The Greenland White-fronted Goose Anser albifrons flavirostris
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