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The Greenland White-fronted Goose Anser albifrons flavirostris

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Mass based on API scores<br />

3600<br />

3200<br />

2800<br />

2400<br />

2000<br />

1600<br />

1200<br />

early Jan<br />

late Jan<br />

early Feb<br />

Wexford<br />

late Feb<br />

early Mar<br />

late Mar<br />

during recesses from the nest by the female, these<br />

bouts are rare and of short duration (see chapter<br />

6). It would therefore seem that, based on observations<br />

of the 'median' female and the calculations<br />

presented here, meeting the energetic and<br />

nutritional costs of laying a clutch and completing<br />

successful incubation is not possible. On this<br />

basis, most females in any given year are unlikely<br />

to attain nutrient and energy thresholds necessary<br />

to reproduce. That said, observations from<br />

several years indicate considerable individual<br />

variation in abdominal profiles between individuals.<br />

Indeed, some birds show considerably faster<br />

rates of accumulation of body mass than do others.<br />

It seems likely, therefore, that only those relatively<br />

few individuals able to accumulate stores<br />

at rates well above the mean throughout the prelude<br />

to breeding will therefore have the potential<br />

to attempt to breed. On this basis, it would appear<br />

that a large proportion of geese could potentially<br />

arrive in west <strong>Greenland</strong> having failed<br />

to reach threshold condition for successful reproduction.<br />

<strong>The</strong> crucial questions, therefore, concern the<br />

mechanisms that are likely to affect the ability of<br />

the individual to acquire the necessary nutrients<br />

early Apr<br />

late Apr<br />

Iceland<br />

early May<br />

<strong>Greenland</strong><br />

late May<br />

clutch of 3<br />

clutch of 6<br />

incubation with 3 eggs<br />

incubation with 6 eggs<br />

Figure 9.2. Estimated fortnightly median body mass<br />

of adult male ( or ) and adult female ( or ) based<br />

on field observations of abdominal profiles and their<br />

observed changes through the first half of the year.<br />

Graph contrasts the slow accumulation of stores at<br />

Wexford (solid symbols, solid line) with the rapid accumulation<br />

in Iceland (open symbols solid line) and<br />

in <strong>Greenland</strong> (solid symbols dotted line). Costs of laying<br />

3 and 6 egg clutches have been subtracted from<br />

the late May median values, and costs of incubation<br />

from these values (based on fat and protein costs from<br />

Table 9.1). Note that this approach underestimates<br />

body mass at all stages because of the effects of using<br />

fortnightly means, hence the differences in some measures<br />

compared to direct mass determinations used<br />

earlier.<br />

early June<br />

late June<br />

for survival and reproduction at each critical<br />

stage. Assuming that the environment is not unlimited<br />

in its ability to supply nutrients, a critical<br />

factor is likely to be the local density of geese.<br />

This factor affects the ability of an individual to<br />

achieve threshold nutrient requirements. What<br />

factors enable some individuals to survive and<br />

breed whilst others cannot? <strong>The</strong>re is abundant<br />

evidence that amongst relatively long lived avian<br />

species such as geese, breeding performance increases<br />

with age (e.g. Owen 1984, Forslund &<br />

Larsson 1992, Cooke et al. 1995). More older birds<br />

attempt to breed than among young age classes<br />

and a greater proportion of older birds breed more<br />

successfully. For example, Raveling (1981) found<br />

although geese 4+ years of age comprised 26% of<br />

the potential breeding population, they produced<br />

more than 50% of young in Giant Canada Geese<br />

Branta canadensis maxima. Specifically, older birds<br />

lay earlier, larger and heavier clutches than young<br />

birds, which ultimately result in more offspring<br />

fledged.<br />

However, reproductive performance in geese generally<br />

increases only for the first 5-6 years of life<br />

(Rockwell et al. 1983, 1993, Forslund & Larsson<br />

1992). This is not consistent with the hypothesis<br />

of reproductive restraint in younger years (Curio<br />

1983), which would predict increasing reproductive<br />

effort throughout life. It would therefore appear<br />

that in many goose populations, young individuals<br />

are constrained from performing well,<br />

perhaps through the lack of social status that permits<br />

access to best feeding opportunities.<br />

Dominance hierarchies have long been recognised<br />

in goose flocks (Boyd 1953, Hanson 1953, Raveling<br />

1970) and rank has been shown to increase with<br />

age (Lamprecht 1986, Black & Owen 1995). However,<br />

amongst birds of the same age class, dominance<br />

explained much of the variation in reproductive<br />

performance, suggesting this was the<br />

overriding factor (Lamprecht 1986, Warren 1994).<br />

Most evidently, dominance determines individual<br />

feeding opportunity through securing and defence<br />

of best feeding opportunities (e.g. Teunissen<br />

et al. 1985, Prop & Loonen 1988, Prop & Deerenberg<br />

1991, Black et al. 1992). This in turn has consequences<br />

for food intake rates, since peck rates<br />

and feeding rates have been found to correlate<br />

positively with dominance (e.g. Warren 1994).<br />

Social status also affects whether a goose pair is<br />

able to obtain and hold a nesting territory. In situations<br />

where predation limits output, hatching<br />

success and fledging rate were also correlated<br />

75

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