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Effects of Great Skuas on Arctic Skuas in Shetland Robert Furness

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Effects of Great Skuas on Arctic Skuas in Shetland Robert Furness Effects of Great Skuas on Arctic Skuas in Shetland Robert Furness Great Skuas, which have increased in numbers in Shetland, kill both adult and young Arctic Skuas. How has this predation affected Shetland's Arctic Skua colonies, and how will they fare in the future? he Great Skua or Bonxie Stercorarius skua was brought close to extinction Tin Britain during the i gth century through extensive human persecution, but since 1900 the numbers breeding in Shetland have increased rapidly (Venables and Venables 1955, Parslow 1973, Cramp et al. 1974). Most colonies there are still growing, although in some cases less rapidly than in earlier years (personal observation). Many authors (e.g. Pennie 1948, Baxter and Rintoul 1953, Venables and Venables 1955) have claimed that this increase has resulted in a decrease in Arctic Skuas S. parasiticus, but this view has been supported only by circumstantial evidence. Bonxies kill both adult and young Arctic Skuas, and may rob adults of food. They return to the breeding colonies earlier in the year than the smaller species and may establish territories in areas formerly held by Arctic Skuas, which are then forced to move elsewhere (Jackson 1966). These activities seem likely to have an adverse effect on Arctic Skua popu­ lations, but have not previously been quantitatively studied. Nor has any continued.... 96 [Brit, Birds, 70: 96-107, March 1977] Great and Arctic Skuas in Shetland 97 attempt been made to collect and interpret the estimates of Shetland Arctic Skua colony sizes recorded by numerous ornithologists since 1890. The 1969 'Seafarer' counts indicated that Shetland held 96% and 71% of the British populations of Bonxies and Arctic Skuas respectively (Bourne and Dixon 1974). Most of the Bonxies breed in three large colonies, on Foula, Unst and Noss. Large numbers of Arctic Skuas also nest on these islands, so it is in these areas that interspecific interactions would be expected. Fortunately, these three areas are extensively documented. This paper examines published and previously unpublished census data for these three colonies in relation to quantitative observations of the inter­ specific interactions. Census data Counting pairs is a relatively simple matter. Arctic Skuas maintain ex­ clusive territories of 20-100 m diameter during the breeding season. They are large, conspicuous birds, aggressive towards human intruders. Feeding occupies only a few hours each day, allowing both of the adults to be in the territory most of the time, with at least one present all the time from laying to shortly before the young are fledged. Several counting techniques have been used. Some estimates are based on single counts of apparently occu­ pied territories, others on mapping of pairs in relation to topographical features, and a few on the number of nests with eggs found and marked during a breeding season. All three methods can underestimate or over­ estimate the colony size. It is unlikely that all nests in a colony will be found, but pairs which lose their eggs usually lay a replacement clutch, often some distance from the first nest, and it is difficult to distinguish these from late first clutches. In practice, these two errors tend to cancel each other out. Counts of occupied territories include non-breeding territory holders, but probably exclude a few pairs which are particularly timid. These errors also tend to cancel each other, so both methods should give close estimates of the number of breeding pairs. Estimates based on a single count may be less accurate, and probably tend to under-record the true number, but most of the counts used in this paper were made by several observers over periods of weeks. In 1965, 1974 and 1975, counts were made independently in the same colony. These gave similar results (table 1), indicating that all three methods are suitable for censuses of Arctic Skua colonies. Table 1. Estimates of pairs of Arctic Skuas Stercorarius parasiticus on Foula, Shetland, by different census methods Difference between Estimate methods Year Source Method (pairs) (%) 1965 Fabritius (1969) mapping pairs 140 15 Brathay count of pairs iaa .} 1974 This study mapping pairs 180 J. W. F. Davis count of pairs 200 "} '975 This study mapping pairs 240 J. W. F. Davis marking nests 253 } 98 Great and Arctic Skuas in Shetland Census results In other parts of Scotland, where there are few Bonxies, Arctic Skua numbers have probably changed little, but with suggestions of decreases in northern Scotland and increases in Orkney (Cramp et al. 1974); census data for Shetland should be examined against this background. Table 2. Estimates of pairs of Arctic Skuas Stercorarius parasiticus on Noss, Shetland Estimate Year Source Method (pairs) 1922 Baxter and Rintoul (1953) count of pairs 45 1929 Perry (1948) count of pairs 60 1934 Perry (1948) count of pairs 50 1939 Perry (1948) count of pairs 25 1946 Perry (1948) marking nests 3i 1947 Baxter and Rintoul (1953) count of pairs 37 '955 Kinnear (1974) count of pairs 25 1957 Gordon (1964a) count of pairs 13 1958 Kinnear (1974) count of pairs 25 1964 Gordon (1964a) count of pairs 17 1969 Kinnear (1974) count of pairs 40 i97o Kinnear (1974) count of pairs 40 1973 Kinnear (1974) mapping pairs 39 '974 Kinnear (1974) marking nests 44 Numbers on Noss have fluctuated considerably, from 13 to 60 pairs, but show no long-term trend, the number of pairs now being the same as that recorded in 1922 (table 2). Data for Unst are less reliable: there has been a considerable decrease at Hermaness, but an increase in other parts of Unst; the total number of pairs over the whole of Unst appears not to have decreased by more than 10% since 1922 (table 3). These two Arctic Skua colonies have maintained themselves despite increases in the Bonxie populations, from a score of pairs on Unst and none on Noss in 1900 to the current totals of 1,000 pairs on Unst and 250 pairs on Noss. Although numbers of Arctic Skuas appear to have changed little, the increase in the numbers of Bonxies at Hermaness has caused a major redistribution of the Arctic Skuas over the rest of Unst. The most thoroughly documented of the three colonies is that on Foula, where the Arctic Skuas have increased slowly and with some fluctuations, from 60 pairs in 1890 to 130 pairs in Table 3. Estimates of pairs of Arctic Skuas Stercorarius parasiticus on Unst, Shetland Density is measured in pairs per square kilometre HERMANESS REST OF UNST Year Source Method Pairs Density Pairs Density 1922 Pitt (1922) mapping pairs 200 29 o o 1937 Baxter and Rintoul (1953) count of pairs 100 1950 Gordon (1964b) marking nests 75 ■ij — - 1958 Gordon (1964b) count of pairs 70 1905 Dott (1967) mapping pairs 60 1969 Bourne and Dixon (1974) count of pairs 80 1974 Bundy (1974) mapping pairs 72 72 121 24 Great and Arctic Skuas in Shetland 99 Table 4. Estimates of pairs of Arctic Skuas Stenorarius parasiticus on Foula, Shetland Colony area Density Year Source Method Pairs (km2) (per km2) 1890 Barrington (1890) count of pairs 60 — .— 1948 Pennie (1948) count of pairs 100 3-o 33 i960 Brathay (unpubl.) mapping pairs I31 2-3 57 1961 Brathay (unpubl.) count of pairs 130 — — 1962 Brathay (unpubl.) count of pairs 120 — — 1963 Brathay (unpubl.) count of pairs 120 — — 1964 Brathay (unpubl.) count of pairs 120 — — 1965 Fabritius (1969) mapping pairs 140 ■— — •965 Brathay (unpubl.) count of pairs 120 — — 1966 Brathay (unpubl.) count of pairs 120 — — 1968 Brathay (unpubl.) mapping pairs 160 2-3 70 1969 Brathay (unpubl.) count of pairs 100 — — 1972 Brathay (unpubl.) mapping pairs 150 2.0 75 1973 This study mapping pairs 130 2.0 65 1974 This study mapping pairs 180 1.8 100 1974 J. W. F. Davis {in litt.) count of pairs 200 — — 1975 This study mapping pairs 240 1.8 133 1975 J. W. F. Davis {in litt.) marking nests 253 — — 1973, then suddenly to 190 pairs in 1974 and 250 pairs in 1975 (table 4). This increase is all the more surprising when it is remembered that the Foula Bonxie colony is the biggest and densest in Britain, having grown continuously at a rate of 6.8% per annum, from only 29 pairs in 1906 to 2,500 pairs in 1973 (Furness 1974a) and 3,000 pairs in 1976. Clearly, any deleterious effects of Bonxies in Shetland have been insufficient to cause a decline in Arctic Skua numbers, which have even increased. Factors determining colony size On Fair Isle, adult Arctic Skuas have an annual survival rate of 0.80 (Davis and O'Donald 1973). Most breed for the first time when four years old (Berry and Davis 1970). Fledgling production varies between colonies, and from year to year, but is usually between 1.0 and 1.3 chicks per pair, so that a first-year mortality of about 30% would result in a stable population. Variations in any of these statistics could tend to cause fluctuations in breeding numbers, but the deferred maturity, and variations in age of first breeding between individuals, would smooth out such fluctuations; it follows that rapid changes in numbers of breeding pairs are likely to be due to high rates of immigration or emigration. Many seabirds show a strong tendency to return to their natal area to breed, but this habit may not be well developed in Arctic Skuas, since many individuals ringed as chicks have been retrapped breeding in different colonies (Broad 1973, 1974, British Trust for Ornithology ringing returns) and in arctic areas this species moves its breeding locality from year to year depending upon the densities of small mammals arid; Pomarine Skuas S.
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