Summary. Few studies have investigated foraging decisions
in collectively foraging social insects with no studies in
termites. In termites predation is assumed to be a key
mortality factor. Therefore, we experimentally investigated
the role of predation pressure in foraging decisions of the
fungus cultivating, mound building termite Macrotermes
bellicosus in two habitats of the Comoé National Park (Ivory
Coast). We used the indirect approach of measuring the
Giving up Density (GUD), which is the amount of food left
when individuals stop foraging in a food patch, whilst exper-
imentally varying predation pressure. Three different condi-
tions were examined: (a) natural predation, (b) no predation,
and (c) experimental predation through artificial removal of
termites. In the shrub savanna, foraging termites responded
to increasing predation with increasing GUDs. By contrast,
in the gallery forest, there was no gradual response. Instead
termites abandoned a food patch immediately after an
attack by predators. Without predation GUDs were lower
in the savanna than in the gallery forest indicating that food
had a higher value in the former habitat. This, together with
the differential behavioral responses to predation, was in
accordance with high availability of food in the gallery
forest and a limited supply of food in the savanna. Thus,
according to our results termites traded off predation pres-
sure differently, according to the availability of food in both
habitats.
Key words: Giving up density, optimal foraging, depreda-
tion, termites.
Introduction
Factors influencing foraging decisions have been studied in a
wide range of solitarily foraging species since the formaliza-
tion of optimal foraging theory (e. g., Stephens and Krebs,
1986; Krebs and Kacelnik, 1991). However, for social insects
that forage collectively, and combine individual and cooper-
ative tasks, fewer data are available (e.g., ants: Breed et al.,
1987; Roces, 1990; Beckers et al., 1993; honeybees: Schmid-
Hempel et al., 1985; Wolf and Schmid-Hempel, 1990;
review: Detrain et al., 1999). Most studies concerned the
influence of food quantity and quality on foraging decisions
made by individual foragers within a foraging group of ants
(e. g. the influence of food quality on the load size of an indi-
vidual; Roces, 1990; Beckers et al., 1993). The combined
effect of these individual decisions on the exploitation of
food patches by a group of collectively foraging individuals
is less well known (Detrain et al., 1999). What is the outcome
of optimal foraging strategies of individuals on foraging pat-
terns of foraging groups, and thus the exploitation of food by
a colony?
Among social insects, termites are truly collective for-
agers. In comparison to ants and bees in which solitary work-
ers often carry out food search and recruit nestmates based
on an individual evaluation of resource quality, termites
appear to forage only in groups (Traniello and Leuthold,
2000). Termites have never been used to test any predictions
of models of optimal foraging theory (Traniello and
Leuthold, 2000). As predation is often assumed to be a key
mortality factor (Lepage, 1981; Abe and Darlington, 1985) in
termites, we investigated the influence of predation on forag-
ing decisions in collectively foraging termites.
To evaluate the influence of predation on foraging deci-
sions the indirect experimental approach of the Giving up
Density (GUD; e.g., Brown, 1988; Thorson et al., 1998) was
used. The GUD measures the amount of food left in a patch
when foraging individuals abandon that patch. This method
is based on the marginal value theorem of optimal foraging
theory (Charnov, 1976; Stephens and Krebs, 1986; Krebs and
Kacelnik, 1991). It assumes that a forager should stay in a
Insectes soc. 49 (2002) 264– 269
0020-1812/02/030264-06
© Birkhäuser Verlag, Basel, 2002
Insectes Sociaux
Research article
Evaluation of predation risk in the collectively foraging termite
Macrotermes bellicosus
J. Korb * and K. E. Linsenmair
Theodor Boveri Institut, Lehrstuhl für Tierökologie und Tropenbiologie, Am Hubland, D-97074 Würzburg, Germany,
e-mail: Judith.Korb@biologie.uni-regensburg.de; ke_lins@biozentrum.uni-wuerzburg.de
* Current address: Judith Korb, Biologie I, Universität Regensburg, D-93040 Regensburg, Germany, Fax: ++49 941 943 3304
Received 11 January 2002; revised 15 March 2002; accepted 10 April 2002.