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Great Spotted Cuckoos Improve Their Reproductive Success by Damaging Magpie Host Eggs

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Great Spotted Cuckoos Improve Their Reproductive Success by Damaging Magpie Host Eggs Anim. Behav., 1997, 54, 1227–1233 Great spotted cuckoos improve their reproductive success by damaging magpie host eggs MANUEL SOLER, JUAN J. SOLER & JUAN G. MARTINEZ Departamento de Biologı´a Animal y Ecologı´a, Facultad de Ciencias, Universidad de Granada (Received 10 July 1996; initial acceptance 4 September 1996; final acceptance 27 February 1997; MS. number: 5290) Abstract. Adult great spotted cuckoos, Clamator glandarius, damage the eggs of their magpie, Pica pica, host without removing or eating them. The number of damaged magpie eggs was recorded in 360 parasitized nests of which 62.2% contained between one and eight damaged magpie eggs. Egg- destroying behaviour may be adaptive if it reduces nestling competition and/or enhances the hatching success of the cuckoo. To clarify the role of egg destruction for the reproductive success of great spotted cuckoos, unparasitized magpie nests were experimentally parasitized (without egg damage) by intro- ducing cuckoo eggs or chicks. Egg damage was common in parasitized nests but the eggs were not damaged by the hosts. Egg damage increased the breeding success of the cuckoos, by both reducing the number of competing host chicks in the nest and increasing the likelihood that late-laid cuckoo eggs would hatch. ? 1997 The Association for the Study of Animal Behaviour Avian brood parasites lay eggs in the nests of parasite removes a host egg at laying because other birds. Many of them remove or damage at parasitized nests contain significantly fewer eggs least one host egg; the female parasite usually does than unparasitized nests (e.g. Post & Wiley 1977; this shortly before she lays her own egg (Friedman Zimmerman 1983; Brooker & Brooker 1989a), 1968; Payne 1977; Sealy 1992). Removal or and that the host egg is eaten by the parasite (Hoy destruction of a host egg has been reported in the & Ottow 1964; Scott et al. 1992). However, as has European cuckoo, Cuculus canorus (Ga¨rtner 1981; been shown previously (Soler 1990), a significantly Wyllie 1981; Davies & Brooke 1988) and in smaller clutch in parasitized than unparasitized almost all other species of the genera Cuculus nests may result from the removal of damaged (Friedman 1968; Jensen & Clinning 1974; eggs by the host, given that many birds eject their Moksnes & Røskaft 1987; Brooker & Brooker own cracked eggs (Kemal & Rothstein 1988; Soler 1989a), Chrysococcyx (Jensen & Clinning 1974; 1990). Gill 1983; Brooker et al. 1988; Brooker & Brooker The great spotted cuckoo, Clamator glandarius, 1989b) and Clamator (Mountfort & Ferguson- is an obligate brood parasite, which parasitizes Lees 1961; Valverde 1971; Soler 1990), as well as mainly magpies, Pica pica, in Europe, the carrion in less well known cuckoos such as Eudynamis and crow, Corvus corone, being its secondary host Scythrops (Brooker & Brooker 1989a). Similarly, (Cramp 1985). Nestlings of the great spotted cowbirds, Molothrus spp. (Mason 1980; Carter cuckoo do not eject the eggs or young of the host, 1986; Wolf 1987; Sealy 1992) and most species but the eggs generally hatch several days before of honey guides, Indicatoridae, cuckoo-weavers, the magpie’s eggs, and magpie chicks frequently Anomalospiza, and widow-birds, Viduinae, also starve in competition for food with the larger remove host eggs during parasitic laying cuckoo chick (Cramp 1985; Soler 1990; Soler & (Friedman 1960; Lack 1968). Soler 1991). Mountford & Ferguson-Lees (1961) Brood-parasitic females can damage, remove showed that great spotted cuckoo females remove or eat host eggs. It is usually inferred that the one or two host eggs at laying. However, in other studies, egg removal was never observed, although Correspondence: M. Soler, Departamento de Biologı´a damaged eggs in parasitized nests were frequent Animal y Ecologı´a, Facultad de Ciencias, Universidad de Granada, E-18071 Granada, Spain (email: (Valverde 1971; Soler 1990). Parasites destroying [email protected]). host eggs without removing or eating them has 0003–3472/97/111227+07 $25.00/0/ar970524 ? 1997 The Association for the Study of Animal Behaviour 1227 1228 Animal Behaviour, 54, 5 also been recorded in different Molothrus species which (58.4%) were parasitized. We weighed great (Hoy & Ottow 1964; Post & Wiley 1977; Mason spotted cuckoo chicks in 1992 and 1993 with a 1980; Carter 1986). 300 g Pesola spring balance 1–2 days before they Egg-damaging behaviour by great spotted left the nest. Although in these years we per- cuckoos may reduce nestling competition or formed several field experiments, we have enhance the hatching success of the cuckoo. The included data only from non-experimental nests. ‘reduction of nestling competition’ hypothesis We determined the number of damaged eggs by states that egg-destruction behaviour increases carefully examining all eggs in each nest. We survival of the parasite chick by reducing the recorded breeding parameters such as laying date, number of competing host chicks in the nest. The clutch size, number of eggs hatched and number ‘enhancement of hatching success’ hypothesis of fledglings from nearly all nests. states that egg-damaging behaviour increases the likelihood that late-laid cuckoo egg(s) will hatch because of the destruction of host eggs that would Experimental Procedures otherwise hatch earlier than those of the cuckoo To clarify the role of egg destruction for the and would cause the host to cease incubation breeding success of great spotted cuckoos we before the cuckoo egg had hatched. experimentally parasitized (without egg damage) We mention only adaptive hypotheses; how- unparasitized magpie nests by introducing real ever, it is not clear whether the crushing of host cuckoo eggs or chicks in three experiments. Since eggs during the laying of parasite eggs is an the success of cuckoos depends on the timing of adaptation. Cuckoo eggs may have become thick- laying relative to the laying sequence of the mag- shelled to prevent breakage during the hurried pie (see below), we simulated parasitism early and laying. This may have incidentally led to more late during the laying sequence. Each experiment frequent destruction of host eggs, but this was not was performed in a different year. The controls the selective force driving the evolution in the first were, in each case, from the same year as the place. If this were the case we would really be experiment. dealing with an exaptation rather than an adap- tation (Gould & Vrba 1982; but see Skelton 1992 for problems with terminology). Experiment 1 Our aims in this paper are to describe the To test the adaptation hypothesis for egg dam- egg-damaging behaviour of great spotted cuck- age by great spotted cuckoo females laying late oos, provide evidence that eggs are not damaged (after clutch completion), we experimentally intro- by the host and test experimentally both the duced one cuckoo chick recently hatched (0–1 reduction of nestling competition and the days old) into each of 15 unparasitized magpie enhancement of hatching success hypotheses. nests, where the oldest magpie chick was between 1 and 7 days old. Three experimentally parasitized nests that were depredated were not considered. MATERIALS AND METHODS In experimentally parasitized nests the cuckoo chick was 5.3 0.61 (X , N=12) days younger Field Work & & than the oldest magpie chick, while in naturally We carried out the field work during the breed- parasitized nests the cuckoo chick was 3.6&1.94 ing seasons of 1990–1995 (although some ob- (N=5) days older than the oldest magpie chick. servations were conducted in 1982–1983 and This difference was statistically significant (Mann– 1986–1988) in Hoya de Guadix, southern Spain Whitney U-test: z=2.8, P<0.01). We used as con- (37)18*N, 3)11*W), a high plateau (1000 m) with trols 13 naturally parasitized nests with only one sparse vegetation, cultivated cereals (especially cuckoo egg matched with respect to laying date barley), and many groves of almond trees, Prunus and area with experimental nests. dulcis, and holm oaks, Quercus rotundifolia. Magpies nest at high density in almond trees but also nest in oaks (Soler 1990). Experiment 2 We inspected nests usually twice weekly. Dur- We introduced one cuckoo egg into 11 unpara- ing 1990–1994 we found 692 magpie nests, 404 of sitized magpie nests again simulating late-laying Soler et al.: Egg damaging by great spotted cuckoos 1229 of the cuckoo egg. In two of these nests, the starvation rate on experimentally parasitized cuckoo egg hatched simultaneously with the mag- magpie chicks (magpie breeding success in experi- pie eggs, and we have not considered these fur- mentally parasitized nests=50.6&4.80% in ther. The eggs used in the experiment were laid experiment 1, 27.0&8.55% in experiment 2 and between 3 and 6 days later than the last magpie 32.7&5.37% in experiment 3). However, the egg in the experimental nest (X&=4.4&0.37, magpie is a brood-reduction strategist and com- N=9). As controls we used seven naturally para- petition among chicks is usually strong and chick sitized nests with only one cuckoo egg matched to starvation frequent (magpie breeding success in experimental nests with respect to laying date and unparasitized nests=50.5&1.89%, N=122, Soler area. et al., in press). Thus, only experiments 2 and 3 produced a higher starvation rate than that natu- rally occurring in unparasitized nests. The effect of Experiment 3 natural parasitism was usually stronger than that To see whether the reduction of nestling com- of experimental parasitism (magpie breeding suc- petition hypothesis also applies when cuckoos lay cess in naturally parasitized nests=8.9&4.11% in early in the breeding cycle of the host, we simu- experiment 1, 17.3&6.15% in experiment 2 and lated early laying of the cuckoo egg by experimen- 14.0&5.19% in experiment 3).
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