Abundance–range size relationships of macrolepidoptera in Britain: the effects of taxonomy and life history variables

1. A positive relationship between the local abundances and regional distributions of species in an assemblage has been observed for a wide variety of taxa, but its underlying mechanisms remain poorly understood.
2. Taxonomically controlled analyses of a group of the British macrolepidoptera (moths) find a strong, abundance–range size relationship.
3. Degree of polyphagy and habitat generalism explain significant within-taxon variation in both range size and abundance, with polyphages and habitat generalists having larger ranges and greater local abundances than specialists.
4. Taxa that feed in niches considered to be associated with a high diversity of parasitoids have significantly smaller range sizes than other taxa.
5. With the possible exception of body size, none of the life history variables examined explain significant variation around the range size–abundance relationship.     

Development of the solitary endoparasitoid Microplitis demolitor: host quality does not increase with host age and size

Abstract.  1. Many studies examining the relationship between host size, an index of host quality, and parasitoid fitness use development time and/or adult parasitoid size as currencies of fitness, while ignoring pre-adult mortality. Because the physiological suitability of the host may vary in different stages, sizes, or ages of hosts, a misleading picture of host quality may therefore be obtained in cases where fitness is based on only one or two developmental traits.
2. The development of the solitary koinobiont endoparasitoid Microplitis demolitor is examined in different larval age-classes of its host the soybean looper Pseudoplusia includens . Hosts were parasitised on days 1–8 after hatching from the egg, and development time, adult body size, and mortality of the parasitoid were compared.
3. A comparison of larval growth trajectories (using dry body mass) of M. demolitor revealed that parasitoid larvae attained over twice as much body mass in old hosts than in younger hosts. Similarly, adult parasitoid size at eclosion generally increased with host size, although parasitoids developing in smaller hosts lost a much lower proportion of mass between pupation and eclosion.
4. Overall egg-to-adult development was most rapid in intermediate-aged hosts, and longer in hosts at opposite ends of the age continuum. Moreover, parasitoid mortality varied non-linearly with host stage, and was generally higher in very young and older hosts.
5. Based on these results and other empirical data for koinobionts, it is argued that fitness functions in this group of parasitoids are not simply a positive function of host size or age, but instead may be distinctly dome-shaped, both patterns reflecting the degree of physiological and nutritional compatibility between the two organisms.     

How do host plant use and seasonal life cycle relate to insect body size: A case study on European geometrid moths (Lepidoptera: Geometridae)

We used European geometrid moths (>630 species) as a model group to investigate how life history traits linked to larval host plant use (i.e., diet breadth and host-plant growth form) and seasonal life cycle (i.e., voltinism, overwintering stage and caterpillar phenology) are related to adult body size in holometabolous insect herbivores. To do so, we applied phylogenetic comparative methods to account for shared evolutionary history among herbivore species. We further categorized larval diet breadth based on the phylogenetic structure of utilized host plant genera. Our results indicate that species associated with woody plants are, on average, larger than herb feeders and increase in size with increasing diet breadth. Obligatorily univoltine species are larger than multivoltine species, and attain larger sizes when their larvae occur exclusively in the early season. Furthermore, the adult body size is significantly smaller in species that overwinter in the pupal stage compared to those that overwinter as eggs or caterpillars. In summary, our results indicate that the ecological niche of holometabolous insect herbivores is strongly interrelated with body size at maturity.     

Variation in trichome density and resistance against a specialist insect herbivore in natural populations of Arabidopsis thaliana

Abstract.  1. The annual herb Arabidopsis thaliana is a prime model organism of plant molecular genetics, and is currently used to explore the molecular basis of resistance to herbivores. However, both the magnitude and the causes of variation in resistance among natural populations of A. thaliana are poorly known. The hypotheses (a) that resistance to a specialist herbivore, the diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), is positively related to the density of leaf trichomes, and (b) that oviposition preference by female moths is positively correlated to larval performance on different populations and maternal lines of the host plant were examined.
2. Variation in leaf trichome density and resistance to P. xylostella within and among six natural populations of A. thaliana in Sweden was quantified . Resistance was quantified by examining the number of eggs laid on plants exposed to ovipositing female moths and by monitoring larval development on plants of different origin.
3. Trichome density varied significantly among populations; for 4-week-old plants (exposed to ovipositing moths), it also varied significantly among maternal families within populations. The rate of oviposition varied significantly both among populations and among families within populations. This variation could partly be explained by a negative relationship between trichome density and egg number, and a positive relationship between plant size and egg number. Time to pupation, pupal mass, and time to adult eclosion did not vary among populations or maternal lines of the host plant, and offspring performance was not related to P. xylostella oviposition preference . The results indicate that A. thaliana populations may respond to selection for increased resistance to P. xylostella , and suggest that trichome production contributes to resistance against this specialist herbivore.     

The effects of body size on mating and reproduction in Brachinus lateralis (Coleoptera: Carabidae)

Abstract. 1. The causes and reproductive consequences of body size variation of Brachinus lateralis Dejean, a parasitic carabid beetle, were investigated.
2. Body size variation occurs within and between sites. Host size has a major influence on body size of the adult.
3. Fecundity is positively correlated with body size. Egg size is not correlated with body size.
4. Mating males tend to be larger than non-mating males. There is a positive correlation of body sizes in mating pairs.
5. Limited opportunity for host choice may maintain size variation despite the advantages of large size.
6. The non-random patterns of mating for a species without obvious intrasexual aggression suggest that subtle means of male-male competition or female choice may be important.     

Influence of host plant heterogeneity on the distribution of a birch aphid

Abstract.  1. The spatial and temporal abundance of the aphid Euceraphis betulae was investigated in relation to heterogeneity in host plant ( Betula pendula ) vigour and pathogenic stress. The performance of aphids feeding on vigorous and stressed foliage was also examined.
2. The plant stress and plant vigour hypotheses have been suggested as opposing ways in which foliage quality influences herbivore abundance. In many plants, however, vigorous growing foliage co-exists with stressed or damaged foliage.
3. There was a negative correlation between branch growth (vigour) and branch stress (leaf chlorosis), with the most vigorous branches displaying little or no stress, and the most stressed branches achieving poor growth. There was a similar negative correlation between vigour and stress at the level of individual trees, which themselves represented a continuum in quality.
4. At the beginning of the season, E. betulae were intermittently more abundant on vigorous branches than on branches destined to become stressed, but aphids became significantly more abundant on stressed branches later in the season, when symptoms of stress became apparent. Similar patterns of aphid abundance were seen on vigorous and stressed trees in the following year.
5. Euceraphis betulae performance was generally enhanced when feeding on naturally stressed B. pendula leaves, but there was some evidence for elevated potential reproduction when feeding on vigorous leaves too.
6. Overall, plant stress probably influences E. betulae distribution more than plant vigour, but the temporal and spatial variability in plant quality suggests that plant vigour could play a role in aphid distribution early in the season.     

Covariance of phenotypically plastic traits induces an adaptive shift in host selection behaviour

Flexibility in adult body size allows generalist parasitoids to use many host species at a cost of producing a range of adult sizes. Consequently, host selection behaviour must also maintain a level of flexibility as adult size is related to capture efficiency. In the present study, we investigated covariance of two plastic traits--size at pupation and host size selection behaviour-using Aphidius ervi reared on either Acyrthosiphon pisum or Aulacorthum solani, generating females of disparate sizes. Natal host was shown to change the ranking of perceived host quality with relation to host size. Parasitoids preferentially attacked hosts that corresponded to the size of the second instar of their natal host species. This resulted in optimal host selection behaviour when parasitoids were exposed to the same host species from which they emerged. Parasitoid size was positively correlated with host size preference, indicating that females use relative measurements when selecting suitable hosts. These coadapted gene complexes allow generalist parasitoids to effectively use multiple host species over several generations. However, the fixed nature of the behavioural response, within a parasitoid's lifetime, suggests that these traits may have evolved in a patchy host species environment.     

Grape variety affects female but also male reproductive success in wild European grapevine moths

Abstract.  1. For insect herbivores the quality of the larval host plant is a key determinant of their fitness. Only little attention, however, has been given to the effects of plants on mating success of males and its consequence for the reproductive output of their mates. In addition, almost all the studies that have investigated the influence of host plants on herbivore fitness components have been done in the laboratory, and less is known of these effects in natural conditions.
2. Using the phytophagous European grapevine moth ( Lobesia botrana Den. & Schiff., Lepidoptera: Tortricidae), we tested the influence of grape cultivars as larval food on the probability of acquiring a mate for both sexes, and on the reproductive output of females and males.
3. Results from this study stress the importance of larval host plants on the reproductive success of both sexes. Larval diet differentially affected mating success and reproductive output of male and female moths. Fecundity, egg size, and egg hatchability were significantly different when larvae were fed on particular grape cultivars.
4. A given cultivar that is of poor quality for females is generally also of poor quality for males. A cultivar, however, could be suitable for females but not for males and vice-versa. Apparently, the nutrients required for adult reproduction are not necessarily the same for males and females.
5. The important conclusion from this study is that evaluating the differential effect of host-plant species on traits associated with reproductive success of herbivores requires that the effects on both sexes be taken into account.     

Exploring stable pattern formation in models of tussock moth populations

1. The western tussock moth ( Orgyia vetusta ) at the University of California Bodega Marine Reserve (Sonoma County, California, USA) exhibits dense, localized populations in the midst of extensive habitats where variation in host plant quality or predator abundance is unable to explain the restricted extent of the outbreaks.
2. Two primary features suggest that the host patterning is intrinsically generated: (i) female tussock moths are wingless, producing a low effective dispersal distance for the hosts; and (ii) the tussock moth population is attacked by several species of widely dispersing wasp and fly parasitoids.
3. We consider a set of spatially explicit host–parasitoid models to examine whether intrinsically generated patterns are possible within this system. These models include a spatially extended Nicholson–Bailey model to examine general features of pattern formation in host–parasitoid systems, and two system-specific models, an individual-based simulation and a population-level analytic model, to examine the details of this empirical system.
4. Both stable patterning and rapid extinction of the host population are initial-condition dependent outcomes of the general and specific models, implying that an intrinsically generated stable host pattern is feasible within the tussock moth system.
5. Stable patterning is enhanced by a large parasitoid-to-host dispersal ratio, local host resource limitation, and increased parasitism at the host patch's edge.     

Age-dependent clutch size in a koinobiont parasitoid

Abstract.  1. The Lack clutch size theory predicts how many eggs a female should lay to maximise her fitness gain per clutch. However, for parasitoids that lay multiple clutches it can overestimate optimal clutch size because it does not take into account the future reproductive success of the parasitoid.
2. From egg-limitation and time-limitation models, it is theoretically expected that (i) clutch size decreases with age if host encounter rate is constant, and (ii) clutch size should increase with host deprivation and hence with age in host-deprived individuals.
3. Clutch sizes produced by ageing females of the koinobiont gregarious parasitoid Microplitis tristis Nees (Hymenoptera: Braconidae) that were provided daily with hosts, and of females ageing with different periods of host deprivation were measured.
4. Contrary to expectations, during the first 2 weeks, clutch size did not change with the age of the female parasitoid, neither with nor without increasing host-deprivation time.
5. After the age of 2 weeks, clutch size decreased for parasitoids that parasitised hosts daily. The decrease was accompanied by a strong decrease in available eggs. However, a similar decrease occurred in host-deprived parasitoids that did not experience egg depletion, suggesting that egg limitation was not the only factor causing the decrease in clutch size.
6. For koinobiont parasitoids like M. tristis that have low natural host encounter rates and short oviposition times, the costs of reproduction due to egg limitation, time limitation, or other factors are relatively small, if the natural lifespan is relatively short.
7. Koinobiont parasitoid species that in natural situations experience little variation in host density and host quality might not have strongly evolved the ability to adjust clutch size.     

Parasitoid load affects plant fitness in a tritrophic system

Plants attacked by herbivorous insects emit volatile compounds that attract predators or parasitoids of the herbivores. Plant fitness increases when these herbivorous insects are parasitized by solitary parasitoids, but whether gregarious koinobiont parasitoids also confer a benefit to plant fitness has been disputed. We investigated the relationship between parasitoid load of the gregarious Cotesia glomerata (L.) (Hymenoptera: Braconidae), food consumption by larvae of their host Pieris brassicae L. (Lepidoptera: Pieridae), and seed production in a host plant, Brassica nigra L. (Brassicaceae), in a greenhouse experiment. Plants damaged by caterpillars containing single parasitoid broods produced a similar amount of seeds as undamaged control plants and produced significantly more seeds than plants with unparasitized caterpillars feeding on them. Increasing the parasitoid load to levels likely resulting from superparasitization, feeding by parasitized caterpillars was significantly negatively correlated with plant seed production. Higher parasitoid brood sizes were negatively correlated with pupal weight of Cotesia glomerata , revealing scramble competition leading to a fitness trade-off for the parasitoid. Our results suggest that in this tritrophic system plant fitness is higher when the gregarious parasitoid deposits a single brood into its herbivorous host. A prediction following from these results is that plants benefit from recruiting parasitoids when superparasitization is prevented. This is supported by our previous results on down-regulation of synomone production when Brassica oleracea was fed on by parasitized caterpillars of P. brassicae . We conclude that variable parasitoid loads in gregarious koinobiont parasitoids largely explain existing controversies about the putative benefit of recruiting these parasitoids for plant reproduction.     

Gamete number and size correlate with adult size in the egg parasitoid Trichogramma euproctidis

In parasitoids, the size of the adult is influenced by the size and quality of the host in which it develops. Body size is generally positively correlated with several adult fitness proxies (fecundity, longevity, and mating capacity). The initial resources available to an individual can influence gamete production (sperm and oocytes), and the number and quality of gametes produced directly influence the expected fitness of both males and females. Gamete production in relation to adult body size was quantified in Trichogramma euproctidis (Girault) (Hymenoptera: Trichogrammatidae), a short‐lived egg parasitoid of lepidopteran species. To avoid host quality variation, male and female parasitoids of different body sizes were produced using superparasitism by allowing mated and virgin female parasitoids to oviposit on Trichoplusia ni Hübner (Lepidoptera: Noctuidae) eggs. Seminal vesicles and ovaries of their offspring were dissected to count oocytes and to measure sperm length and oocytes volume. Tibia length was also measured to estimate body size. The number of oocytes, volume of oocytes, maternal investment index [= (number of oocytes × mean volume of oocytes)/10 000] and sperm length were all significantly positively correlated to body size. These results show that initial resources acquired during larval stage induce phenotypic plasticity in gamete production in both male and female T. euproctidis. Whereas number of sperm and oocytes can influence the fitness of males and females through increased mating capacity and fecundity, variation in gamete size (sperm length and oocyte volume) could also affect the fitness of an individual through sperm and larval competition.     

Testing the resource concentration hypothesis with tarnished plant bug on strawberry: density of hosts and patch size influence the interaction between abundance of nymphs and incidence of damage

Abstract. 1. Attributes of patches with strawberry plants were manipulated experimentally to assess the impact of patch size and host density on the abundance of tarnished plant bug (Hemiptera: Miridae) and incidence of fruit damage.
2. The density of nymphs per inflorescence increased with patch size and host density for some but not all generations of plant bug, providing partial support for the resource concentration hypothesis, while emphasising the importance of replicating experiments across several generations of herbivore.
3. Increasing density of nymphs with increasing patch size and host density did not translate into a high incidence of damage, which may be due to the relatively low feeding impact of nymphs in patches with numerous fruits and to the relatively high perimeter-to-area ratio in small patches resulting in a relatively high proportion of damaged fruits.
4. The decreasing proportion of damaged fruits with increasing density of fruits per patch suggests that nymphs exhibit a saturating functional response to density of strawberry fruits. Functional responses of herbivores may be a critical yet overlooked component that influences interactions between insects and their host plant. In particular, concentrated plant resources may result in increased numbers of herbivores yet sill have a negligible influence on plant damage and/or fitness.     

Maternal effects and their consequences for offspring fitness in the Yellow Dung Fly

1. Maternal adult diet and body size influence the fecundity of a female and possibly the quality and the performance of her offspring via egg size or egg quality. In laboratory experiments, negative effects in the offspring generation have often been obscured by optimal rearing conditions.
2. To estimate these effects in the Yellow Dung Fly, Scathophaga stercoraria , how maternal body size and adult nutritional status affected her fecundity, longevity and egg size were first investigated.
3. Second, it was investigated how female age and adult nutritional experience, mediated through the effects of egg size or egg quality, influenced the performance of offspring at different larval densities.
4. Maternal size was less important than maternal adult feeding in increasing reproductive output. Without food restriction, large females had larger clutch sizes and higher oviposition rates, whereas under food restriction this advantage was reversed in favour of small females.
5. Offspring from mothers reared under nutritional stress experienced reduced fitness in terms of egg mortality and survival to adult emergence. If the offspring from low-quality eggs survived, the transmitted maternal food deficiency only affected adult male body size under stressful larval environments.
6. Smaller egg sizes due to maternal age only slightly affected the performance of the offspring under all larval conditions.     

How two different host species influence the performance of a gregarious parasitoid: host size is not equal to host quality

1. Hyssopus pallidus Askew (Hymenoptera, Eulophidae) is a gregarious ectoparasitoid of the two tortricid moths species Cydia molesta Busck and C. pomonella L. (Lepidoptera, Tortricidae). It paralyses and parasitizes different larval instars of both species inside the apple fruit, which leads to the death of the caterpillar. 2. We assessed the influence of host species characteristics and host food on the performance of the parasitoid female in terms of clutch size decisions and fitness of the F(1) generation. 3. A comparison of clutch size revealed that female parasitoids deposited similar numbers of eggs on the comparatively smaller C. molesta hosts as on the larger C. pomonella hosts. The number of parasitoid offspring produced per weight unit of host larva was significantly higher in C. molesta than in C. pomonella, which is contrary to the general prediction that smaller hosts yield less parasitoid offspring. However, the sex ratio was not influenced by host species that differed considerably in size. 4. Despite the fact that less host resources were available per parasitoid larva feeding on C. molesta caterpillars, the mean weight of emerging female wasps was higher in the parasitoids reared on C. molesta. Furthermore, longevity of these female wasps was neither influenced by host species nor by the food their host had consumed. In addition we did not find a positive relationship between adult female weight and longevity. 5. Parasitoid females proved to be able to assess accurately the nutritional quality of an encountered host and adjust clutch size accordingly. These findings indicate that host size is not equal to host quality. Thus host size is not the only parameter to explain the nutritional quality of a given host and to predict fitness gain in the subsequent generation.     

Testing the plant stress hypothesis: stressed plants offer better food to an insect herbivore

Temperature and nutrition are among the most important environmental factors affecting ectotherm growth. As temperature and host‐plant quality often co‐vary in nature, the interaction between the two is of potentially high ecological importance for herbivorous insects. We here use the temperate‐zone butterfly Pieris napi L. (Lepidoptera: Pieridae) to investigate interactive effects of larval rearing temperature and host‐plant quality (by manipulating water availability) on larval growth. As growth rates have been hypothesized to govern stress tolerance, we additionally assessed adult starvation resistance. Butterflies followed the ‘temperature‐size rule’, which states that body size increases at lower developmental temperatures, proximately caused by differences in growth increment, which resulted from increased consumption at the lower temperature. Larvae benefitted from feeding on stressed plants from the low‐water regime by having higher body mass, growth rate, and food conversion efficiency, thus supporting the plant stress hypothesis, which predicts that plant quality for herbivores should increase if stress is imposed on plants. Some effects of host‐plant quality on larval growth parameters were as strong as or even stronger than effects of temperature, whereas interactive effects between temperature and food quality were scarce. At the low temperature, adult starvation resistance was higher than at the higher temperature and females were more resistant than males, whereas plant water regime had no clear impact. No evidence was found for a trade‐off between growth rate and starvation resistance. This study illustrates the importance of considering effects of host‐plant quality along with variation in other environmental factors for estimating the impact of environmental changes on herbivorous species.     

Patterns of host use in solitary parasitoids (Hymenoptera, Ichneumonidae): field evidence from a homogeneous habitat

Teder, T., Tammaru, T. and Pedmanson, R. 1999. Patterns of host use in solitary parasitoids (Hymenoptera, Ichneumonidae): field evidence from a homogeneous habitat. - Ecography 22: 79-86.
We detected a significant inter- and intraspecific host preference on the level of individual host use in a system, in which three moth species (Lepidoptera: Noctuidae), feeding on a cattail Typha latifolia , are parasitized by three solitary parasitoid species (Hymenoptera: Ichneumonidae). The biology of the host species is similar but they exhibit remarkable inter- and intraspecific variance in body size. All the parasitoid species preferred the largest host species in this system whereas other host species were used only occasionally. We found that parasitoids which emerged from females of the preferred host species were larger than those which developed in males of the same species. Accordingly, two of the parasitoid species had a significant within-host-species preference: females of the largest moth species were used more often than males. No dependence of the preference pattern on host density was found. This pattern of host use is discussed in the light of the switching theory and the optimal host selection theory. Our results indicate that non-random host use by parasitoids may have significant effects on host populations and communities, and forms a potential selective factor against large body size in herbivorous insects. Unlike the majority of ichneumonid wasps, these three parasitoid species have no remarkable female-biased sexual size dimorphism, in accordance with the predictions of Charnov's sex allocation theory for this case, we did not observe any significant host quality dependent biases in sex allocation: there was no association between host sex and parasitoid sex, neither did parasitoid sex ratio differ between years with different host quality.     

Does immune function influence population fluctuations and level of parasitism in the cyclic geometrid moth?

Populations of the autumnal moth, Epirrita autumnata, exhibit cycles with high amplitudes in northernmost Europe, culminating in devastating outbreak densities at favourable sites. Parasitism by hymenopteran parasitoids has been hypothesised to operate with a delayed density dependence capable of producing the observed dynamics. It has also been hypothesised that insects in crowded conditions invest greatly in their immunity as a counter-measure to increased risk of parasitism and pathogen infections. Furthermore, inducible plant defences consequent to grazing by herbivorous insects may be linked to the performance of parasitoids and pathogens through increased immunocompetence of the herbivore feeding on the foliage, in which the defence induction has taken place. At ten sampling sites, we quantified larval abundance, outbreak status and percentage larval parasitism during an extended peak phase of a population cycle. These population level covariates, together with an individual pupal mass, were used to explain differences in the immune defence, measured as an encapsulation reaction to artificial antigen. We also conducted a field study for an investigation of the susceptibility of autumnal moth pupae to naturally occurring pupal parasitoids. We did not find obvious differences between the encapsulation rate of autumnal moths originating from the sites with different past and current larval densities and risks for parasitism. The best ranked statistical models included pupal mass and outbreak status as explanatory variables, although both showed only slight effects on the encapsulation rate. The host resistance test revealed positive relationships between the encapsulation rate, body size and percentage parasitism of the exposed pupae, indicating that pupal parasitoids chose, and/or survived better, in large host individuals irrespective of their encapsulation ability. Thus, our results do not provide support for the hypothesis that variation in the immune function drives or modulates population cycles of autumnal moths. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorised users.     

Optimum and maximum host sizes at parasitism for the endoparasitoid Hyposoter didymator (Hymenoptera: Ichneumonidae) differ greatly between two host species

Host size is considered a reliable indicator of host quality and an important determinant of parasitoid fitness. Koinobiont parasitoids attack hosts that continue feeding and growing during parasitism. In contrast with hemolymph-feeding koinobionts, tissue-feeding koinobionts face not only a minimum host size for successful development but also a maximum host size, because consumption of the entire host is often necessary for successful egression. Here we study interactions between a generalist tissue-feeding larval endoparasitoid, Hyposoter didymator Thunberg (Hymenoptera: Ichneumonidae) and two of its natural hosts, Spodoptera exigua Hübner and Chrysodeixis chalcites Esper (Lepidoptera: Noctuidae). Larvae of C. chalcites are up to three times larger than corresponding instars of S. exigua and also attain much higher terminal masses before pupation. We hypothesized that the range of host instars suitable for successful parasitism by H. didymator would be much more restricted in the large host C. chalcites than in the smaller S. exigua. To test this hypothesis, we monitored development of H. didymator in all instars of both host species and measured survival, larval development time, and adult body mass of the parasitioid. In contrast with our predictions, C. chalcites was qualitatively superior to S. exigua in terms of the survival of parasitized hosts, the proportion of parasitoids able to complete development, and adult parasitoid size. However, in both hosts, the proportion of mature parasitoid larvae that successfully developed into adults was low at the largest host sizes. Our results suggest that qualitative, as well as quantitative, factors are important in the success of tissue-feeding parasitoids.