Functional
Ecology 2001
15, 24–28
© 2001 British
Ecological Society
24
Blackwell Science, Ltd
Mortality risk of rapid growth in the spider
Nephila clavipes
L. E. HIGGINS*† and M. A. RANKIN‡
*Department of Entomology, University of Massachusetts, Amherst MA 01003, USA, and ‡Department of
Zoology, University of Texas at Austin, Austin, TX 78712, USA
Summary
1. Female Nephila clavipes from univoltine populations have greatly reduced repro-
ductive success if they grow slowly and reach maturity late in the growing season.
Although such fitness costs are expected to select for rapid increases in mass, several
authors have presented models and data describing physiological costs of rapidly
increasing mass.
2. In order to test the hypothesis that there are inherent costs of rapid growth
(increasing mass), laboratory-reared juveniles of the orb-weaving spider Nephila clavipes
were randomly assigned to receive daily feedings ranging from 2·5% to 23% of their
initial mass.
3. Spiders receiving higher amounts of food were more likely to die at or immediately
before the next moulting cycle.
4. These results indicate that there may be inherent physiological costs of rapidly
increasing mass. In opportunistic feeders such as spiders that tend to gorge when
prey are abundant, this could present a real cost to a common foraging strategy.
Key-words : Aranea, fitness costs, growth rates, plasticity, Tetragnathidae
Functional Ecology (2001) 15, 24–28
Introduction
Although variation in growth rate has traditionally
been assumed to reflect non-adaptive phenotypic re-
sponses to the availability of food (e.g. Roff 1980, 1983;
Stearns & Koella 1986), experimental work with some
insects has demonstrated adaptive variation in rate
of growth (reviewed in Gotthard et al. 1994). Low
rate of increasing mass in larval arthropods increases
the duration of each larval stage, and is associated
with both increased risk of prereproductive mortality
(Higgins & Rankin 1996) and reproductive costs of
delayed maturation (Stearns & Koella 1986; Higgins
& Rankin 1996; Higgins 2000). Although very high
rates of growth have been often assumed to increase
fitness, allowing animals to avoid the above-mentioned
costs of slow growth, there is increasing evidence that
there are ecological and physiological costs asso-
ciated with rapidly increasing mass (Stockhoff 1991;
Wiklund et al. 1991; Gotthard et al. 1994; Abrams
et al. 1996; Arendt 1997).
Ecological costs of rapid growth are primarily
assumed to reflect risk of predation and parasitism
associated with the increased foraging necessary to
increase mass more rapidly (Abrams et al. 1996). Such
costs are postulated to underlie observed patterns
of declining investment into foraging when prey
are abundant (Lubin & Henschel 1996). The best
demonstrated physiological cost of rapid growth is
decreased resistance to starvation and other environ-
mental stresses (Stockhoff 1991; Gotthard et al. 1994).
One can postulate other physiological costs of rapidly
increasing mass. For example, rapidly increasing mass
may decrease resource allocation to other functions,
such as development (Arendt 1997). Under these
circumstances, high rates of growth alone could
increase mortality in the absence of external environ-
mental stresses.
In populations of the orb-weaving spider Nephila
clavipes (Linnaeus) (Araneae: Tetragnathidae) inhabit-
ing strongly seasonal environments, there are strong
fitness costs associated with slow growth and develop-
ment in females (Higgins 2000). The duration of each
juvenile instar is negatively correlated with the rate
of increasing mass, and the number of juvenile
instars is variable (Higgins 1992, 1993). Size at matur-
ity is determined in large part in this species by the
number of juvenile instars (Higgins & Rankin 1996),
and rapidly growing females pass through several
instars and reach maturity at a large size earlier
than slowly growing females. There is a strong repro-
ductive advantage to early maturation: large, early
maturing females have disproportionally greater
reproductive success, producing multiple large clutches. †Author to whom correspondence should be addressed.