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The Greenland White-fronted Goose Anser albifrons flavirostris

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8 Survival<br />

8.1 Introduction<br />

Almost all goose populations have increased in<br />

the Western Palearctic in the last 45 years (Madsen<br />

et al. 1999). Since many of these are discrete and<br />

closed populations, the increases cannot be accounted<br />

for by immigration of individuals from<br />

elsewhere. <strong>The</strong>se increases have not always been<br />

brought about by enhanced reproductive output,<br />

indeed for many populations, recruitment has<br />

actually fallen with increasing abundance (e.g. the<br />

Russian-breeding Barnacle <strong>Goose</strong> population<br />

Ebbinge 1991, and see also chapter 6). Population<br />

modelling and simulation demonstrates that<br />

as relatively long-lived birds, northern nesting<br />

geese are sensitive to small changes in annual<br />

adult survival - very much larger changes in productivity<br />

rates are necessary to effect similar<br />

changes in the rate of population change (e.g.<br />

Tombre et al. 1997, Pettifor et al. 1999). <strong>The</strong> size<br />

of a population is determined by the relative annual<br />

gain (birth rate and immigration) balanced<br />

against loss (death and emigration), hence it is<br />

natural to accredit recent increases in abundance<br />

to declining mortality rates. As a very high proportion<br />

of individually marked geese has been<br />

recovered as a result of hunting, it is assumed that<br />

hunting is responsible for a high proportion of<br />

deaths (e.g. Ebbinge 1991). Restrictions and complete<br />

banning of hunting on particular goose populations<br />

have resulted in immediate increases in<br />

some species including the <strong>Greenland</strong> <strong>White</strong><strong>fronted</strong><br />

<strong>Goose</strong> (MS14). Others have shown less<br />

immediate responses after protective legislation<br />

(e.g. Russian Barnacle Geese Ebbinge 1991 and<br />

Svalbard Barnacle Geese Owen & Black 1999).<br />

Others still have shown long-term increases despite<br />

apparent increasing mortality (e.g. Russian<br />

<strong>White</strong>-<strong>fronted</strong> Geese <strong>Anser</strong> a. <strong>albifrons</strong>, Mooij et<br />

al. 1999) or sudden rapid increases not linked in<br />

any way to changes in hunting restriction (e.g.<br />

the Iceland/<strong>Greenland</strong> Pink-footed <strong>Goose</strong>, Mitchell<br />

et al. 1999). Hence, while it has been suggested<br />

that increases in numbers of some goose populations<br />

are primarily due to the decreased mortality<br />

rates resulting from reduction in shooting, this<br />

is unlikely to be the sole factor influencing<br />

changes in population size. However, there is<br />

evidence that adult annual survival rates have<br />

been greater under protection than in earlier pe-<br />

riods when the geese were subject to hunting exploitation<br />

(Ebbinge 1991, MS14).<br />

This relationship is important in order to understand<br />

the nature of hunting mortality if restriction<br />

on shooting kill is to be used justifiably as a<br />

management tool to achieve nature conservation<br />

management goals. It is necessary to understand<br />

the extent to which the number of deaths caused<br />

directly through hunting add to natural mortality<br />

(additive mortality), rather than being compensated<br />

for through a consequent reduction in<br />

natural loss through some density-dependent<br />

function (compensatory mortality, see discussion<br />

in Anderson & Burnham 1976, Nichols et al. 1984,<br />

Newton 1998). In order that hunting loss is directly<br />

compensated for by reductions in natural<br />

mortality, natural loss must already be density<br />

dependent and the kill cannot take place after the<br />

main period of natural loss. Hence the impact of<br />

the hunting bag in terms of the mortality in addition<br />

to natural loss depends on both the numbers<br />

killed in the hunt, the seasonal timing of both<br />

losses and the degree of density dependence involved<br />

in natural mortality. However, to demonstrate<br />

such a mechanism is important – if hunting<br />

mortality were completely compensatory,<br />

protection of a hunted population would not result<br />

in an increase of numbers. Conversely, demonstrating<br />

that mortality is completely additive<br />

enables restriction of hunting kill to be used as a<br />

tool to directly influence population size, controlling<br />

for reproduction rates. Hunting mortality<br />

may ultimately become partially additive to natural<br />

losses, i.e. at low levels, this would have no<br />

effect on the total annual death loss but above a<br />

certain density threshold, extra hunting mortality<br />

does contribute to a reduction in survival.<br />

Serious declines in the numbers and range of the<br />

<strong>Greenland</strong> <strong>White</strong>-<strong>fronted</strong> <strong>Goose</strong> were attributed<br />

to habitat loss and the effects of hunting in the<br />

1980s (Ruttledge & Ogilvie 1979). This led to the<br />

protection of the population at most haunts on<br />

the wintering grounds and ultimately to the drafting<br />

of a management plan for the population<br />

(Stroud 1992). <strong>The</strong> geese had been legal quarry<br />

throughout its range (West <strong>Greenland</strong> breeding<br />

grounds, Iceland staging areas and wintering<br />

quarters in Britain and Ireland) until the early<br />

65

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