SON. 2007. Hot oceanography: Planktivo- fish as a probable cause of a major seabird rous seabirds reveal ecosystem responses to breeding failure in the North Sea. Marine warming of the Bering Sea. Marine Ecol- Ecology Progress Series 294:1–8. ogy Progress Series 352:289–297. WATANUKI, Y., M. ITO, T. DEGUCHI, AND S. STEMPNIEWICZ, L. 2007. The Polar Bear Ursus MINOBE. 2009. Climate-forced seasonal maritimus feeding in a seabird colony in mismatch between the hatching of Rhinoc- Frans Josef Land. Polar Biology 12:33–36. eros Auklets and the availability of STENHOUSE, I. J., G. J. ROBERTSON, AND H. G. anchovy. Marine Ecology Progress Series GILCHRIST. 2004. Recoveries and survival 393:259–271. rate of Ivory Gulls banded in Nunavut. WIESE, F., AND G. J. ROBERTSON. 2004. Assess- Waterbirds 27:486–492. ing seabird mortality from chronic oil dis- STENHOUSE, I. J., S. STUDEBAKER, AND D. charges at sea. Journal of Wildlife ZWIEFELHOFER. 2008. Kittlitz’s Murrelet Management 68:627–638. Brachyramphus brevirostris in the Kodiak WILHELM, S. I., C. J. WALSH, AND A. E. Archepelago, Alaska. Marine Ornithology STOREY. 2008. Time budgets of Common 36:59–66. Murres vary in relation to changes in SURYAN, R. M., D. B. IRONS, E. D. BROWN, P. inshore Capelin availability. Condor G. R. JODICE, AND D. D. ROBY. 2006. Site- 110:316–324. specific effects on productivity of an upper trophic-level marine predator: Bottom-up, top-down, and mismatch effects on reproduction in a colonial seabird. Progress in Oceanography 68:303–328.

160