Mate availability contributes to maintain the mixed‐mating system in a scolytid beetle

We investigated the mating system and population genetic structure of the beetle, Coccotrypes dactyliperda, with life history characteristics that suggest the presence of a stable mixed‐mating system. We examined the genetic structure of seven populations in Israel and found significant departures from the Hardy–Weinberg equilibrium and an excess of homozygosity. Inbreeding coefficients were highly variable across populations, suggesting that low levels of outbreeding occur in nature. Experiments were conducted to determine whether the observed high inbreeding in these populations is the result of a reproductive assurance strategy. Females reared in the laboratory took longer to mate with males from the same population (inbreeding) than with males from a different population (outbreeding). These results suggest that females delayed inbreeding, and were more inclined to outbreed when possible. Thus inbreeding, which predominates in most populations, may be due to a shortage of mates for outbreeding rather than a preference for inbreeding. We conclude that C. dactyliperda has a mixed‐mating system that may be maintained by a reproductive assurance strategy.     

Red anemone guild flowers as focal places for mating and feeding by Levant glaphyrid beetles

Several species of glaphyrid (Scarabaeoidea: Glaphyridae) beetles forage and mate on Mediterranean red bowl‐shaped flowers. In red anemones and poppies in Israel, female beetles occupy only a subset of the flowers, do not aggregate, and are hidden below the petals. This raises the question of how males find their mates. In the present study, we investigated the hypothesis that males and females orient to similar plant‐generated cues, thereby increasing their mate encounter prospects. Previous studies have demonstrated that beetle attraction to red models increases with display area. Choice tests with flowers and with models indicate that both male and female beetles prefer large displays. In anemones, beetles rest, feed, and mate mainly on male‐phase flowers, which are larger than female‐phase flowers. Poppies that contain beetles are larger than the population average. These findings support the hypothesis that males and females meet by orienting to large red displays. Corolla size correlates with pollen reward in both plant species, suggesting that visits to large flowers also yield foraging benefits. Male beetles often jump rapidly among adjacent flowers. By contrast to the preference for large flowers by stationary individuals, these jump sequences are random with respect to flower sex‐phase (in anemone) and size (in poppy). They may enable males to detect females at close range. We hypothesize that males employ a mixed mate‐searching strategy, combining orientation to floral signals and to female‐produced cues. The glaphyrids' preference for large flowers may have selected for extraordinarily large displays within the ‘red anemone’ pollination guild of the Levant. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 808–817.     

Trans‐generational effects of maternal rearing density on offspring development time in a parasitoid wasp

Maternal pre‐reproductive experience can impose phenotypic changes on offspring traits. These modifications may result from physiological constraints, although they can also increase the adaptation of offspring to their anticipated environment. Distinguishing between the two interpretations is often difficult. The effects of virgin female rearing density on their longevity and the characteristics of their male offspring are explored in the polyembryonic parasitoid wasp Copidosoma koehleri (Blanchard) (Encyrtidae: Hymenoptera). High rearing density may adversely affect maternal physiology or, alternatively, act as a cue for anticipated competition during the lives of the mothers and their offspring. Male offspring of group‐reared females reach pupation significantly sooner than male offspring of females reared alone. This accelerated development may provide an advantage when competition from superparasitising individuals is expected. The lifespan of high‐density females is longer than that of singly‐reared females, and their male offspring survive longer, suggesting that crowded rearing does not reduce the fitness of females or offspring. The shortened development time of male offspring may reflect an adaptive epigenetic response to predicted competitive conditions.     

REVIEW: The evolution of polyembryony in parasitoid wasps

Polyembryony has evolved independently in four families of parasitoid wasps. We review three main hypotheses for the selective forces favouring this developmental mode in parasitoids: polyembryony (i) reduces the costs of egg limitation; (ii) reduces the genetic conflict among offspring; and (iii) allows offspring to adjust their numbers to the quality of the host. Using comparative data and verbal and mathematical arguments, we evaluate the relative importance of the different selective forces through different evolutionary stages and in the different groups of polyembryonic wasps. We conclude that reducing the cost of egg limitation is especially important when large broods are favoured. Reducing genetic conflict may be most important when broods are small, thus might have been important during, or immediately following, the initial transition from monoembryony to polyembryony. Empirical data provide little support for the brood‐size adjustment hypothesis, although it is likely to interact with other selective forces favouring polyembryony.     

Time limitation affects offspring traits and female's fitness through maternal oviposition behaviour

Plasticity of various life‐history traits has evoked continuing interest among biologists. For example, the plasticity of offspring characteristics as well as maternal effects may be affected by time limitation and by limitation caused by changing environmental conditions. However, it is difficult to tell apart the effect of a time constraint, experienced by the mother, from food limitation, which is experienced by the offspring at the end of the season. In this study, we controlled for food limitation and simulated a time constraint for the mother. We tested how the seed beetle, Coccotrypes dactyliperda, adapts its reproductive investment after encountering a period of low availability of seeds as oviposition sites, as compared with females that encountered a seed at an early adult stage, while maintaining a similar food supply for offspring of both groups. We show that time limitation has a significant effect on the reproductive investment patterns of females. Females that were prevented from ovipositing, but provided with abundant food and later given oviposition sites, produced more, but smaller offspring than control females. Although the number of offspring increased, there was no indication of competition for food between offspring. We propose that, in order to compensate for the loss of time, mothers that experienced a shortage of oviposition sites influence their offspring to mature faster at the cost of a smaller than average body size. This study emphasizes the importance of considering more than one offspring generation in order to correctly estimate female fitness. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 728–736.     

Can clone size serve as a proxy for clone age?An exploration using microsatellite divergence in Populus tremuloides

In long‐lived clonal plants, the overall size of a clone is often used to estimate clone age. The size of a clone, however, might be largely determined by physical or biotic interactions, obscuring the relationship between clone size and age. Here, we use the accumulation of mutations at 14 microsatellite loci to estimate clone age in trembling aspen (Populus tremuloides) from southwestern Canada. We show that the observed patterns of genetic divergence are consistent with a model of increasing ramet population size, allowing us to use pairwise genetic divergence as an estimator of clone age. In the populations studied, clone size did not exhibit a significant relationship with microsatellite divergence, indicating that clone size is not a good proxy for clone age. In P. tremuloides, the per‐locus per‐year neutral somatic mutation rate across 14 microsatellite loci was estimated to lie between 6 × 10^?7 (lower bound) and 4 × 10^?5 (upper bound).     

Costs and consequences of superparasitism in the polyembryonic parasitoid Copidosoma koehleri (Hymenoptera: Encyrtidae)

Abstract.  1. Polyembryonic wasps provide dramatic examples of intra-specific developmental conflict. In these parasitoids, each egg proliferates into a clonal lineage of genetically identical larvae. If more than one egg is laid in a host (superparasitism), individuals of different clones may compete for food resources.
2. In the polyembryonic encyrtid Copidosoma koehleri , one larva per clone can differentiate into a sterile soldier. It is shown that soldiers are always females, and that they attack intra-specific competitors.
3. Research hypotheses were that (a) clones that develop in superparasitised hosts suffer heavier mortality than clones that develop in singly parasitised hosts, and (b) female clones cause higher mortality to their competitors than male clones, hence larval survival is lower in superparasitised hosts that contain females than in male-only broods.
4. The potential frequency of superparasitism in C. koehleri was manipulated by varying parasitoid–host ratios and exposure durations.
5. As parasitoid densities and exposure durations increased, the frequency of superparasitism rose, brood sizes increased, but the number of hosts that completed development was reduced. The number of offspring per parasitoid female decreased with increasing parasitoid–host ratios. Offspring size and longevity were inversely correlated with brood size. As superparasitism rates increased, fewer all-male broods were produced. Male–female broods were female-biased, suggesting selective killing of males by female soldiers. All-female broods were significantly smaller than all-male broods at high parasitoid densities only, possibly reflecting aggression among soldiers of competing clones.
6. The results support the working hypotheses, and suggest that female larvae outcompete males in superparasitised hosts.     

Host choice decisions in the polyembryonic wasp Copidosoma koehleri (Hymenoptera: Encyrtidae)

Female parasitoids often reject hosts of poor quality, where the survival and fitness of their offspring are expected to be low. In polyembryonic parasitoid wasps, a clone of genetically identical embryos develops from one egg in a host. In the wasp Copidosoma koehleri, each female clone produces one soldier larva that attacks competing clones inside the host. Aggression by soldiers is directed usually towards unrelated clones. Accordingly, it may be predicted that females will prefer nonparasitized over parasitized hosts, especially if the latter have been parasitized previously by a mated unrelated female, as a result of the reduced chances of survival for their offspring inside these hosts. In accordance with these predictions, females prefer nonparasitized hosts over self‐parasitized hosts when they are presented simultaneously. By contrast to the predictions, females prefer hosts parasitized by an unrelated conspecific over nonparasitized hosts when presented simultaneously. Females do not distinguish hosts parasitized by conspecifics from self‐parasitized hosts when presented simultaneously. They reject self‐parasitized hosts significantly more often than hosts parasitized by conspecifics when each host type is presented alone. Females faced with two previously parasitized hosts are not affected in their choice by the mating status (i.e. virgin or mated) of the previous parasitizing females. The combined results suggest that females are limited in their ability to assess the risk that their offspring will be attacked by a soldier, or that this risk is balanced by the relative advantages of ovipositing in a host parasitized by conspecifics. A possible advantage may be increased out‐breeding opportunities for the emerging offspring.