We reviewed information on the demands of incubation to examine
whether these could influence the
optimal clutch size of birds. The results indicate that appreciable
metabolic costs of incubation commonly
exist, and that the incubation of enlarged clutches can impose penalties
on birds. In 23 studies on 19 species,
incubation metabolic rate (IMR) was not elevated above the metabolic
rate of resting non-incubating birds
(RMR), but contrary to the physiological predictions of King and
others, IMR was greater than RMR in
15 studies on 15 species. Across species, IMR was substantially
above basal metabolic rate (BMR), averaging
1.606×BMR. Of six studies on three species performed under
thermo-neutral conditions, none found IMR
to be in excess of RMR. IMRs measured exclusively within the
thermo-neutral zone averaged only
1.08×BMR contrasting with the significantly higher figure
of 1.72×BMR under wider conditions. 16 of 17
studies on procellariiforms found IMR below RMR, indicating a
significant difference between this and
other orders. We could find no other taxonomic, or ecological
factors which had clear effects on IMR. Where
clutch size was adjusted experimentally during incubation,
larger clutches were associated with: significantly
lower percentage hatching success in 11 of 19 studies; longer
incubation periods in eight of ten studies;
greater loss of adult body condition in two of five studies;
and higher adult energy expenditure in eight of
nine studies. Given that incubation does involve metabolic
costs and given that the demands of incubation
increase sufficiently with clutch size to affect breeding performance,
we propose that the optimal clutch size
of birds may in part by shaped by the number of eggs the parents can afford
to incubate.