Changes in function and temporal variation in a guild of
gall-parasitoids across a temperature gradient in Australian
subtropical rainforest
CASEY R. HALL,
1,
* CHRIS J. BURWELL
1,2
AND ROGER L. KITCHING
1
1
Environmental Futures Research Institute (EFRI), Griffith School of Environment, Griffith University,
170 Kessels Road, 4111 (Email: casey.hall@griffithuni.edu.au), and
2
Natural Environments Program,
Queensland Museum, Brisbane, Queensland, Australia
Abstract Parasitoids play an important role in ecosystem functioning through their influence on herbivorous
insect populations. Theoretical and experimental evidence suggest that increased species richness can enhance and
stabilize ecosystem function. It is important to understand how richness-driven functional relationships change
across environmental gradients. We investigated how temperature affected the relationship between parasitoid
richness and parasitism rate in a guild of gall-parasitoids along an elevational gradient.We collected galls at 15 sites
along five elevational gradients (between 762 m and 1145 m asl) on six occasions over a year. A total of 1902
insects, including 1593 parasitoids, were reared from 12 402 galls. Parasitism rate increased significantly with
temperature on all sampling occasions, except December and February.We found a significant, positive richness–
parasitism relationship. This relationship, however, was weaker at higher elevations which may be linked to
decreased functional efficiency of parasitoids at lower temperatures. Temporal variability in parasitism rate and
parasitoid richness were significantly related, regardless of temperature. A stable functional guild of this kind may
provide a more reliable ecosystem service under environmental changes.
Key words: Cecidomyiidae, elevation gradients, parasitism, rainforest, trophic interactions.
INTRODUCTION
Environmental gradients, by selecting for certain func-
tional traits, affect species in a non-random manner
and can determine how species interact with each
other (Hodkinson 2005). Elevational gradients, in
particular, create pronounced patterns in species dis-
tributions due to the contrasting environmental con-
ditions found over short geographical distances
(Körner 2007). In general, decreasing temperatures
with increasing elevation may constrain insect traits,
such as thermal tolerance and phenology, leading to
reduced species richness, turnover and functional
diversity (Hodkinson 2005). Usually studies of
elevational patterns in insect richness and assemblage
turnover focus on a single taxon or trophic level (e.g.
moths, Ashton et al. 2011; ants, Burwell & Nakamura
2011; dung beetles, Escobar et al. 2005), and
elevational effects on species interactions are seldom
taken into account (but see Maunsell et al. 2014).
Parasitoids provide an interesting case study as they
play significant roles in ecosystem functioning (LaSalle
& Gauld 1993), yet are constrained in their responses
to environmental gradients by changes in host traits.
Few studies have looked explicitly at how elevation
affects host–parasitoid interactions. Those that have,
have found that the decrease in parasitism rates with
increasing elevation is disproportionately greater than
that of their host populations (Virtanen & Neuvonen
1999; Maunsell et al. 2014). Such asymmetrical
responses may explain why most studies report a
decline in parasitism rates with increasing elevation
(Hodkinson 2005; Péré et al. 2013). In some cases,
parasitism rates reach zero before the upper elevational
limits of the host species, thereby creating high eleva-
tion refuges from parasitism (Randall 1982; Maunsell
et al. 2014). Even when present, the efficacy of
parasitoids may be reduced at higher elevations.This is
supported by laboratory experiments demonstrating
decreased parasitoid functional efficiency at lower
temperatures (Enkegaard 1994; Menon et al. 2002).
Other factors, however, may also be important
determinants of parasitism rates. Complementary use
of host resources by a guild of parasitoids may lead to
increased rates of parasitism (Snyder et al. 2008). Dif-
ferent parasitoid species may be both temporally (e.g.
egg vs. larval parasitoids) and spatially (e.g. ecto- vs.
endoparasitoids) separated on the same host species
(Bonsall et al. 2002; Hackett-Jones et al. 2009).
In addition, the relative abundances of different
parasitoids change seasonally (Sorribas et al. 2010).
Such complementary host use may lead to a positive
relationship between parasitoid richness and host
*Corresponding author.
Accepted for publication May 2015.
Austral Ecology (2015) ••, ••–••
© 2015 Ecological Society of Australia doi:10.1111/aec.12283