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The Greenland White-fronted Goose Anser albifrons flavirostris

The Greenland White-fronted Goose Anser albifrons flavirostris

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9 Synthesis<br />

9.1 Anticipatory acquisition of nutrients,<br />

density dependence and constraints<br />

upon fitness<br />

In spring, every female <strong>Greenland</strong> <strong>White</strong>-<strong>fronted</strong><br />

<strong>Goose</strong> of breeding age must clear three nutritional<br />

hurdles in order to breed successfully. <strong>The</strong> annual<br />

cycle involves completion of two spring migration<br />

episodes. Both require the bird to make the<br />

necessary physiological and anatomical modifications<br />

and lay down fuel reserves to sustain two<br />

flights of 1,500 km, one over the sea to Iceland,<br />

and the second crossing the sea and the <strong>Greenland</strong><br />

Ice Cap. <strong>The</strong> ease with which an individual<br />

can clear these hurdles has various fitness consequences.<br />

Failure to construct large enough flight<br />

muscles or energy stores to sustain the flight will<br />

result in death en route – this much natural selection<br />

will ensure. However, the ability to accumulate<br />

the necessary resources to only just complete<br />

the two flights leaves no stores for investment in<br />

reproduction. Slow accumulation of adequate<br />

stores will delay departure from staging areas and<br />

Lipid costs of egg laying and incubation<br />

Costs � body<br />

mass (kg)<br />

BMR<br />

(kJ)<br />

Lipid equivalent<br />

of 1xBMR<br />

(g/day)<br />

DEE<br />

(xBMR)<br />

time of arrival to the nesting grounds, so condition<br />

mediated timing of breeding may also affect<br />

reproductive output in this way. To complete the<br />

journey to the breeding grounds early enough<br />

with some extra stores remaining is likely to contribute<br />

to the investment in reproduction, and<br />

this, together with efficiency in finding food during<br />

the pre-nesting period, is likely to determine,<br />

to a major extent, the reproductive success of that<br />

individual.<br />

On the other hand, there must be some upper<br />

limit on the amount of energy or other nutrient<br />

stores, set by the cost of carrying such excess body<br />

mass (e.g. predation risk and enhanced energy<br />

use induced by heavier flying weight, see review<br />

in Witter & Cuthill 1993). Nevertheless, the ability<br />

to acquire specific nutrients to store for use at<br />

key points in the winter and spring has consequences<br />

for the ability of a bird to reproduce, or,<br />

in the extreme, to survive each migration. <strong>The</strong><br />

efficiency (and therefore the rate) of accumulation<br />

of such 'capital' through a series of acquisi-<br />

Table 9.1. Protein and lipid energy requirements for a laying <strong>Greenland</strong> <strong>White</strong>-<strong>fronted</strong> <strong>Goose</strong>, her clutch of<br />

3 or 6 eggs and subsequent incubation. Analysis follows methods of Meijer & Drent (1999), using Basal<br />

Metabolic Rate estimated from the relationships for non-passerines derived by Aschoff & Pohl (1970), i.e.<br />

BMR = 330.W 0.722 (where W = body weight in kg.) and based on the assumption that daily energy expenditure<br />

(DEE) during laying is equivalent to 1.7 x BMR. Protein costs were calculated using the modified<br />

formula of Robbins (1981) related to body weight according to the formula 2.68.W 0.75 g protein day -1 (see<br />

Meijer & Drent 1999 for full explanation).<br />

Laying/<br />

incubation<br />

period<br />

(days)<br />

Lipid used in<br />

laying/<br />

incubation<br />

period<br />

(g)<br />

Lipid<br />

used in<br />

clutch<br />

(g)<br />

Total<br />

minimum<br />

and<br />

minimum<br />

lipid<br />

investment<br />

(g)<br />

Laying 2.78 690 18.2 1.7 4-8 123.5-247.1 49-98 172.5-345.1<br />

Incubation 2.78 690 18.2 1.1 26 519.6 519,6<br />

Protein costs of egg-laying and incubation<br />

Costs � body Maintenance Laying/ Protein used in Protein used in Total<br />

mass (kg) protein incubation period laying/incubation clutch minimum<br />

(g/day)<br />

(days)<br />

period (g)<br />

(g)<br />

and<br />

minimum<br />

protein<br />

investment<br />

(g)<br />

Laying 2.78 5.7 4-8 23.1-46.2 56-112 79.1-158.2<br />

Incubation 2.78 5.7 26 165.0 165.0<br />

73

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