Theoretical models predict that brood guarding may evolve in situations where eggs are costly to produce or when handling times are long. This study reveals that females of the secondary hyperparasitoid Trichomalopsis apanteloctena guarded cocoon broods of Cotesia kariyai, a gregarious endoparasitoid. Hyperparasitoid females also monopolized host resources and protected their offspring by driving away other conspecific hyperparasitoid females. The females exhibited antagonistic behavior towards competitors through threatening body postures, biting and chasing. Using a video camera to determine how long a hyperparasitoid female attended and parasitized cocoons within a single host brood, it was found that after about 4 days, cocoon guarding behavior became much less apparent. Moreover, more than 90% of hosts were typically parasitized by a hyperparasitoid female over the course of 4 days after she commenced brood guarding. Observations of egg production during a female's lifetime revealed a physiological interval rhythm that typically lasted 3-4 days, which correlates almost exactly with the period during which the cocoons were guarded. To confirm the giving-up time for a host cocoon brood, hyperparasitoid females were given access to 24 h-old cocoon clusters, each containing 60-100 individual cocoons. Ninety percent of the females remained on cocoons for approximately 72 h. Furthermore, twenty-five percent of wasps continued attending and presumably guarding host cocoon broods for more than 138 h after the female first attended the brood. C. kariyai larvae pupate within a few hours of egression from their host and emerge as adults about 5 days (120 h) later. Therefore, many hyperparasitoid females continued to guard older host cocoons of greatly reduced quality as a resource for their progeny and some even after eclosion of the primary parasitoid. Late-brood guarding enabled a hyperparasitoid female to protect her own progeny from other hyperparasitoid females that readily attacked and killed them when she was removed. Our study thus reveals that extended guarding behavior is an adaptive mechanism that probably plays an important role in the survival of the original brood.
Eurytoma robusta Mayr (Chalcidoidea) exploits host galls either as a primary or secondary parasitoid, an entomophytophagous inquiline or occasionally even as a predator. We present data on its ecology and impact on gall densities and population trends of the gall fly Urophora cardui (L.) on Cirsium arvense (L.) Scop. Habitat preference, host gall selection, clutch size, and high incidence of superparasitism causing empty gall cells show that E. robusta, a generalist with a broad host spectrum, is relatively poorly adapted to parasitising U. cardui. The influence of E. robusta on U. cardui in the Belfort-Sundgau region (1970-2004), in the Upper Rhine Valley (1973-2004) and in north-eastern Bavaria (1977-2004), differed considerably. In the forests of the Upper Rhine Valley and the Belfort-Sundgau region, where U. cardui has relatively stable source-sink populations, E. robusta is present but not the dominant mortality factor of the gall fly. In most areas of north-eastern Bavaria U. cardui occurs in fragmented populations and short lived non-equilibrium metapopulations. In these systems E. robusta became more abundant over the last five years, which resulted in a high incidence of superparasitism, an increase in the number of empty gall cells and reduced gall quality. The greatly increased degree of parasitism and an excess of empty cells resulted recently in the collapse of most local populations of U. cardui in the study area south of Bayreuth (north-eastern Bavaria). Together with earlier records the data presented here suggest that in north-eastern Bavaria E. robusta cause fluctuations in the abundance of U. cardui, which have a periodicity of 5-7 years. A remarkable feature of the oligophagous E. robusta is its high fidelity to formerly abundant U. cardui populations, which, with declining host densities, leads to overexploitation, resulting in a high incidence of superparasitism and high larval mortality. The possible influence of the habitat structure on the effect of E. robusta on the population dynamics of U. cardui is discussed. Our data plus that of other authors suggest that, with regard to U. cardui, E. robusta can develop a temporary local host specialisation.
Parasitoid females may adjust offspring sex allocation according to the number and quality of hosts available. Because in solitary species only one offspring survives per host, already parasitized hosts are of low quality and generally rejected. Superparasitism (i.e., sequential oviposition by the same or different females) results in aggressive interactions and competition for nutritional resources among larvae. We examined variations in the offspring sex ratio of Dendrocerus carpenteri (Curtis) (Hymenoptera: Megaspilidae), a solitary ectoparasitoid developing as a hyperparasitoid on the prepupae and pupae of primary aphid parasitoids inside mummified aphids. Mated females produced a female-biased sex ratio of 0.433 (proportion of sons) when caged singly and provided with 12 mummies for 2 h; they parasitized an average of four mummies/h and rarely superparasitized. Superparasitism increased when two females were caged together and provided with 12 mummies, from 1.18 to 1.24 and 1.38 eggs/host parasitized in 1, 2 and 3 h, respectively. The offspring sex ratio became increasingly more female-biased with increase in superparasitism; however, sex ratio variations were not correlated with cohort size. One mated and one unmated female provided with 12 mummies and caged together for 1 h produced a mean cohort sex ratio of 0.645, which differed from the one predicted (0.717) by an algebraic model incorporating the assumptions that both females contribute equal numbers of offspring and that the mated female does not change her offspring-sex allocation strategy. The observed shift in the cohort sex ratio to an increased female-bias indicates that mated females of D. carpenteri change their behaviour when encountering parasitized mummies or a conspecific competitor in the same patch. By depositing fertilized rather than unfertilized eggs, a female can increase the proportion of her daughters among parasitoids competing for a diminishing host supply., Manfred Mackauer, Andrew Chow., and Obsahuje bibliografii
On the basis of a twenty-year investigation, the life-cycle of Torymus cyanimus Boheman (Hymenoptera: Torymidae), a hyperparasitoid of a gall-forming fly in the Volga-Kama region is described. This parasitoid is the top-consumer in a food chain on Cirsium setosum (Willdenow) Iljin, in which the herbivore is Urophora cardui L. (Diptera: Tephritidae) and the primary parasitoids belong to the genus Eurytoma (Hymenoptera: Eurytomidae). Mating and oviposition behaviour were studied, and the superparasitism and larval cannibalism investigated in the second parasitoid generation. The superparasitism in T. cyanimus evolved in connection with the ovipositor elongation, leading to eggs being laid later when the fly host has already been completely consumed by larvae of Eurytoma serratulae F. Hyperparasitism and larval cannibalism in the second generation of T. cyanimus might account for the evolution of hyperparasitism in this species.