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The Greenland White-fronted Goose Anser albifrons flavirostris

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7 Moult of flight feathers<br />

7.1 Introduction<br />

Considering the annual cycle of birds in the context<br />

of periods of nutritional stress, the period of<br />

replacement of the body plumage represents one<br />

such stage. <strong>The</strong> maintenance of plumage has considerable<br />

consequences for the individual, not just<br />

in terms of flight and aerodynamic efficiency, but<br />

also because feathers provide effective thermal<br />

insulation and, in the case of waterbirds, buoyancy.<br />

<strong>The</strong> Anatidae have evolved a pattern of replacement<br />

of feathers throughout the annual cycle,<br />

and indeed, ducks may have multiple<br />

plumages and moult throughout the year. In<br />

northern geese, replacement of feathers occurs<br />

throughout the annual cycle, except during the<br />

breeding period (e.g. Gates et al. 1993), although<br />

it is the period of flightlessness during the postbreeding<br />

phase which has attracted most research<br />

attention (Hohman et al. 1992).<br />

In common with most of the Anatidae, and all<br />

northern-nesting geese, <strong>Greenland</strong> <strong>White</strong>-<strong>fronted</strong><br />

Geese shed their flight feathers simultaneously<br />

and become flightless for a period of 3-4 weeks<br />

(Belman 1981). Flightlessness constrains feeding<br />

opportunities and denies flying as a means of<br />

predator escape, hence it seems highly likely that<br />

selection will have reduced the flightless period<br />

as much as food resources and predation risk<br />

would permit. Whether the flightless period represents<br />

a period of stress (i.e. where the nutrient<br />

demands of a bird exceed the supply derived from<br />

ingestion, resulting in catabolism of body tissue<br />

to meet that demand, sensu Ankney 1979) has<br />

been the subject of considerable debate (see<br />

Hohman et al. 1992). Hanson (1962) demonstrated<br />

that captive Canada Geese, fed ad libidum throughout<br />

wing moult, lost weight. Ankney (1979)<br />

pointed out that goslings increase their body<br />

weight 20 fold, grow leg and breast muscles and<br />

ossify a skeleton to almost adult size and grow a<br />

full set of body tail and flight feathers. Hence, he<br />

considered that it would be surprising if adult<br />

Lesser Snow Geese could not meet the nutrient<br />

demands of wing moult over the same time period.<br />

Nevertheless, Owen & Ogilvie (1979)<br />

showed significant declines in body mass with<br />

moult stage in Barnacle Geese in Svalbard<br />

amongst adult males and yearling females. Gates<br />

et al. (1993) demonstrated significant declines in<br />

lipid reserves during wing moult in breeding females<br />

and non-breeding Canada Geese, suggesting<br />

that energy stores (i.e. nutrients accumulated<br />

in advance of periods of demand, sensu van der<br />

Meer & Piersma 1994) may be built up in advance<br />

of wing moult. Non-breeding Greylag Geese<br />

<strong>Anser</strong> anser moulting on the Danish island of<br />

Saltholm selected the most protein rich food available<br />

(Fox et al. 1998) and showed modifications<br />

to their nitrogen metabolism (Fox & Kahlert 1999),<br />

yet still lost 12-26% of their body weight (MS17).<br />

Subsequent dissection of birds obtained at this<br />

site throughout wing moult has shown that most<br />

of this involves use of extensive abdominal, mesenteric<br />

and sub-cutaneous fat deposits which are<br />

completely depleted by the time flight feathers<br />

are regrown (unpublished data). <strong>The</strong> study of van<br />

Eerden et al. (1998) also demonstrated that Greylag<br />

Geese moulting in the Netherlands could not<br />

meet their daily energetic requirements during<br />

moult. In combination with use of field scores of<br />

abdominal profiles to assess declines in fat stores<br />

(Owen 1981, Loonen et al. 1991), these authors<br />

concluded that geese relied upon fat deposits to<br />

meet their energy requirements during moult.<br />

Analysis of stable isotope ratios in the new grown<br />

feathers of the geese from Saltholm also suggest<br />

strongly that some of the protein involved in<br />

growth of feather tissue must originate from body<br />

stores accumulated prior to the moult period (unpublished<br />

data). Greylag Geese moulting in Iceland<br />

do not show significant changes in body<br />

mass during moult (A. Sigfusson & C. Mitchell<br />

in litt.), hence, it seems that this species shows a<br />

flexible response to nutrient acquisition and wing<br />

moult, depending on local conditions.<br />

Is wing moult a period of energetic stress (sensu<br />

Ankney 1979) for the <strong>Greenland</strong> <strong>White</strong>-<strong>fronted</strong><br />

<strong>Goose</strong>? What do the geese do during moult? What<br />

habitats are utilised? Are there any indications<br />

that the nutrient requirements of the period and<br />

finite habitat availability during this stage of the<br />

annual cycle could create density dependent limitations<br />

on the population in the future?<br />

7.2Moulting distribution and habitat<br />

<strong>The</strong> non-breeding element of the <strong>Greenland</strong><br />

<strong>White</strong>-<strong>fronted</strong> <strong>Goose</strong> population moults in close<br />

59

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