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The lesser black-backed gull, Larus fuscus, in Greenland.

ABSTRACT. The lesser black-backed gull (Larus fuscus) became a frequent visitor in Greenland in the mid-1980s. Breeding was confirmed in 1990, and today the species is a common breeder in at least two areas in Southwest Greenland between 60[degrees] and 66[degrees] N. The current breeding population is estimated at more than 700 pairs. Even though the colonization of Greenland by this species may be primarily the result of an increasing source population in Northwest Europe, climate change most likely will facilitate its expansion farther north in Greenland and probably also farther west to eastern Canadian coasts.

Key words: lesser black-backed gull, Larus fuscus, Greenland, range expansion, North Atlantic

RESUME. Le goeland brun (Larus fuscus) est un visiteur frequent du Greenland depuis le milieu des annees 1980. La reproduction de cette espece y a d'ailleurs ete confirmee en 1990 et de nos jours, elle se reproduit regulierement dans au moins deux regions du sud-ouest du Groenland, entre 60[degrees] et 66[degrees] N. En ce moment, la population de reproduction est evaluee a plus de 700 paires. Bien que la colonisation du Groenland par cette espece puisse etre attribuable a une population source de plus en plus grande dans le nord-ouest de l'Europe, le changement climatique facilitera vraisemblablement son foisonnement plus au nord du Groenland et probablement plus a 1'ouest vers la cote est canadienne.

Mots cles: goeland brun, Larus fuscus, Groenland, expansion de l'aire de repartition. Atlantique Nord

Traduit pour la revue Arctic par Nicole Giguere.

INTRODUCTION

The bird fauna of Greenland is young, as all the breeding birds are supposed to be post-glacial immigrants that found their way to this large island during approximately the last 10000 years (Bennike, 1999). Immigration is ongoing, and many bird niches still seem to be unoccupied in Greenland, judging from the low diversity there compared to the adjacent Baffin Island or other Arctic areas (Boertmann, 1994, 1999).

In recent decades, two bird species have immigrated to Greenland and established large and viable populations. Canada geese (Branta canadensis) have moved in from the North American continent, and the current breeding population numbers probably several thousand pairs. This invasion has been the focus of several studies because of concern about possible effects of interspecific competition on the indigenous and discrete population of Greenland white-fronted geese (Anser albifrons flavirostris) (Fox et al., 1996; Boertmann and Egevang, 2002; Kristiansen and Jarrett, 2002). However, the establishment in Greenland of the lesser black-backed gull (Larus fuscus), which occurred almost simultaneously, has gone unnoticed.

The lesser black-backed gull has shown a remarkable population increase and range expansion in northwest Europe since the early 1900s (Cramp and Simmons, 1983). The increase has accelerated in Iceland, Britain, and Ireland in recent decades (Calladine, 2004; A. Petersen, pers. comm. 2005), coinciding with an increase of vagrant lesser black-backed gulls along the Atlantic borders of the Nearctic-Greenland, Canada, and the United States (Boertmann, 1994; Post and Lewis, 1995). Breeding was confirmed in Greenland in 1990, and today lesser black-backed gulls breed abundantly in the Southwest Greenland region. Recently breeding has also occurred in the high Arctic as far north as 74[degrees] N. This paper documents the development of the lesser black-backed gull breeding population in Greenland.

SOURCES

Records of lesser black-backed gulls in Greenland have been published in general avifaunistic works published through the years (H[empty set]rring and Salomonsen, 1941; Salomonsen, 1950, 1967; Boertmann, 1994), and many records were also published in reports from the Rarity Committee of the Danish Ornithological Society (Olsen, 1987, 1989, 1991). Lesser black-backed gulls were observed during extensive ship- and aircraft-based surveys for breeding and moulting seabirds in West and East Greenland over the past 15 years (Boertmann and Mosbech, 1992, 1999; Boertmann, 1994, 2004; Boertmann et al., 1996; Gilg, 2005). Long-term monitoring projects in Northeast Greenland have also recorded lesser black-backed gulls (Hansen and Meltofte, 2006). In addition to these published records, the documentation reported here includes my own observations and many observations reported directly to me. Negative information is also relevant in this context, and I have been on the lookout for the species during all my research in Greenland since 1974.

Breeding records were accepted if nests with eggs or chicks were reported or if adults were observed feeding chicks. Records of territorial birds without location of nests or chicks are treated here as suspected breeding.

OBSERVATIONS

Until 1984, only six records of lesser black-backed gulls were known from Greenland, all of them from the west coast (Table 1). The species was indeed a rare bird then: in the summer of 1974, I surveyed extensive coastal areas of Southwest Greenland between 60[degrees] and 64.5[degrees] N without recording a single sighting (Boertmann, 1979). However, the observations of six adults off the southern tip of Greenland in 1966 (Brown, 1968) were notable exceptions.

From 1985 onwards, numbers of records increased rapidly (Table 1). The first breeding record was documented in 1990, on the same island in South Greenland where breeding had been suspected as early as 1986 and again in 1989 (Table 1). In 1992, the entire coastal region between Paamiut and Disko Bay (62[degrees] to 69[degrees] N) was surveyed for breeding seabirds, and lesser black-backed gulls were found at two sites (Table 1). No breeding gulls were observed during that survey in the extensive archipelagos to the west of Nuuk (64[degrees] N), where six years later I observed large flocks of the species (up to about 50 individuals) with several breeding pairs. Today, however, the region has several breeding colonies holding up to 28 pairs, both in the archipelagoes and in the fjordland (C. Egevang, pers. comm. 2006). In 2003, during another extensive survey for breeding seabirds covering the South Greenland region (60-61[degrees] N), breeding lesserblack, backed gulls were recorded on at least 20 islands and in colonies numbering up to 350 pairs, for a total of more than 600 pairs (Table 1).
TABLE 1. Records of lesser black-backed gull in Greenland.
Observations of non-breeding birds in recent years are not included.

Year       Latitude            Status         Note          Source

West Greenland

1850       60[degrees]         ZMUC  (1)      immature      Boertmann.
                               specimen                     1994

1920       64[degrees]         ZMUC           Adult         Horring and
                               specimen                     Salomonsen,
                                                            1941

1961       64[degrees]         ZMUC           Immature      Salomonsen,
                               specimen       (2)           1967

1966       59[degrees]         sight          6 offshore    Brown. 1968
                               records        sight
                                              records of
                                              adults

1971       70[degrees]         specimen       Adult         Boertmann,
                               (private                     1994
                               collection)

1979       67[degrees]         sight record   Adult         Fox and
                                                            Stroud,
                                                            1981

1985       61-64.5[degrees]    many sight     adults, many  Pers.
                               records        in pairs and  obser.;
                                              flocks        Olsen, 1987

1986       61-64.5[degrees]    many sight     adults, many  Pers.
                               records        in pairs and  obser.
                                              flocks

1986       60.5[degrees]       suspected      territorial   K. Falk,
                               breeding (3)   pair          pers. comm.
                                                            1991

1988       61[degrees]         sight record   Adult         Olsen, 1989

1988       69[degrees]         sight record   Adult         Olsen. 1991

1989       60.5[degrees]       suspected      territorial   K. Falk,
                               breeding (3)   pair          pers. comm.
                                                            1991

1990       60.5[degrees]       confirmed      one pair      Frich and
                               breeding (3)                 Nordbjerg,
                                                            1992

1991       60.5[degrees]       confirmed      one pair      Frich and
                               breeding (3)                 Nordbjerg,
                                                            1992

1992       66[degrees]         confirmed      nest with     Boertmann
                               breeding       eggs          and Mosbech,
                                                            1999

1992       67[degrees]         probable       territorial   Boertmann
                               breeding       pair          and Mosbech,
                                                            1999

1994       60.5[degrees]       probable       2-3 pairs     K. Falk,
                               breeding (3)                 pers. comm.
                                                            1995

1994       64[degrees]         sight record   Juvenile,     Pers.
                                              recently      obser.
                                              Hedged

1998       63-65[degrees]      probable       many pairs    Pers.
                               breeding       seen          obser.

2003       60.5[degrees]       probable       > 10 pairs    K. Falk,
                               breeding (3)                 pers. comm.
                                                            2004

2003       64.5[degrees]       common         [greater      C. Egevang,
                               breeder        than or       pers. comm.
                                              equal to] 5   2006
                                              colonies, in
                                              total
                                              [greater
                                              than or
                                              equal to]60
                                              pairs

2003       60-61[degrees]      common         [greater      Boertmann.
                               breeder        than or       2004
                                              equal to]22
                                              colonies, in
                                              total
                                              [greater
                                              than or
                                              equal to]600
                                              pairs

2006       63.5-64.5[degrees]  common         [greater      C. Egevang,
                               breeder        than or       pers. comm.
                                              equal to]11   2006
                                              colonies

2007       65[degrees]         probable       territorial   Pers.
                               breeding       pair          obser.

East Greenland

1969       70.5[degrees]       sight record   Adult         Boertmann,
                                                            1994

1974       72[degrees]         sight record   Adult         Ferns. 1978

1985       77[degrees]         sight record   Adult         Boertmann.
                                                            1994

1992       66[degrees]         band           Banded 1988   Lyngs, 2003
                               recovery       in UK

1993       77[degrees]         sight record   Adult         Boertmann,
                                                            1994

1998       66[degrees]         band           Banded 1997   Lyngs, 2003
                               recovery       in UK

2001-06    74.5[degrees]       almost annual  Adults        Hansen and
                               observations                 Meltofte,
                                                            2006

2001-05    72.5[degrees]       almost annual  Adults        O. Gilg.
                               observations                 pers. comm.
                                                            2005

2003       74.5[degrees]       confirmed      2 pairs and   M. Elander,
                               breeding (4)   nest with     pers. comm.
                                              eggs          2003

2004       74.5[degrees]       probable       territorial   Gilg, 2005
                               breeding (4)   pair +
                                              single bird

2004       73.5[degrees]       confirmed      nest with     Gilg, 2005
                               breeding       eggs

2005       74.5[degrees]       probable       territorial   Hansen and
                               breeding (4)   pair          Meltofte,
                                                            2006

2005       70.5[degrees]       probable       territorial   R. Burton,
                               breeding       pair          pers. comm.
                                                            2005

2007       81.5[degrees]       sight record   2 adults, 6   C.
                                              July          Nordstr0m,
                                                            pers. comm.
                                                            2007

(1) ZMUC = Zoological Museum of Copenhagen.
(2) Banded on Faroe Islands as chick in 1958.
(3) Island at Eqaluit near Qaqortoq.
(4) Sand Island in Young Sound.


Two extensive recent breeding seabird surveys covering the central and northern part of the west coast of Greenland (69-74[degrees] N in July 1994 and 67.5-70[degrees] N in July 2005) failed to find any lesser black-backed gulls (Boertmann et al., 1996; Boertmann, 2006).

There are far fewer records from the sparsely populated East Greenland (Table 1). Since 2001, however, vagrant lesser black-backed gulls have been observed almost annually at two research sites in High Arctic Northeast Greenland, and in 2007 they were observed as far north as at 81.5[degrees] N (Table 1). The most recent development is that lesser black-backed gulls have been present on a small island at 74.5[degrees] N since the summer of 2003 and (at least in 2003) attempted breeding (Table 1). Breeding and territorial pairs have also been recorded recently at another two small islands in Northeast Greenland (Table 1). No observations from Southeast Greenland have been published, but local hunters recognize the species and observe it regularly along all of the coasts (A. Rosing-Asvid, pers. comm. 2006). Moreover, two of the total three band recoveries of birds banded abroad were recovered in southeast Greenland (Table 1), confirming the occurrence of the species in this part of Greenland. Figure 1 summarizes all the breeding records.

Population Size

The total number of lesser black-backed gull pairs recorded in South Greenland in 2003 is about 600, but the region was not surveyed systematically (i.e., the first known breeding site, where many additional pairs may have occurred in the area, was not visited). Except for a single colony, the numbers in the other core area in the archipelago, around Nuuk, are unknown; but that colony may hold at least 100 pairs. A conservative estimate of the total 2003 population in Southwest Greenland is at least 700 pairs. However, since the coast between the two core areas has not been surveyed recently, and there is no reason to believe that the lesser black-backed gull should be absent there, the breeding population may be considerably larger.

Origin and Subspecies

Band recoveries indicate the origin of the immigrating lesser black-backed gulls. One bird banded as a chick in the Faroe Islands and two birds ringed as immatures in England have been recovered in Greenland one to six years later (Lyngs, 2003). But as Iceland is much closer to Greenland, many immigrants undoubtedly originate there. These birds belong to the pale-mantled northwest European subspecies graelsii. Most sight records also refer to this subspecies. However, two reports, one from West Greenland (Fox and Stroud, 1981) and one from East Greenland (Ferns, 1978) specifically mention birds with darker mantle and upper wings, which suggests that those birds probably belonged to the subspecies intermedius.

Phenology

The earliest date in the year on which lesser black-backed gulls were observed was 12 April, when an adult was observed near Nuuk in West Greenland in 2000 (M. Kviesgaard, pers. comm. 2001), and the latest observation date was 2 November, when two juveniles were seen in 2005, also near Nuuk (C. Egevang, pers. comm. 2006). In 2003, recently fledged juveniles still fed by the parent birds in the breeding colonies were observed in South Greenland on 28 July, and in 2006, a juvenile with still growing primaries and also fed by a parent bird was seen away from the breeding site in Nuuk on 10 September. These dates indicate that egg laying occurs from late May until at least late June (brooding period 25 days and fledging period 35 days; Cramp and Simmons, 1983).

Breeding Habitat

All lesser black-backed gulls breeding in West Greenland were found on small, low islands more or less covered with grass and dwarf scrub vegetation. Here they often nested among other Larus species, usually Iceland gulls (L. glaucoides) or great black-backed gulls (L. marinus). However, the breeding site in Young Sound in Northeast Greenland is a low, almost barren bank of gravel and sand, where the nest was found among large numbers of breeding arctic terns (Sterna paradisaea) and Sabine's gulls (Larus sabini). The two other breeding or supposed breeding sites in Northeast Greenland also hold arctic tern colonies.

DISCUSSION

Although lesser black-backed gulls apparently occurred in waters off South Greenland as early as the 1960s (Brown, 1968), they were considered rare vagrants in Greenland until at least the mid-1970s. Since then, however, their frequency of occurrence has shown a rapid increase. The first breeding record was documented in the mildest part of Greenland, where the inland is ascribed to the Subarctic zone (average July temperature > 10[degrees] C).

Thirteen years later, the species had expanded its range in the Low Arctic part of West Greenland as far north as 66[degrees] N. Farther north in West Greenland, the species still seems to occur only as a scarce summer visitor. In East Greenland, the lesser black-backed gull is now a regular visitor to many areas in both the High and the Low Arctic parts, and breeding occasionally takes place.

[FIGURE 1 OMITTED]

The southern part of Greenland's west coast appears well suited for breeding lesser black-backed gulls: the climate is generally mild and oceanic, and sea ice is absent from most of the region throughout the year. It is more puzzling why the gulls have found their way to the much more inhospitable High Arctic of Northeast Greenland, where heavy drift ice usually blocks the coasts in summer. But the breeding record there coincided with extremely light ice conditions (Rasch and Caning, 2004). In 2006, when the ice conditions were severe, only non-breeding lesser black-backed gulls were recorded at the breeding island in Young Sound (A. T[emptyset]ttrup, pers. comm. 2006).

The increase in lesser black-backed gulls in northwestern Europe during the first half of the 20th century is usually ascribed to increased protective legislation and reduced exploitation, whereas the increase in the latter half of the century may be due to increased food resources (fishery discards, garbage, etc.) and a reduced tendency to migrate (Calladine, 2004). In the early 20th century, the population increase of these gulls also coincided with a general amelioration of the climate. However, the colonization of Greenland took place during a period of regional cooling, when sea temperatures were decreasing around Greenland, which also coincided with the disappearance of the Atlantic cod (Gadus morhua) from the banks off West Greenland (Pedersen and Rice, 2002). This timing indicates that the lesser black-backed gull niche in Southwest Greenland may have been available for a long time, but the colonization did not take place until there was a sufficiently large population surplus from Iceland and northwest Europe to sustain a founder population. Given that the 20th-century range expansion also extended southward to France and the Iberian Peninsula (Cramp and Simmons, 1983), it is likely that the primary cause for the species expansion was not the climate, but an increasing source population in combination with the migratory tendencies of the species.

Two other gull species, the herring gull (Larus argentatus) and the black-headed gull (Larus ridibundus), have followed the same route, using Iceland as a stepping stone when expanding their breeding range across the North Atlantic. These species colonized Iceland in the early 20th century (Cramp and Simmons, 1983) and subsequently found their way to Southwest Greenland, where today they breed irregularly and in very low numbers (Salomonsen, 1979; Boertmann, 1994, 2004). Black-headed gulls also colonized Labrador and Newfoundland (Montevecchi et al., 1987; Chaulk et al., 2004). Lesser black-backed gulls, except for a single odd breeding record from the interior United States, have so far been reported only as vagrants in North America (Post and Lewis, 1995); however, I find it likely that they will follow the black-headed gulls and establish at least temporary small breeding populations along the shores of Labrador or Newfoundland within the next few decades. And if the reduction in sea ice distribution in the Arctic summer continues as predicted by the climate-change models (Walsh, 2005), the potential for a further northward expansion of the breeding range of coastal boreal species (including lesser black-backed gulls) in Greenland and other Arctic areas will increase.

ACKNOWLEDGEMENTS

Thanks go to all reporters of unpublished records mentioned in the text, to Hans Meltofte who commented on the manuscript, to Peter W. Post for sending me photocopies of his papers, and to three anonymous referees who reviewed and improved the manuscript.

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D.BOERTMANN (1)

(1) Department of Arctic Environment, Aarhus University, National Environmental Research Institute, P.O. Box 358, DK-7000, Roskilde, Denmark; dmb@dmu.dk

(Received 27 March 2007; accepted in revised form 4 September 2007)

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