Ecological Entomology (2007), 32, 201–210
© 2007 The Authors
Journal compilation © 2007 The Royal Entomological Society 201
Introduction
Inbreeding depression is common in many taxa (see reviews in
Husband & Schemske, 1996; Lynch & Walsh, 1998), and is
predicted in large outbreeding populations as a consequence of
the accumulation of deleterious recessive mutations (Davenport,
1908; Bruce, 1910; Haldane, 1927). Furthermore, Lynch and
Walsh (1998) note that inbreeding depression tends to affect
traits directly related to fitness such as viability, rather than
morphometric traits. Inbreeding depression may be prevalent
for traits that are highly related to fitness, because such traits
often have low heritability and are strongly influenced by domi-
nance (Mousseau & Roff, 1987; Crnokrak & Roff, 1995;
DeRose & Roff, 1999).
Historically, inbreeding in bark beetles has been of interest
because of its regularity in the tribes Xyleborini, Dryocoetini,
Cryphalini, Corthylini, and Hyorrhynchini (Hamilton, 1967;
Kirkendall, 1993). These inbreeding scolytids generally feed on
ambrosia in xylem of trees or in small plant parts like seeds. Sib
mating occurs on hosts before dispersal. Such inbreeding groups
are often characterised by female-biased sex ratios, flightless
males, and haplodiploid sex determination by arrhenotoky
(Kirkendall, 1993) or paternal genome elimination (Brun et al.,
1995). Inbreeding-related behaviour has not been investigated
outside of these exclusively inbreeding tribes. Many species of
scolytids display the ancestral trait of phloem feeding (Normark
et al., 1999), where outbreeding is expected (Kirkendall, 1993).
Despite similarity in resource utilisation, such bark beetle sys-
tems can vary considerably with respect to the sex that initiates
host colonisation, the degree of polygamy, and the population
dynamics of outbreaks (Wood, 1982).
Considering this variation, the engraver beetle, I. pini, is a
fairly typical phloem-feeding species that normally does not
experience inbreeding. After dispersal, males colonise hosts
and begin constructing nuptial galleries under the bark (Pitman
et al., 1965), while releasing aggregation pheromones in frass
Correspondence: Michael J. Domingue, Department of Entomology,
College of Agricultural Sciences, Pennsylvania State University, College
Park, PA 16802, US.A. E-mail: mjd29@psu.edu
Inbreeding depression and its effect on intrinsic
population dynamics in engraver beetles
MICHAEL J . DOMINGUE and STEPHEN A . TEALE Department of Environmental and
Forest Biology, College of Environmental Science and Forestry, State University of New York, Syracuse, U.S.A.
Abstract. 1. Phloem-feeding bark beetles (Coleoptera: Curculionidae, Scolytinae)
generally disperse before mating, leading to expectations of outbreeding. New York and
British Columbia populations of engraver beetles ( Ips pini) were tested for inbreeding
depression using different methods. Among several traits measured, only the number of
offspring surviving to adulthood was strongly reduced by inbreeding.
2. There was no evidence of avoidance of inbreeding depression in two possible
mechanisms considered: differential male and female emergence times within full sib
broods, and early termination of brood construction in forced sib mating.
3. Sib-mated females lay more eggs and have longer galleries than those in outbred
crosses, despite a low rate of survival to adulthood for such eggs. This difference may
be due to the ability of engraver beetles to assess crowding in broods as larvae begin to
feed, and allows partial compensation for the effects of inbreed depression.
4. Population models assuming density-dependent generational effects were modified
to account for inbreeding depression. Inbreeding depression makes populations less
prone to cyclical behaviour, particularly at lower carrying capacities.
5. Inbreeding depression has not been previously measured in scolytids, nor has
inbreeding-related behaviour been explicitly considered outside of exclusively
inbreeding tribes.
Key words. Bark beetle, genetic load, inbreeding depression, Ips pini, population
ecology , sib mating.