David J. Thompson

FLEXIBLE LARVAL GROWTH ALLOWS USE OF A RANGE OF HOST SIZES BY A PARASITOID WASP

Recent optimality models of host-parasitoid associations have assumed that host quality varies with host size or age at parasitism. This is based on the fact that larger hosts provide more resources, making size a reliable indicator of the amount of resources available for parasitoid development. Few studies have examined this in parasitoids that allow their hosts to continue development after parasitism (koinobiont parasitoids). In this study we compared growth trajectories of the koinobiont ichneumonid endoparasitoid Venturia canescens developing in four larval instars of one of its hosts, the moth Plodia interpunctella. Hosts were reared with excess food and parasitized as late second, third, fourth, and fifth instars. Hosts were dissected at intervals after parasitism, and host and parasitoid dry mass determined. The survivorship, development time, and size of eclosing adult wasps were also monitored. Hosts parasitized by Venturia continued to grow and become prepupae, although their maximum masses were progressively smaller the earlier that the hosts were parasitized: second-instar (L2) hosts grew to 70% of control (unpar- asitized) size, while L4 and L5 hosts reached control size. The maximum larval dry mass, as well as eclosing adult size, varied with host instar at parasitism. Venturia larvae spent prolonged periods as first instars when developing from L2 hosts and grew most rapidly during early stages in L5 larvae. The data show that for Venturia, host quality is not a direct function of host size at parasitism. Furthermore, the pattern of development shown by Venturia differs markedly from that shown by aphidiid koinobiont parasitoids.

Lifetime mating success in the damselfly Coenagrion puella

Abstract Lifetime mating success of male azure damselflies (Coenagrion puella) was measured in a natural population. The major determinant of mating success is the number of days a male spends at the breeding site, which is mostly determined by a males adult lifespan. Long-lived males have a higher mating rate than short-lived males, and daily mating rate increases with age up to 6 days, then falls. Large males live longer, but have a lower daily mating rate than small males. These effects of size are very weak, accounting for no more than 2% of the daily variance in mating success. The only overall effect of size on lifetime mating success is that males at both extremes of the size distribution are more likely to fail to mate. Chance differences in the number of females encountered are sufficient to account for the remaining variance in mating success. The weather is also shown to have a major effect on mating success. We draw attention to the ways in which it may be misleading to draw conclusions about the action of sexual selection from studies of daily, rather than lifetime, reproductive success. We provide evidence to support the view that variance in male reproductive success is neither evidence for sexual selection, nor a measure of its intensity.

Compatible genetic and ecological estimates of dispersal rates in insect (Coenagrion mercuriale: Odonata: Zygoptera) populations: analysis of 'neighbourhood size' using a more precise estimator.

Genetic and demographic estimates of dispersal are often thought to be inconsistent. In this study, we use the damselfly Coenagrion mercuriale (Odonata: Zygoptera) as a model to evaluate directly the relationship between estimates of dispersal rate measured during capture–mark–recapture fieldwork with those made from the spatial pattern of genetic markers in linear and two‐dimensional habitats. We estimate the ‘neighbourhood size’ (Nb) — the product of the mean axial dispersal rate between parent and offspring and the population density — by a previously described technique, here called the regression method. Because C. mercuriale is less philopatric than species investigated previously by the regression method we evaluate a refined estimator that may be more applicable for relatively mobile species. Results from simulations and empirical data sets reveal that the new estimator performs better under most situations, except when dispersal is very localized relative to population density. Analysis of the C. mercuriale data extends previous results which demonstrated that demographic and genetic estimates of Nb by the regression method are equivalent to within a factor of two at local scales where genetic estimates are less affected by habitat heterogeneity, stochastic processes and/or differential selective regimes. The corollary is that with a little insight into a species’ ecology the pattern of spatial genetic structure provides quantitative information on dispersal rates and/or population densities that has real value for conservation management.

Molecular and ecological evidence for small-scale isolation by distance in an endangered damselfly, Coenagrion mercuriale

Coenagrion mercuriale (Charpentier) (Odonata: Zygoptera) is one of Europes most threatened damselflies and is listed in the European Habitats directive. We combined an intensive mark‐release‐recapture (MRR) study with a microsatellite‐based genetic analysis for C. mercuriale from the Itchen Valley, UK, as part of an effort to understand the dispersal characteristics of this protected species. MRR data indicate that adult damselflies are highly sedentary, with only a low frequency of interpatch movement that is predominantly to neighbouring sites. This restricted dispersal leads to significant genetic differentiation throughout most of the Itchen Valley, except between areas of continuous habitat, and isolation by distance (IBD), even though the core populations are separated by less than 10 km. An urban area separating some sites had a strong effect on the spatial genetic structure. Average pairwise relatedness between individual damselflies is positive at short distances, reflecting fine‐scale genetic clustering and IBD both within‐ and between‐habitat patches. Damselflies from a fragmented habitat have higher average kinship than those from a large continuous population, probably because of poorer dispersal and localized breeding in the former. Although indirect estimates of gene flow must be interpreted with caution, it is encouraging that our results indicate that the spatial pattern of genetic variation matches closely with that expected from direct observations of movement. These data are further discussed with respect to possible barriers to dispersal within the study site and the ecology and conservation of C. mercuriale. To our knowledge, this is the first report of fine‐scale genetic structuring in any zygopteran species.

The effects of a sublethal baculovirus infection in the Indian meal moth, Plodia interpunctella

1. Baculoviruses are often highly pathogenic to their insect hosts, but theoretical studies have shown that sublethal infections may also play an important role in host population dynamics. Previous experiments have suggested that deleterious effects are caused by sublethal infections in a number of host-pathogen interactions, but these studies have been criticized on the basis of the methodology used. 2. Using an improved bioassay technique, the dose- and age-dependent response of the Indian meal moth, Plodia interpunctella, to sublethal levels of its granulosis virus was investigated. For the first time, significant sublethal effects have been unequivocally demonstrated in the development rate and reproductive capacity of the host

Lifetime reproductive success in the solitary endoparasitoid, Venturia canescens

Parasitoid wasps have long been considered excellent organisms in studies examining the evolution of reproductive and life-history strategies. In examining the lifetime reproductive success of parasitoids in the laboratory, most investigations have provided the insects with excess hosts and food, where they exist in a relatively constraint-free environment. Importantly, these conditions may not accurately reflect the true heterogeneity of natural systems, where suitable hosts and food sources are likely to be limiting. This study examines the influence of differences in host and food availability on reproductive and life-history parameters in an asexual strain of the solitary endoparasitoid, Venturia canescens (Hymenoptera: Ichneumonidae). Lifetime reproductive success in V. canescens was measured in response to temporal variations in host and food (honey solution) access. Cohorts of parasitoids were provided with 200 fifth-instar larvae of the Indian meal moth, Plodia interpunctella (Lepidoptera: Pyralidae), and food for variable periods daily after eclosion. V. canescens is synovigenic, and host-deprived wasps continued to mature eggs over the first few days after eclosion until the egg storage capacity was reached in the oviducts. When these wasps were subsequently provided with hosts, oogenesis resumed and continued until later in adult life. Constantly fed wasps lived longer and produced more progeny than wasps from cohorts which were alternately fed and starved or were starved from eclosion. Moreover, wasps with constant host and food access produced most progeny early in life and usually experienced prolonged periods of postreproductive survival. In contrast, the reproductive period of wasps with limited host access was more evenly distributed throughout the adult life. Consequently, the cumulative progeny production by V. canescens with constant food access was fairly uniform irrespective of host availability. Longevity and fecundity in V. canescens were positively correlated with adult size. However, variable host access had little effect on the longevity of wasps which were constantly supplied with honey. Over the first 2 days of adult life, variation in food access also had no effect on progeny production by V. canescens. We argue that manipulating temporal host and food access to parasitoids in the laboratory more closely approximates natural conditions, where these resources are likely to be spatially separated. Moreover, our findings suggest that many highly synovigenic parasitoids like V. canescens, which produce microtype (=hydropic) eggs, have a considerably higher reproductive potential than ovary dissections have revealed. Our findings are discussed in relation to life-history evolution in the parasitic Hymenoptera.

The impact of specialized enemies on the dimensionality of host dynamics

Although individual species persist within a web of interactions with other species, data are usually gathered only from the focal species itself. We ask whether evidence of a species’ interactions be detected and understood from patterns in the dynamics of that species alone. Theory predicts that strong coupling between a prey and a specialist predator/parasite should lead to an increase in the dimensionality of the preys dynamics, whereas weak coupling should not. Here we describe a rare test of this prediction. Two natural enemies were added separately to replicate populations of a moth. For biological reasons that we identify here, the prediction of increased dimensionality was confirmed when a parasitoid wasp was added (although this increase had subtleties not previously appreciated), but the prediction failed for an added virus. Thus, an imprint of the interactions may be discerned within time-series data from component species of a system.

Long-term population dynamics of the Indian meal moth Plodia interpunctella and its granulosis virus

1. Theoretical studies suggest that disease may play an important role in the population dynamics of insects, but there have been no long-term empirical studies that have examined this interaction in the absence of other important biotic and abiotic factors. 2. In the laboratory, three virus-free (VF) populations of Plodia interpunctella were compared with three virus-infected (VI) populations that were continuously exposed to a granulosis virus. Data sets lasted for between 10 and 17 generations. Census data of dead adults were collected twice weekly and spectral analysis of the data was used to complement the observed population fluctuations

The effect of superparasitism on development of the solitary parasitoid wasp, Venturia canescens (Hymenoptera: Ichneumonidae)

Abstract. 1 The effects of superparasitism on the rate of development, adult size and mortality of Venturia canescens (Hymenoptera: Ichneumonidae) were investigated. Parasitoids were reared from third (L3) and fifth (L5) instars of one of its hosts, Plodia interpunctella (Lepidoptera: Pyralidae) containing one, two or four parasitoid eggs. 2 Superparasitism increased the development time of Venturia reared from both instars, but the developmental delay was more evident in wasps from L5 hosts. 3 The size of parasitoids from L3 hosts was unaffected by egg number, but wasps from both superparasitized L5 treatments were significantly smaller than those from singly parasitized hosts. 4 Parasitoid mortality was significantly higher in L5 than L3, but within instars did not differ significantly with egg number. 5 The results confirm that superparasitism may affect the fitness of both the adult female wasp and her progeny, and should therefore be incorporated into models of superparasitism as an adaptive foraging strategy.

Transmission Dynamics of Bacillus thuringiensis Infecting Plodia interpunctella: A Test of the Mass Action Assumption with an Insect Pathogen

Central to theoretical studies of host-pathogen population dynamics is a term describing transmission of the pathogen. This usually assumes that transmission is proportional to the density of infectious hosts or particles and of susceptible individuals. We tested this assumption with the bacterial pathogen Bacillus thuringiensis infecting larvae of Plodia interpunctella, the Indian meal moth. Transmission was found to increase in a more than linear way with host density in fourth and fifth instar P. interpunctella, and to decrease with the density of infectious cadavers in the case of fifth instar larvae. Food availability was shown to play an important part in this process. Therefore, on a number of counts, the usual assumption was found not to apply in our experimental system.