Biological Journal of the Linnean Society, 2007, 90, 509–516. With 1 figure
© 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 90, 509–516 509
Blackwell Publishing LtdOxford, UKBIJBiological Journal of the Linnean Society0024-4066© 2007 The Linnean Society of London? 2007
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Original Article
IMMUNE FUNCTION AND FORCEPS SIZE
M. J. RANTALA
ET AL
.
*Corresponding author. E-mail: marrant@cc.jyu.fi
Forceps size and immune function in the earwig
Forficula auricularia L.
MARKUS J. RANTALA
1,2
*, DEREK A. ROFF
1
and LIISA M. RANTALA
3
1
Department of Biology, University of California, Riverside, CA 92521, USA
2
Department of Biology, Section of Ecology, University of Turku, FIN-20014 Turku, Finland
3
Department of Biological and Environmental Science, University of Jyväskylä, PO Box 35, FIN-40351,
Jyväskylä, Finland
Received 6 October 2005; accepted for publication 20 June 2006
Females of many species select their mates on the basis of the size or intensity of sexual ornaments, and it has been
suggested that these provide reliable signals of a male’s ability to resist parasites and pathogens. European earwigs,
Forficula auricularia, are sexually dimorphic in forceps shape and length. Male forceps are used as weapons in male
contests for access to females, but recent findings suggest that females also choose males on the basis of their forceps
length. In the present study, we tested the hypotheses that in the European earwig, F. auricularia, the size of forceps
is correlated with immune function and that immune function differs between the sexes. We found that encapsula-
tion rate was not correlated with the length of forceps in either sex, but was negatively correlated with body size. By
contrast, lytic activity of the haemolymph increased with overall body size in both sexes but, in females, lytic activity
increased with relative forceps length whereas, in males, it decreased with relative forceps length. After accounting
for effects of body size, there was no remaining significant correlation in females but the negative correlation in
males remained. Furthermore, we found that males had higher encapsulation rate and higher lytic activity than
females, suggesting that males have stronger immune defence. The results of the study indicate that the size of
forceps in male earwigs does not reliably reflect male immune defence. © 2007 The Linnean Society of London,
Biological Journal of the Linnean Society, 2007, 90, 509–516.
ADDITIONAL KEYWORDS: encapsulation rate – immunocompetence – lytic activity – sexual ornament –
sexual selection.
INTRODUCTION
Indicator models of sexual selection propose that sec-
ondary sexual ornaments preferred by females may
reliably signal male health and vigour by acting as
handicaps (i.e. characters that impose costs to their
bearer) (Zahavi, 1975, 1977; Heywood, 1989; Grafen,
1990; but see also Getty, 2002). It has been suggested
that one such possible cost is an increased susceptibil-
ity to parasites and pathogens (Folstad & Karter,
1992), which could influence the development and
maintenance of secondary sexual ornaments through
two major mechanisms. First, in cases where the
acquisition of important metabolites for ornamental
development results in increased exposure to para-
sites and pathogens, ornaments may reveal an indi-
vidual’s ability to tolerate parasite exposure (Folstad
et al., 1994). Second, because development and/or
maintenance of secondary sexual ornaments may
impose costs to a male’s immune defence, they may
honestly signal a male’s parasite resistance (Folstad &
Karter, 1992). For example, if resources must be
diverted away from immune function to maximize the
expression of a trait, males may suffer increased
susceptibility to parasites and pathogens (Folstad &
Karter, 1992; Sheldon & Verhulst, 1996). Thus, only
males in good condition or with high immunocompe-
tence, or both, can afford to allocate resources to large
ornaments, which should produce a positive correla-
tion between immune defence and the expression of
costly secondary sexual characters (Westneat &
Birkhead, 1998; Rantala, Kortet & Vainikka, 2003b).