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The Greenland White-fronted Goose Anser albifrons flavirostris

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during the period when the population was legal<br />

quarry in <strong>Greenland</strong>, Iceland, Ireland and Britain,<br />

although we have little idea of the precise<br />

size and distribution of the harvest at that time.<br />

By way of comparison, the survival rate for the<br />

period 1990-1997 (using only the recovery data<br />

generated from the collar-marking scheme) was<br />

analysed using the Haldane methods and gave<br />

an annual survival rate for the period of 81.7% (±<br />

0.8 SE, H. Boyd, in litt.). <strong>The</strong> 5% difference in survival<br />

rate is very similar to the mean hunting<br />

mortality rate prior to protection (see Figure 8.1).<br />

It was considered that the survival rate of <strong>Greenland</strong><br />

<strong>White</strong>-<strong>fronted</strong> Geese prior to protection was<br />

too low to sustain the then level of hunting kill<br />

(e.g. Owen 1978). For that reason much bureaucratic<br />

and political effort was put into removing<br />

the subspecies from the quarry list, especially on<br />

the wintering grounds. As described earlier, this<br />

led to the effective protection of the population<br />

from hunting on the wintering areas from 1982<br />

onwards (see MS14, Stroud 1992 and Fox et al.<br />

1999 for full details). Evidence from counts of<br />

birds at a number of wintering sites strongly suggested<br />

that the increase in numbers that followed<br />

the cessation of winter hunting was due to the<br />

increase in return rate of birds, not to changes in<br />

reproductive success (MS14). Indeed, we are now<br />

aware that since protection, breeding success has<br />

actually decreased amongst the Wexford and Islay<br />

wintering elements of the population, which<br />

provides more evidence that restriction on hunting<br />

has increased annual survival.<br />

<strong>The</strong>re was no extensive visible marking programme<br />

in effect before and after the implementation<br />

of protection in winter. This would have<br />

allowed a more sensitive monitoring of changes<br />

in survival based on individual bird histories and<br />

thus enable the interpretation of subsequent<br />

changes in overall population size. <strong>The</strong> capturemark-recapture<br />

study using neck-collared birds<br />

initiated in Ireland only commenced after protection<br />

had been implemented. Using the resightings<br />

of neck-collared birds marked at Wexford during<br />

1984-1989, Bell et al. (MS10) calculated annual<br />

adult survival using SURGE4 models (Clobert et<br />

al. 1987, Pradel et al. 1990) to generate maximum<br />

likelihood estimates of 78.5% (± 1.4 SE). This compares<br />

with 72.4% (± 7.3 SE) based on ringing recoveries<br />

from the same ringing programme using<br />

BROWNIE (Brownie et al. 1985). Using the<br />

Haldane method on birds marked in the winter<br />

period gives a survival estimate of 72.1% (± 1.1<br />

Estimated annual adult survival rate<br />

1<br />

0.9<br />

0.8<br />

0.7<br />

0.6<br />

0.5<br />

0.4<br />

0.3<br />

0.2<br />

0.1<br />

0<br />

1982 1984 1986 1988 1990 1992 1994 1996 1998<br />

Figure 8.3. Annual adult survival rate (+ 95% CL) for<br />

<strong>Greenland</strong> <strong>White</strong>-<strong>fronted</strong> Geese caught at Wexford,<br />

1983/84-1997/98 based on observations and recoveries<br />

of neck-collared individuals using the MARK suite<br />

of programs (see text and Appendix 1 for details) 12 .<br />

Open symbols indicate those seasons when the hunting<br />

season was opened at Wexford for a limited shoot.<br />

<strong>The</strong> unusually low survival estimate and large confidence<br />

intervals for 1983 probably reflect small sample<br />

sizes in that year.<br />

SE) for the period 1984-1989 (H. Boyd in litt.) <strong>The</strong><br />

SURGE models use much more fine-grained information<br />

and provide year- and age-specific<br />

maximum likelihood estimates, based on datarich<br />

repeated resighting histories of individual<br />

birds. <strong>The</strong> Brownie techniques utilise the geese<br />

caught, ringed and never retrieved again to estimate<br />

reporting and recovery rates in a more sophisticated<br />

survival estimation than the Haldane<br />

method. It is not possible to use the Brownie et<br />

al. methods on the 1940s data, because some of<br />

the original capture and ringing data do not exist.<br />

More recent analysis has been carried out using a<br />

combination of recoveries and resightings of collared<br />

birds at Wexford using MARK (<strong>White</strong> &<br />

Burnham 1999, based on the recovery-recapture<br />

models of Burnham 1993 and multi-stage models<br />

of Hestbeck et al. 1991). <strong>The</strong> selected model<br />

was one of survival, which varied with year independently<br />

for adults (weighted mean 78.5%,<br />

see Figure 8.3) and juveniles (67.8%, but which<br />

showed greater variability, Figure 8.4), with a<br />

mean of 7% permanent emigration per year (M.<br />

Fredriksen & A.D. Fox unpublished). <strong>The</strong>re was<br />

no effect of hunting on annual adult or juvenile<br />

survival estimates in the two winters (1985/86<br />

and 1989/90) when the moratorium was lifted at<br />

Wexford, although the survival rates in the year<br />

following 1989/90 were the lowest for adults and<br />

juveniles (Figures 8.3 and 8.4). In 1990/91, only 7<br />

out of a marked cohort of 33 first winter juve-<br />

69

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