Functional
Ecology 2006
20, 1037–1044
1037
© 2006 The Authors.
Journal compilation
© 2006 British
Ecological Society
Blackwell Publishing Ltd
Ant stoichiometry: elemental homeostasis in
stage-structured colonies
A. D. KAY,*†‡ S. ROSTAMPOUR* and R. W. STERNER*
*Department of Ecology, Evolution and Behavior, University of Minnesota, St Paul, MN, USA
Summary
1. Organisms facing variation in food quality maintain elemental composition within
limited bounds. Such stoichiometric homeostasis has often been considered a species-
specific parameter, but stoichiometry can also vary intraspecifically across life stages,
sexes and sizes. In colonial organisms with overlapping generations, stoichiometric vari-
ation among stages could lead to flexibility in colony-level elemental composition due
to changes in internal demography.
2. We examine how the balance of energy (sucrose) and nutrients (prey) affects growth
rate and carbon : nitrogen : phosphorus (C : N : P) homeostasis in a eusocial insect,
the pavement ant Tetramorium caespitum.
3. Colony growth depended heavily on prey availability. However, sucrose scarcity led
to higher worker mortality and production of smaller workers, suggesting sucrose
availability will affect colony-level performance in a competitive environment.
4. In contrast, C : N : P stoichiometry of larvae, pupae, and workers varied mostly
with sucrose availability. Biomass P content within life stages was lower in colonies
receiving less access to sucrose. We suggest this difference arose primarily from shifts
in individual ant mass coupled with negative P-body mass relationships.
5. Life stages differed considerably in elemental composition, and resource conditions
affected colony stage structure. Nevertheless, variation in colony-level stoichiometry
primarily reflected compositional differences within stages rather than shifts in internal
demography.
Key-words: carbohydrates, growth rate hypothesis, nutrient balance, phosphorus, RNA
Functional Ecology (2006) 20, 1037–1044
doi: 10.1111/j.1365-2435.2006.01187.x
Introduction
Homeostasis has served as a conceptual framework for
animal physiologists for over a century (Bernard 1872;
Randall, Burggren & French 2001). More recently, it
has emerged as a central idea in ecological stoichiom-
etry, the study of the balance of energy and materials
in living systems (Sterner & Elser 2002). The degree to
which organisms regulate elemental composition may
influence the magnitude and nature of their ecological
impact. For example, strict elemental homeostasis
may result in consumers disproportionately depleting
resources containing scarce materials and increasing
recycling rates of materials ingested in excess of demand
(Vanni 2002). Alternatively, elemental composition
may be flexible, and stoichiometric shifts in individuals
could reflect changes in functional capabilities (Kay
et al . 2005). For example, the growth rate hypothesis
(GRH) predicts a causal link between variation in
whole-body P concentration and growth rate, a key
life-history attribute, due to the role that P-rich ribo-
somal RNA plays in regulating protein synthesis (Elser
et al . 1996). Elucidating the degree of homeostasis in
different taxa and the factors that explain regulatory
variation should help clarify the mechanisms that link
food quality to consumer traits that affect ecological
interactions (Sterner & Elser 2002).
In this paper, we examine elemental homeostasis at
the individual- and colony-level in the pavement ant,
Tetramorium caespitum . No previous study has exam-
ined the C : N : P stoichiometry of a social insect, but
this group may prove to be an important system for
exploring the maintenance and ecological relevance of
elemental homeostasis. Relative to solitary animals,
ants and other social insects may be more likely to
vary in chemical composition in response to changing
food quality due to within-colony specializations.
Social insect colonies contain multiple life stages with
†Author to whom correspondence should be addressed.
E-mail: adkay@stthomas.edu
‡Present address: Department of Biology, University of St
Thomas, St Paul, MN, USA, 55105