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Why Should Greylag Goose Anser Anser Parents Rear Offspring of Others?

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Why Should Greylag Goose Anser Anser Parents Rear Offspring of Others? 25 Why should Greylag Goose Anser anser parents rear offspring of others? L. Nilsson & H. Kampe-Persson Department of Animal Ecology, Lund University, Ecology Building, SE-223 62 Lund, Sweden. leif. nilsson@zooekol. lu. se, hakonpersson@hotmail. com The benefits and costs of rearing large broods were studied in a popula­ tion of neck collared Greylag Geese in Scania, southernmost Sweden. Families were divided into four groups according to brood size: 1-4, 5-8, 9-12 and >12 young. Gosling survival showed a significant positive rela­ tionship with brood size on arrival at the rearing area. First-year and third-year local survival, as well as breeding recruitment, showed a sig­ nificant positive relationships with brood size at fledging. For adults rearing large broods, no costs were detected during the year following that when they fledged >8 young; both survival rate and reproductive success exceeded the long-term averages. Key Words: neck collared population, adoption, survival, recruitment, south Sweden Greylag Geese Anser anser are 1983). In addition to these very large sometimes seen with very large broods broods, there are also many cases of small young, some broods being so where just one or two eggs are added to large (up to 55 young) that it is impossi­ a clutch, or one or a few young are ble for a single female to have adopted into a brood. incubated so many eggs (Karlsson et al. The habit of intra- and inter-specific 1982; Jensen 2000; Persson 2002). It is egg parasitism and the adoption of well-known that Greylag Geese, like a small young is widespread among dif­ number of other waterfowl species, ferent species of waterfowl (Eadie et ai sometimes lay their eggs in the nest of 1988; Lank et ai 1989; Weigmann & another female, but excessively large Lamprecht 1991; Williams 1994; clutches laid by two or more females Beauchamp 1997; Andersson & Åhlund rarely hatch (Hauff 1982; Witkowski 2001). The adoption of foreign young is ©Wildfowl & Wetlands Trust Wildfowl [2003) 54: 25-37 26 Why should Greylag parents rear offspring of others? a prevalent habit of the Common Eider (Gregoire & Ankney 1990). Somateria mollissima and has been In this study, these questions are studied by a number of workers, who addressed for the Greylag Goose by have discussed the advantages for studying pre- and post-fledging sur­ females that Leave their young in the vival rates and recruitment rate in a care of others (eg Bustnes & Erikstad neck collared population that had been 1991; Pöysä 1995; Kilpi et al. 2001; studied intensively since 1985. Bustnes et al. 2002). Eadie & Lumsden (1985; cf. also Arnat 1987) concluded Study population that nest parasitism would sometimes The study population has never not be disadvantageous for the host but been manipulated during nesting, might be of advantage. because of the risk of influencing the Modern DNA techniques have breeding performance (cf. Witkowski revealed that there is a high frequency 1983). of mixed parentage due to nest para­ Pre-hatch brood amalgamation has sitism and adoption of young both in not been studied in this population. For ducks (Andersson & Åhlund 2001) and the present, there is only circumstan­ geese [Choudhury et al. 1993; Larsson tial evidence of pre-hatch brood et al. 1995). Among geese, parental amalgamation, such as the occurrence care in the form of vigilance and pro­ of nests containing >12 eggs and the tection can serve all members of a nest of a Barnacle Goose Branta leu­ brood, and the acceptance of extra copsis which contained four of its own young would therefore have no negative eggs and six Greylag Goose eggs effects on the host family (Lazarus & (Nilsson & Persson 1994; Persson Inglis 1986). Cooch (1991) found a 1997). A clutch of 12 eggs is widely faster growth rate in large families and accepted as the largest a Greylag suggested that the adoption of young Goose could lay naturally (Kampe- could lead to positive effects for the Persson 2002). In a German study, 7% host family. Larger families have a bet­ of 467 nests contained >12 eggs, and ter chance of detecting predators at an 18% of all eggs were found in nests early stage, and the strategy of adopt­ containing 13-40 eggs (Hauff 1982). In a ing foreign goslings may dilute the Polish study, only two clutches out of predation on their own offspring (Eadie 629 contained >12 eggs; all clutches & Lumsden 1985; Eadie et al. 1988). containing 1-6 eggs were laid by a sin­ Larger families are socially dominant gle female, while more than half of over smaller families (Black & Owen those containing 9-12 eggs were laid by 1989; Boyd 1953; Hanson 1953; two females [Witkowski 1983). Overall, Raveling 1970), which was found to be 4% of all nests contained eggs laid by of advantage in wintering Lesser Snow two females; in 30% of these, two Geese Anser caerulescens caerulescens females had laid two complete clutches Why should Greylag parents rear offspring of others? 27 in a single nest, while in the remaining area, and early-formed broods always 70%, one female had laid a complete consisted of similar-aged goslings clutch and another female had added (Persson 2002). In the rearing areas, on 1-3 eggs. The Swedish study popula­ the other hand, permanent brood mix­ tion, established in the late 1960s, grew ing was an extremely rare event, while on average 15. 3% per annum (Nilsson pairs frequently lost or gained one or a et al. 2002), and now numbers >1, 000 few goslings. Warhurst & Bookhout breeding pairs. As a consequence, (1983) suggested that post-hatch brood pairs nest close together, either in amalgamation in Canada Geese Branta reedbeds or on small islands (Nilsson canadensis occurs due to accidental & Persson 1994), which increases brood mixing shortly after hatching in opportunities to lay eggs in other areas of high brood density. The same females' nests (Brown 1984). seems to apply to the study population. Consequently, there is no reason to During the study period, the most assume pre-hatch brood amalgama­ common brood size of families arriving tion to be less frequent in the study at the rearing area was five, but four population than in other goose species was almost as common (Figure 1). The (Lank et al. 1989; Weigmann & commonest brood size at fledging was Lamprecht 1991). three, closely followed by two and four Almost all families with >12 young young. The largest brood seen during formed before they reached the rearing the study had 27 young, followed by Figure 1. Distribution of brood sizes in the study area, Scania, south Sweden, on arrival to the brood- rearing area (light columns) and at fledging (dark columns), pooled over 1984-2002. All families having at least one neck collared parent were included. 28 Why should Greylag parents rear offspring of others? families of 22, 21 and 17 young. The area on the coast, where a large pro­ largest family at fledging had 22 young. portion of the Greylag Geese from the A total of nine broods numbering >12 study area gather (cf. Nilsson & young when first seen at the rearing Persson 1992). area was found, six at Lake Yddingen For analysis, families were divided and three at Lake Börringesjön. The into four groups according to brood mean number of young in these broods size, on arrival at the rearing area and decreased during brood-rearing, from at fledging, respectively: 1-4, 5-8, 9-12 17. 4±1. 63 to 14. 6+1. 4-9. and >12 young. Rearing survival was expressed as Method the number of geese that fledged as a percentage of the number that had Greylag Goose families were cap­ reached the brood-rearing area. tured and neck collared annually in a Calculations were based on the num­ study area in southwestern Scania, ber of young in families having at least southernmost Sweden, from 1985 one neck collared parent. All such fam­ onwards (Nilsson & Persson 1992; ilies recorded in a rearing area within Persson 1994). A total of 575 adults and ten days of hatching were included. 1, 700 goslings had been neck collared Account was taken of pairs that lost all by 2000 (Persson 2000a). Goslings were their young and left the study area to neck collared on average 22±0 days moult elsewhere, as well as of pairs before they fledged (Nilsson et al. 1997). that left the main rearing area before Individuals marked with poor quality fledging. Permanent brood mixing was neck collars (mainly 1984-1985) were accounted for by assuming that gains excluded from the analyses (cf. Persson and losses of goslings were indepen­ dent of family size. In this way, survival 2000b). During 1985-2002, regular searches estimates were not influenced as the for neck collared geese were undertaken individual was used as a sampling unit from their arrival in spring until their when following changes in brood size. autumn departure. In spring, the study During the latter half of the rearing area was visited several times a week period, goslings commonly left their in order to establish the return of parents without joining another family for shorter or longer periods (up to marked birds and their breeding per­ formance. From late May until the last three weeks), but all recorded separa­ young were fledged, the breeding area tions ended with a complete was visited almost daily to establish the re-grouping of the family. As signifi­ number of fledglings in each family. In cantly fewer small young survived to summer and autumn, weekly searches fledging at Lake Fjöllfotasjön than at were extended to include neighbouring the three lakes, Klosterviken, staging areas, especially the Foteviken Börringesjön and Yddingen (45% vs 68- Why should Greylag parents rear offspring of others? 29 71%; Nilsson et at.
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