The predatory mite Phytoseiulus persimilis manipulates imprinting among offspring through egg placement

A crucial question with respect to imprinting is how animals ensure that kin imprint on kin but not on non-kin. Imprinting takes place in a sensitive ontogenetic phase, usually in an early period of life or when offspring are produced, at which time the recipient imprints on the first referents met. In the predatory mite Phytoseiulus persimilis, imprinting among immature individuals happens in the larval stage immediately after hatching. I tested the hypothesis that adult P. persimilis females manipulate offspring imprinting by influencing the likelihood of encounters among recipients and referents via egg placement and egg aggregation. I conducted two experiments, one of which addressed imprinting and cannibalism, and the other addressed egg placement and egg aggregation. The imprinting experiment suggests that larvae imprint on any conspecific individual met in a sensitive ontogenetic phase and later on treat this individual as kin, irrespective of relatedness. After molting to protonymphs, imprinted individuals preferentially cannibalized unfamiliar to familiar larvae. Irrespective of familiarity, kin were cannibalized earlier than non-kin, suggesting the involvement of self-referent phenotype matching. The egg-placement experiment provides evidence that females adjust the aggregation level of their own eggs according to the degree of relatedness to present eggs from a previously ovipositing female. Both experiments in concert suggest that egg placement is a maternal strategy influencing imprinting among immature individuals. Apart from avoiding kin cannibalism, egg placement and imprinting by larvae may have relevance to other behaviors influenced by kin recognition, such as mate choice, prey-patch choice and dispersal.Communicated by L. Simmons     

Intraspecific hyperparasitism in a primary hymenopteran parasitoid

In solitary parasitoids, in which only one individual can emerge per host, the adaptive value of conspecific superparasitism is a function of the survival probability of the egg laid by the superparasitizing female. In the few cases which these probabilities are compared, the oldest immature has an advantage over the other individuals. We measured the acceptance rate of parasitized hosts and survival rate of supernumerary larvae in Anaphes victus (Hymenoptera: Mymaridae) in relation to the interval between ovipositions. When this interval was 5–7 days, the first immature was at the prepupa and pupa stage respectively, and female Anaphes victus changed their oviposition behavior markedly. They killed the developing parasitoid of their own species before ovipositing in it. The progeny of these females, which are normally primary parasitoids, developed thereafter as hyperparasitoids. Indeed, in contrast with other species, the survival of the second female's progeny increased with the time interval between ovipositions. This type of facultative intraspecific hyperparasitism is different from autoparasitism in Aphelinidae and has never been mentioned in other parasitoids; it would be adaptive if females of this short-lived species encounter low-quality patches.     

Female oviposition-site preference and egg hatching success in the damselfly Calopteryx splendens xanthostoma

By removing males and controlling the rate of water flow past oviposition patches in an experimental area, we showed that female Calopteryx splendens xanthostoma preferred to oviposit in fast flowing water rather than slow flowing water. A series of manipulations revealed the fitness benefits to females for this preference: eggs placed in fast-flowing water developed significantly faster and showed significantly lower mortality than eggs placed in slow-flowing water. A major factor determining this difference was the deposition and growth of encrusting algae which prevented the successful hatching of eggs in slow-flowing water.     

On the consequences of aggressive male mate-locating behaviour and micro-climate for female host plant use in the butterfly <Emphasis Type="Italic">Lycaena hippothoe</Emphasis>

The distribution of ecological resources and their significance for males and females may vary considerably. Intersexual behavioural interactions may lead, combined with particular resource configurations, to sexual spatial segregation. We investigated this issue relative to host plant use in females of the purple-edged copper butterfly, Lycaena hippothoe. Males exhibited nectar resource-based territoriality, which is an uncommon mate-locating system in butterflies. They perched and patrolled in large territories harassing every passing female. In our study system, the percentage of spatial dimension shared for adult and larval resources was estimated at 50%, and males monopolised 28% of the nectar-rich zones. Under these conditions of harassment, females travelled between nectar-rich zones for feeding and zones with suitable host plants for egg laying, but often without nectar and hence with low male density. This is likely to limit their time budget and, potentially, their realised fecundity as suggested by the low number of eggs found relative to population size. Females were also highly specialised in selecting host plants under particular environmental conditions. Using test choice in experimental cages, we showed that, in the absence of males, only micro-climatic conditions may significantly influencing egg-laying decisions. Moreover, results of egg-rearing experiments under different temperature treatments suggested that eggs were laid in thermally suitable micro-environments. The highly selective egg-laying behaviour can be viewed as a preference-performance choice. Knowledge of individuals' behaviour, including sexual interactions, can be highly significant for our understanding of habitat use, which in turn can be essential for conservation. We discuss this for L. hippothoe, a species of regional conservation concern.     

Parasitoid learning during interactions with ants: how to deal with an aggressive antagonist

Pauesia picta, P. pinicollis and P. silvestris (Hymenoptera: Aphidiidae) are common parasitoids of the conifer aphid Cinara pinea, which is regularly attended by red wood ants (Formica spp.). In this study, I tested whether females of these parasitoid species learned during interactions with honeydew-collecting Formica polyctena workers that caution is not necessary if searching behaviour is adapted, and whether parasitoids benefit from being able to learn. When searching on Scots pines, naive females of P. picta and P. pinicollis generally retreated to a pine needle when making contact with a honeydew-collecting ant, did not approach ants from the side or from the front and kept a ”safe distance” from ants when sitting on needles. After some non- aggressive ant encounters, experienced female parasitoids changed their behaviour: they reduced their searching speed, approached ants from the side and even from the front, retreated less often in response to an approaching ant and reduced the ”safe distance”. These experienced females had a significantly higher rate of oviposition than naive females or females foraging for an unattended host. Thus, the ability of the parasitoid to learn during interactions with an antagonist led to a prolonged retention time and a higher oviposition rate. By contrast, there was no evidence of learning in P. silvestris. Females of this species showed no behavioural change in response to ant encounters, and there was no difference in the foraging success of naive and experienced female parasitoids. Received: 7 December 1999 / Revised: 23 September 2000 / Accepted: 10 March 2000     

Larval host plant origin modifies the adult oviposition preference of the female European grapevine moth <Emphasis Type="Italic">Lobesia botrana</Emphasis>

According to the 'natal habitat preference induction' (NHPI) hypothesis, phytophagous insect females should prefer to lay their eggs on the host species on which they developed as larvae. We tested whether this hypothesis applies to the breeding behaviour of polyphagous European grapevine moth, Lobesia botrana, an important pest in European vineyards. We previously found that different grape cultivars affect several life history traits of the moth. Because the different cultivars of grapes are suspected to provide different plant quality, we tested the NHPI hypothesis by examining oviposition choice of L. botrana among three Vitis vinifera cultivars (Pinot, Chasselas and Chardonnay). In a choice situation, females of L. botrana that had never experienced grapes were able to discriminate between different grape cultivars and preferentially selected Pinot as an oviposition substrate. This 'naive' preference of oviposition could be modified by larval environment: Females raised on grapes as larvae preferred to lay eggs on the cultivar that they had experienced. Furthermore, experience of the host plant during adult emergence could be excluded because when pupae originating from our synthetic diet were exposed to grapes, the emerging adults did not show preference for the cultivar from which they emerged. The NHPI hypothesis that includes the two sub-hypothesis "Hopkins host selection principle" and "chemical legacy" may thus be relevant in this system.     

Daphne gnidium, a possible native host plant of the European grapevine moth Lobesia botrana, stimulates its oviposition. Is a host shift relevant?

Summary. We tested the hypothesis that Daphne gnidium is an ancestral host plant of Lobesia botrana. To this end, we studied the oviposition response of this moth to various aerial organs of this plant. Our results show that females prefer to lay eggs on daphne berries rather than on grapes (cv. chasselas) and that polar surface extracts of daphne berries and leaves strongly stimulate oviposition in a dosedependant way, whereas flower extracts are weakly stimulant and branch extracts are inactive or deterrent for oviposition. Furthermore, a total extract of daphne berries stimulates oviposition in semi-natural conditions when applied onto fresh grapes. Oviposition stimulants from berries were isolated by HPLC coupled with a dual-choice oviposition bioassay. The ecological value of D. gnidium for L. botrana is discussed and the possible use of oviposition stimulants from daphne, contrasting with the oviposition signal of the cultivated host plant, is proposed in the perspective of developing behavioural manipulation methods such as stimulo-deterrent diversionary strategies compatible with IPM programs.     

Host location behavior of Cotesia plutellae Kurdjumov (Hymenoptera: Braconidae) in ambient and moderately elevated ozone in field conditions

In field O₃-enrichment experiments increased herbivore densities have been reported, which could be due to negatively affected host location behavior of natural enemies. We addressed the impact of doubling background O₃ on the host location of the parasitoid Cotesia plutellae by conducting 24-h trials in an open-air O₃-fumigation system during two consecutive years. Two circles (radii 1.40 and 4.00 m) of Plutella xylostella-infested potted cabbage plants were placed in the O₃ and ambient plots. Female wasps were released into each plot from the center, and observed 5 times over a 24-h period to assess their host location capability. Thereafter, plants were kept in laboratory conditions until larvae pupation to determine parasitism rates. No significant differences were detected between ambient and O₃-enriched environments either in the number of wasps found in the field, or in the percentages of parasitized larvae. This suggests that moderately elevated O₃ will not affect the behavior of this parasitoid.     

The larval parasitoid <Emphasis Type="Italic">Microplitis croceipes</Emphasis> oviposits in conspecific adults

Microplitis croceipes (Hymenoptera: Braconidae) is a larval parasitoid of Helicoverpa/Heliothis spp. In the course of mass rearing of M. croceipes, we found that females oviposited in the conspecific adults in rearing cages. When 20 pairs of inexperienced females and males or of experienced females and males were reared in a cage, the males lived for 14–15 days and the females for 18–20 days on average. At their death, 37–42% of the males and 50–57% of the females contained conspecific eggs or first instar larvae in their abdominal cavity. When two of inexperienced females met on a host-infested leaf of soybean, they attempted to sting each other. Of the attacked females, 30% contained a conspecific egg laid in their abdomen. In abdominal cavity of the adults parasitized by a conspecific female, the majority of the parasitoid eggs laid disappeared within 1 day after oviposition. Only 10–30% of the parasitoid eggs laid in conspecific adults hatched 3–4 days after oviposition, but those larvae never molted to second instar. When the adults were stung by one or two conspecific females, their subsequent longevity was significantly shorter than that for the control adults. Oviposition in conspecific adults may be prevalent in other parasitic wasps that quickly oviposit without intensive host examination, and have cuticle and size of abdomen to be stung by conspeicifcs.     

Adaptive host choice and avoidance of superparasitism in the spawning decisions of bitterling (Rhodeus sericeus)

Choice of a site for oviposition can have fitness consequences. We investigated the consequences of female oviposition decisions for offspring survival using the bitterling, Rhodeus sericeus, a freshwater fish that spawns inside living unionid mussels. A field survey of nine bitterling populations in the Czech Republic revealed a significantly lower rate of release of juvenile bitterling from Anodonta cygnea compared to three other mussel species. A field experiment demonstrated that female bitterling show highly significant preferences for spawning in A. anatina, Unio pictorum, and U. tumidus. Within a species, female bitterling avoided mussels containing high numbers of bitterling embryos. Mortality rates of bitterling embryos in mussels were strongly density dependent and the strength of density dependence varied significantly among mussel species. Female preferences for mussels matched survival rates of embryos within mussels and females distributed their eggs among mussels such that embryo mortalities conformed to the predictions of an ideal free distribution model. Thus, female oviposition choice is adaptive and minimizes individual embryo mortality. Received: 6 October 1999 / Received in revised form: 7 January 2000 / Accepted: 13 March 2000