Journal of Animal Ecology 2008, 77, 1242–1249 doi: 10.1111/j.1365-2656.2008.01442.x
© 2008 The Authors. Journal compilation © 2008 British Ecological Society
Blackwell Publishing Ltd
Lizards combine stored energy and recently acquired
nutrients flexibly to fuel reproduction
Daniel A. Warner
1
*, Xavier Bonnet
2
, Keith A. Hobson
3
and Richard Shine
1
1
School of Biological Sciences, University of Sydney, New South Wales 2006, Australia;
2
Centre d’Etudes Biologiques de
Chizé, CNRS, F-79360 Villiers en Bois, France; and
3
Environment Canada, 11 Innovation Boulevard, Saskatoon,
Saskatchewan, Canada, S7N 3H5
Summary
1. Energy allocation strategies for reproduction are viewed typically as a continuum between
reliance on ‘income’ (recently acquired energy) vs. ‘capital’ (stored reserves) for fuelling reproduction.
Because ectothermy facilitates long-term energy storage and often involves low feeding rates,
traditional views suggest that many ectotherms rely heavily on stored reserves for egg production.
2. We explored the temporal relationship between energy intake and expenditure in a multi-clutching
lizard (Amphibolurus muricatus) by evaluating the effect of maternal nutrition on reproductive
output and by contrasting δ
13
C measurements of the maternal diet and endogenous energy stores
with that of the eggs produced.
3. Our experiment revealed that females utilize both endogenous energy stores and recently
acquired food to fuel reproduction; this pattern did not shift seasonally from first to second clutches
produced. Importantly, however, egg lipid was derived primarily from capital, whereas egg protein
was derived about equally from both income and capital.
4. Overall, these results suggest that the energy allocation strategy used for reproduction differs
among egg components, and that the use of recently acquired energy for reproduction may be more
widespread in ectotherms than thought previously.
Key-words: Amphibolurus muricatus, capital breeding, carbon-13, income breeding, stable isotopes
Journal of Animal Ecology(2007) doi: 10.1111/j.1365-2656.2007.0@@@@.x
Introduction
The temporal relationship between the acquisition of energy
and its expenditure for reproduction is an important axis of
life history variation. Although energy acquisition and
allocation tactics cover a continuum, they can be viewed
usefully in terms of two end-points: reliance on ‘capital’ vs.
‘income’ to fuel reproductive expenditure (Bonnet, Bradshaw
& Shine 1998; Meijer & Drent 1999). ‘Income’ breeders
expend energy for reproduction soon after that energy is
acquired, whereas ‘capital’ breeders gather energy over long
periods prior to utilizing these stored reserves for reproduction
(reviewed by Jönsson 1997). These two strategies of energy
allocation can influence selection on life-history attributes
(e.g. the number and quality of offspring produced, or the
frequency of reproduction: Jönsson 1997; Bonnet et al. 2001;
Brown & Shine 2002; Lourdais et al. 2002), as well as the
sensitivity of reproductive output to local conditions. For
example, local resource availability should influence immediate
reproductive output strongly in income breeders, but not in
capital breeders (due to the temporal separation of energy
acquisition and expenditure in the latter group). Clearly, both
types of energy allocation strategies have associated costs and
benefits; long-term energy storage may allow capital breeders
to cope with unpredictable environments or low food availability
(Calow 1979; Santos & Llorente 2004), whereas income
breeders can increase their reproductive output more rapidly
in response to an increase in local food supply (Jönsson 1997).
Most research on these issues has focused upon avian and
mammalian species (Drent & Daan 1980; Festa-Bianchet,
Gaillard & Jorgenson 1998; Kunkele 2000; Veloso & Bozinovic
2000; Voigt 2003), and detailed studies have revealed that
these high-energy systems rely on income breeding to fuel
some aspect of reproductive expenditure (Bronson &
Manning 1991; Klaassen et al. 2001; Gauthier, Bêty &
Hobson 2003; Wheatley et al. 2008). On the other hand,
features associated with ectothermy may facilitate long-term
energy storage, such that capital breeding may be more
energetically efficient than income breeding in ectotherms
(Bonnet et al. 1998). Consistent with this idea, stored energy
reserves (i.e. fat bodies) are used to fuel reproduction in many
ectothermic animals (Schultz, Clifton & Warner 1991;
*Correspondence author and present address: Daniel A. Warner,
Department of Ecology, Evolution and Organismal Biology, Iowa
State University, Ames, Iowa 50011, USA. E-mail: dwarner@iastate.edu