Summary1. Reproduction and parasite defence are assumed to be costly activities for hosts, and therefore trade-offs might exist between reproduction and parasite defence. 2. Brood sizes were manipulated in a population of Great Tits (Parus major L.) to assess trade-offs between reproduction and parasite defence. Blood samples were taken from males and females during the late nestling phase, and parasite prevalence was compared among parents raising enlarged, reduced and control broods. 3. A higher prevalence of protozoan blood parasites dominated by Haemoproteus majoris was found among parents with enlarged broods, as compared with parents with control and reduced broods, respectively. Brood size manipulation did not affect parasite prevalence between the sexes or between age classes differently. 4. Mean parasite intensity (number of parasites per microscope field) among adult birds only infected with H. majoris was higher for parents with experimentally enlarged broods than controls and reduced broods, respectively. 5. These results support the hypothesis of a trade-off between parasite defence and reproduction.
Key-words: Blood parasites, brood size manipulation, Haemoproteus majoris, Parus major, trade-offFunctional Ecology (1997) 11, 358-364 * Present address: ETH Zürich, Experimental Ecology ETH-Zentrum NW, CH 8092 Zürich, Switzerland.
359Reproductive investment and parasite susceptibility the notion that the effects of an increased intensity of parasites may operate for a considerable time. Thus, the trade-off between reproductive effort and parasite defence could be one mechanism that operates in the observed patterns of costs of reproduction (reviewed by Lindén & Møller 1989).The aim of this study was to investigate whether experimentally increased reproductive effort (achieved by altering brood size) might affect the prevalence and intensity of avian blood parasites in Great Tits P. major. This particular population of Great Tits has previously been found to be infected by a variety of blood parasites with Haemoproteus majoris being by far the most common, infecting around 75% of the adults depending on year and age of the birds (Allander & Bennett 1994). The paper also compares results from similar experiments of trade-offs between reproductive effort and parasite defence in other Great Tit populations exposed to different environments and parasites (i.e. Norris et al. 1994;Richner et al. 1995).
Methods
STUDY SPECIES AND STUDY AREAThe study was carried out in 1993 using a population of Great Tits breeding on the island of Gotland in the Baltic, SE Sweden (5710'N, 1820'E). The study area contains about 950 nestboxes distributed among 13 different plots. Twelve plots consist of deciduous woodland with ash (Fraxinus excelsior), oak (Quercus robur), birch (Betula spp.), hazel (Corylus avellana) and hawthorn (Crataegus spp.), whereas one plot is mainly coniferous forest, dominated by pine (Pinus sylvestris) with some birch.The Great Tit is a small (c. 20 g), mainly insectivorous passerine bird, resident in ...