Population responses to extrinsic mortality can yield no change in number of survivors (compensation) or an increase in number of survivors (overcompensation) when the population is regulated by negative density-dependence. This intriguing response has been the subject of theoretical studies, but few experiments have explored how the source of extrinsic mortality affects the response. This study tests abilities of three functionally diverse predators, alone and combined, to induce (over)compensation of a prey population. Larval (Diptera: Culicidae) were exposed to predation by (Crustacea: Copepoda) (Diptera: Culicidae) (Diptera: Corethrellidae), all three in a substitutive design, or no predation. The number of survivors to adulthood, female size and development time, and a composite index of performance () were analysed. Predator treatment did not have a significant effect on total number of survivors, nor on number of males, suggesting mortality by predation was compensatory. Predation significantly affected number of female survivors, with a trend of more females produced with predation, though no tests were significant. Predation significantly increased female development rate and relative to no-predator control. A sensitivity analysis indicated that the change in the number of female adults produced was the largest contributing factor to the differences in among cohorts. While predation did not significantly increase overall production of adults, it did release survivors from density-dependent effects sufficiently to increase population performance. This study provides an empirical test of mechanisms by which predation may yield positive effects on a population of victims, a phenomenon predicted to occur across many taxa and food webs.

1. Condition-dependent traits can act as honest signals of mate quality, with fitter individuals being able to display preferred phenotypes. Nutrition is known to be an important determinant of individual condition, with diet known to affect many secondary sexual traits. 2. In butterflies, male chemical signalling plays an important role in female mate choice. Potential male sex pheromone components have been identified previously, although it is unclear what information they convey to the female. 3. In the present study, the effect of diet on androconial and genital compound production is tested in male . To manipulate larval diet, larvae are reared on three different host plants: , the preferred host plant, Passiflora vitifolia and Passiflora platyloba. To manipulate adult diet, adult butterflies are reared with and without access to pollen, a key component of their diet. 4. No evidence is found to suggest that adult pollen consumption affects compound production in the first 10 days after eclosion. There is also a strong overlap in the chemical profiles of individuals reared on different larval host plants. The most abundant compounds produced by the butterflies do not differ between host plant groups. However, some compounds found in small amounts differ both qualitatively and quantitatively. Some of these compounds are predicted to be of plant origin and the others synthesised by the butterfly. Further electrophysiological and behavioural experiments will be needed to determine the biological significance of these differences.

1. We investigated the effects of strong density-dependence on larval growth, development, and survival of the mosquito (Theobald). We tested the hypothesis that density reduction early in larval development could result in as many or more individuals surviving to adulthood (compensation or overcompensation, respectively), or increased reproductive performance via rapid development and greater adult size. 2. In a field study of a natural population of we tested for the effects of a 75% lower density on percent survivorship to adulthood, number of adults, development time, adult size, adult longevity, and size dependent fecundity. 3. We found no evidence for compensation or overcompensation in adult production, nor for effects of lower density on percent survivorship. Low density yielded significant increases in adult size, adult longevity, and size-dependent fecundity, and a decrease in development time. 4. Estimated per capita population growth rate was significantly greater in the low-density treatment than in the high-density treatment. We infer this difference resulted from greater per capita resources increasing female size and fecundity, and reducing development time. Greater per capita population growth could therefore result from early mortality of larvae, meaning that the hydra effect, which predicts greater equilibrium population with, as opposed to without, extrinsic mortality, may be possible for these mosquitoes.

1. Every year billions of insects engage in long-distance, seasonal mass migrations which have major consequences for agriculture, ecosystem services and insect-vectored diseases. Tracking this movement in the field is difficult, with mass migrations often occurring at high altitudes and over large spatial scales. 2. As such, tethered flight provides a valuable tool for studying the flight behaviour of insects, giving insights into flight propensity (e.g. distance, duration and velocity) and orientation under controlled laboratory settings. By experimentally manipulating a variety of environmental and physiological traits, numerous studies have used this technology to study the flight behaviour of migratory insects ranging in size from aphids to butterflies. Advances in functional genomics promise to extend this to the identification of genetic factors associated with flight. Tethered flight techniques have been used to study migratory flight characteristics in insects for more than 50 years, but have never been reviewed. 3. This study summarises the key findings of this technology, which has been employed in studies of species from six Orders. By providing detailed descriptions of the tethered flight systems, the present study also aims to further the understanding of how tethered flight studies support field observations, the situations under which the technology is useful and how it might be used in future studies. 4. The aim is to contextualise the available tethered flight studies within the broader knowledge of insect migration and to describe the significant contribution these systems have made to the literature.

Parasitoid wasps sting and inject venom in arthropod hosts, which alters host metabolism and development while keeping the host alive for several days, presumably to induce benefits for the parasitoid young.Here we investigate the consequences of host envenomation on development and fitness of wasp larvae in the ectoparasitoid , by comparing wasps reared on live unstung, previously stung, and cold-killed hosts. Developmental arrest and suppression of host response to larvae are major venom effects that occur in both stung and cold-killed hosts, but not unstung hosts; while cold-killed hosts lack venom effects that require a living host. Thus, cold-killed hosts mimic some of the effects of venom, but not others.Eggs placed on live unstung hosts have significantly higher mortality during development, however successfully developing wasps from these hosts have similar lifetime fecundity to wasps from cold-killed or stung hosts. Therefore, although venom is beneficial, it is not required for wasp survival.While wasps developing on cold-killed versus stung hosts have similar fitness, multiple generations of rearing on cold-killed hosts results in significant fitness reductions of wasps.We conclude that the largest benefits of venom are induction of host developmental arrest and suppression of host response to larva (e.g. immune responses), although more subtle benefits may accrue across generations, or under stressful conditions.

1. Plants are frequently under attack by multiple insect herbivores, which may interact indirectly through herbivore-induced changes in the plant's phenotype. The identity, order, and timing of herbivore arrivals may influence the outcome of interactions between two herbivores. How these aspects affect, in turn, subsequently arriving herbivores that feed on double herbivore-induced plants has not been widely investigated. 2. This study tested whether the order and timing of arrival of two inducing herbivores from different feeding guilds affected the preference and performance of a subsequently arriving third herbivore, caterpillars of Mamestra brassicae L. (Lepidoptera: Noctuidae). Aphids [Brevicoryne brassicae L. (Hemiptera: Aphididae)] and caterpillars [Plutella xylostella L. (Lepidoptera: Yponomeutidae)] were introduced onto wild Brassica oleracea L. (Brassicaceae) plants in different sequences and with different arrival times. The effects of these plant treatments on M. brassicae caterpillars were assessed in pairwise preference tests and no-choice performance tests. 3. The caterpillars of M. brassicae preferred to feed from undamaged plants rather than double herbivore-induced plants. Compared with undamaged plants, they preferred plant material on which aphids had arrived first followed by caterpillars, whereas they avoided plant material with the reverse order of herbivore arrival. Performance of the caterpillars increased with increasing arrival time between herbivore infestations in double herbivore-induced plants. Although M. brassicae grew faster on plants induced by aphids than on those induced by caterpillars alone, its performance was not affected by the order of previous herbivore arrival. 4. These results imply that the timing of colonisation by multiple herbivores determines the outcome of plant-mediated herbivore-herbivore interactions.

Per-capita resource availability in aquatic habitats is influenced directly by consumer density via resource competition and indirectly via delayed resource competition when temporally non-overlapping cohorts of larvae exploit the same resources. In detritus-based systems, resources are likely to be influenced by the age of the aquatic habitat, as detritus changes in quality over time and may be replenished by new inputs.For aquatic insects that exploit detritus-based habitats, feeding conditions experienced during immature stages can influence fitness directly via effects on development and survivorship, but also indirectly by influencing adult traits such as fecundity and longevity.Larval habitat age and prior resource exploitation were manipulated in a field experiment using the container mosquito .It was found that from older habitats had greater larval survival, faster development and greater adult longevity. Exploitation of larval habitats by a prior cohort of larvae had a significant negative effect on subsequent cohorts of larvae by delaying development.It is suggested that extended conditioning of detritus probably resulted in conversion of recalcitrant resources to more available forms which improved the quality of the habitat.In a parallel study, evidence was found of carry-over effects of habitat age and prior exploitation on adult longevity for and collected from unmanipulated aquatic habitats.These results indicate the importance of detritus dynamics and the discontinuous nature of resource competition in these mosquito-dominated aquatic systems.

General theory from aquatic ecology predicts that smaller aquatic habitats have shorter hydroperiods favouring species that are better resource competitors and complete development quickly. Larger habitats are predicted to have longer hydroperiods enabling longer-lived predators to persist. Habitats with long hydroperiods and predators are predicted to favour slower-developing, predator resistant species, rather than competitive species.In a field experiment, we manipulated independently habitat size and hydroperiod in water-filled containers, to test these hypotheses about processes structuring aquatic communities. We used human-made containers that are dominated by mosquitoes that vary in desiccation resistance, competitive ability, and predation resistance.Habitat size and drying had significant effects on abundances of larvae of the common species in these communities. There was sorting of species by habitat size and by drying, with species that are better competitors relatively more abundant in smaller, more ephemeral habitats, and predator resistant, slower-developing species relatively more abundant in larger or permanently flooded habitats. There were no detectable effects of habitat size or drying on the dominant predator.Habitat size and its interaction with drying affected inputs of eggs to containers. Habitat size also affected relative abundances of the two dominant species in the egg population.Although habitat size and hydroperiod significantly affected composition of these communities, these impacts did not appear to be mediated through effects on predator abundance. Species specific differences in habitat size and drying regime preferences, and habitat-dependent larval performance appear to be the main forces shaping these communities.

The preference-performance hypothesis predicts that organisms lacking parental care should oviposit in habitats that optimize offspring performance. We investigated preference-performance relationships for the Asian tiger mosquito ( Skuse) and the southern house mosquito ( Say) (Diptera:Culicidae), two medically important container-breeding species, in response to an organic chemical blend mimicking decaying plant matter. Additionally, we evaluated the effects of long-term laboratory colonization of by using wild and laboratory strains.Oviposition bioassays were conducted by releasing gravid mosquitoes into field enclosures with automobile tires containing low and high concentrations of the chemical blend, and water controls. The offspring were then reared in water collected from the tires in which they were deposited. and wild laid more eggs in the chemical blend than water controls but did not differentiate between the low and high concentrations. Conversely, laboratory only preferred the high concentration to the low concentration. No statistical associations between oviposition preference and larval survival were found, as the chemical blend did not affect survivorship of either species.The oviposition preference for the chemical blend over water controls suggests that both species oviposit in the best available resource environment, but further studies are needed before conclusions regarding preference-performance relationships can be drawn.We found that long-term laboratory colonization affects oviposition behavior in , suggesting that behavioral studies on laboratory strains are not always applicable to wild populations.

1. Birds are considered to be the primary selective agents for warning colouration in butterflies, and select for aposematic mimicry by learning to avoid brightly coloured prey after unpleasant experiences. It has long been thought that bright colouration plays an important role in promoting the avoidance of distasteful prey by birds. 2. The hypothesis that warning colouration facilitates memorability and promotes predator avoidance was tested by means of a field experiment using distasteful model butterflies. Artificial butterflies with a Heliconius colour pattern unknown to local birds were generated using bird vision models, either coloured or achromatic, and hung in tree branches in a tropical forest. Two sequential trials were conducted at each site to test avoidance by naïve and experienced predators. 3. There was a significant reduction in predation in the second trial. Also, coloured models were attacked less than achromatic models. Specifically, coloured butterflies were attacked significantly less in the second trial, but there was no significant decrease in predation on achromatic models. 4. The present results imply an important role for colour in enhancing aversion of aposematic butterflies. It has also been demonstrated that previous experience of distasteful prey can lead to enhanced avoidance in subsequent trials, supporting mimicry theory.

Few studies have taken a comprehensive approach of measuring the impact of inter- and intra-specific larval competition on adult mosquito traits. In this study, the impact of competition and was quantified over the entire life of a cohort.Competitive treatments affected hatch-to-adult survivorship and development time to adulthood of females for both species, but affected median wing length of females only for . Competitive treatments had no significant effect on the median adult female longevity nor were there any effects on other individual traits related to bloodfeeding and reproductive success.Analysis of life table traits revealed no effect of competitive treatment on net reproductive rate (R) but there were significant effects on cohort generation time (T) and cohort rate of increase () for both species.Inter-specific and intra-specific competition among larvae may produce individual and population-level effects that are manifest in adults; however, benign conditions may enable resulting adults to compensate for some impacts of competition, particularly those affecting blood feeding success, fecundity, and net reproductive rate, R. The effect of competition, therefore, affects primarily larva - to - adult survivorship and larval development time, which in turn impact the cohort generation time, T and ultimately cohort rate of increase, .The lack of effects of larval rearing environment on adult longevity suggests that effects on vectorial capacity due to longevity may be limited if adults have easy access to sugar and blood meals.

Interactions between species can vary from mutually beneficial to evolutionarily neutral to antagonistic, even when the same two species are involved. Similarly, social interactions between members of the same species can lie on a spectrum from conflict to cooperation.The aim of the present study was to investigate whether variation in the two types of social behaviour are interconnected. Is the fitness of the various classes of social partner within species (such as parent and offspring, or male and female) differently affected by interactions with a second species? Moreover, can inter-specific interactions influence the outcome of social interactions within species?The present experiments focus on the interactions between the burying beetle Herbst and the phoretic mite G. Canestrini & R. Canestrini. The approach was to measure the fitness of burying beetle mothers, fathers, and offspring after reproduction, which took place either in the presence or absence of mites.We found that male, female, and larval burying beetles derive contrasting fitness costs and benefits from their interactions with the mite, despite sharing a common family environment. From the mite's perspective, its relationship with the burying beetle can, therefore, be simultaneously antagonistic, neutral, and possibly even mutualistic, depending on the particular family member involved. We also found that mites can potentially change the outcome of evolutionary conflicts within the family.We conclude that inter-specific interactions can explain some of the variation in social interactions seen within species. It is further suggested that intra-specific interactions might contribute to variation in the outcome of interactions between species.

In 2013, an opportunity arose in England to develop an agri-environment package for wild pollinators, as part of the new Countryside Stewardship scheme launched in 2015. It can be understood as a 'policy window', a rare and time-limited opportunity to change policy, supported by a narrative about pollinator decline and widely supported mitigating actions. An agri-environment package is a bundle of management options that together supply sufficient resources to support a target group of species. This paper documents information that was available at the time to develop such a package for wild pollinators. Four questions needed answering: (1) Which pollinator species should be targeted? (2) Which resources limit these species in farmland? (3) Which management options provide these resources? (4) What area of each option is needed to support populations of the target species? Focussing on wild bees, we provide tentative answers that were used to inform development of the package. There is strong evidence that floral resources can limit wild bee populations, and several sources of evidence identify a set of agri-environment options that provide flowers and other resources for pollinators. The final question could only be answered for floral resources, with a wide range of uncertainty. We show that the areas of some floral resource options in the basic Wild Pollinator and Farmland Wildlife Package (2% flower-rich habitat and 1 km flowering hedgerow), are sufficient to supply a set of six common pollinator species with enough pollen to feed their larvae at lowest estimates, using minimum values for estimated parameters where a range was available. We identify key sources of uncertainty, and stress the importance of keeping the Package flexible, so it can be revised as new evidence emerges about how to achieve the policy aim of supporting pollinators on farmland.

1. Africa faces serious challenges in feeding its rapidly growing human population owing to the poor productivity of maize and sorghum, the most important staple crops for millions of smallholder farmers in the continent, with yields being among the lowest in the world. 2. A complex of lepidopterous stemborers attack cereals in Africa. However, their effective control is difficult, largely as a result of the cryptic and nocturnal habits of moths, and protection provided by host stem for immature pest stages. Moreover, current control measures are uneconomical and impractical for resource-poor farmers. 3. An ecological approach, based on companion planting, known as 'push-pull', provides effective management of these pests, and involves combined use of inter- and trap cropping systems where stemborers are attracted and trapped on trap plants with added economic value ('pull'), and are driven away from the cereal crop by antagonistic intercrops ('push'). 4. Novel defence strategies inducible by stemborer oviposition have recently been discovered, leading to the attraction of egg and larval parasitoids, in locally adapted maize lines but not in elite hybrids. We also established that landscape complexity did not improve the ecosystem service of biological control, but rather provided a disservice by acting as a 'source' of stemborer pests colonising the crop. 5. Here we review and provide new data on the direct and indirect effects of the push-pull approach on stemborers and their natural enemies, including the mechanisms involved, and highlight opportunities for exploiting intrinsic plant defences and natural ecosystem services in pest management in smallholder farming systems in Africa.

1. was first recorded in Britain in 2004. Two subsequent earlier records were received from 2003. 2. The UK Ladybird Survey, a citizen science initiative involving online recording, was launched in 2005 to encourage people across Britain to track the spread of . Tens of thousands of people have provided records of and other species of ladybirds, creating an invaluable dataset for large-scale and long-term research. Declines in the distribution of seven (of eight assessed) native species of ladybird have been demonstrated, and correlated with the arrival of , using the records collated through the UK Ladybird Survey. 3. Experimental research and field surveys have also contributed to our understanding of the ecology of and particularly the process of invasion. arrived in Britain through dispersal and introduction events from regions in which it was deliberately released as a biological control agent. The rapid spread of this species has been attributed to its high natural dispersal capability by means of both flight and anthropogenic transport. A number of factors have contributed to the successful establishment and indeed dominance of this polymorphic species within aphidophagous guilds, including high reproductive capacity, intra-guild predation, eurytopic nature, high resistance to natural enemies within the invaded range, and potentially phenotypic plasticity. 4. The global invasion by and subsequent research on this species has contributed to the general understanding of biological invasions.

1. Moths (Lepidoptera) are the major nocturnal pollinators of flowers. However, their importance and contribution to the provision of pollination ecosystem services may have been under-appreciated. Evidence was identified that moths are important pollinators of a diverse range of plant species in diverse ecosystems across the world. 2. Moth populations are known to be undergoing significant declines in several European countries. Among the potential drivers of this decline is increasing light pollution. The known and possible effects of artificial night lighting upon moths were reviewed, and suggest how artificial night lighting might in turn affect the provision of pollination by moths. The need for studies of the effects of artificial night lighting upon whole communities of moths was highlighted. 3. An ecological network approach is one valuable method to consider the effects of artificial night lighting upon the provision of pollination by moths, as it provides useful insights into ecosystem functioning and stability, and may help elucidate the indirect effects of artificial light upon communities of moths and the plants they pollinate. 4. It was concluded that nocturnal pollination is an ecosystem process that may potentially be disrupted by increasing light pollution, although the nature of this disruption remains to be tested.

1. Organisms can respond to changing climatic conditions in multiple ways including changes in phenology, body size or morphology, and range shifts. Understanding how developmental temperatures affect insect life-history timing and morphology is crucial because body size and morphology affect multiple aspects of life history, including dispersal ability, while phenology can shape population performance and community interactions. 2. We experimentally assessed how developmental temperatures experienced by aquatic larvae affected survival, phenology, and adult morphology of dragonflies (). Larvae were reared under 3 environmental temperatures: ambient, +2.5 °C, and +5 °C, corresponding to temperature projections for our study area 50 and 100 years in the future, respectively. Experimental temperature treatments tracked naturally-occurring variation. 3. We found clear effects of temperature in the rearing environment on survival and phenology: dragonflies reared at the highest temperatures had the lowest survival rates, and emerged from the larval stage approximately 3 weeks earlier than animals reared at ambient temperatures. There was no effect of rearing temperature on overall body size. Although neither the relative wing nor thorax size was affected by warming, a non-significant trend towards an interaction between sex and warming in relative thorax size suggests that males may be more sensitive to warming than females, a pattern that should be investigated further. 4. Warming strongly affected survival in the larval stage and the phenology of adult emergence. Understanding how warming in the developmental environment affects later life-history stages is critical to interpreting the consequences of warming for organismal performance.

1. Modification of behaviors in the presence of predators or predation cues is widespread among animals. Costs of a behavioral change in the presence of predators or predation cues depend on fitness effects of lost feeding opportunities and, especially when organisms are sexually dimorphic in size or timing of maturation, these costs are expected to differ between the sexes. 2. Larval (Say) (Diptera: Culicidae) were used to test the hypothesis that behavioral responses of the sexes to predation cues have been selected differently due to different energy demands. 3. Even in the absence of water-borne predation cues, hungry females (the larger sex) spent more time browsing than did males, indicating a difference in energy needs. 4. In the presence of predation cues, well-fed larvae of both sexes reduced their activity more than did hungry larvae, and males shifted away from high-risk behaviors to a greater degree than did females, providing the first evidence of sex-specific antipredator behavior in foraging mosquito larvae. 5. Because sexual size dimorphism is common across taxa, and energetic demands are likely correlated with size dimorphism, this research demonstrates the importance of investigating sex specific behavior and behavioral responses to enemies and cautions against generalizing results between sexes.

1. Classical studies of succession, largely dominated by plant community studies, focus on intrinsic drivers of change in community composition, such as interspecific competition and changes to the abiotic environment. They often do not consider extrinsic drivers of colonization, such as seasonal phenology, that can affect community change. 2. We investigated both intrinsic and extrinsic drivers of succession for dipteran communities that occupy ephemeral pools, such as those in artificial containers. By initiating communities at different times in the season and following them over time, we compared the relative importance of intrinsic (i.e., habitat age) vs. extrinsic (i.e., seasonal phenology) drivers of succession. 3. We placed water-filled artificial containers in a deciduous forest with 20 containers initiated in each of three months. Containers were sampled weekly to assess community composition. Repeated-measures mixed-effects analysis of community correspondence analysis (CA) scores enabled us to partition intrinsic and extrinsic effects on succession. Covariates of temperature and precipitation were also tested. 4. Community trajectories (as defined by CA) differed significantly with habitat age and season, indicating that both intrinsic and extrinsic effects influence succession patterns. Comparisons of AICcs showed that habitat age was more important than season for species composition. Temperature and precipitation did not explain composition changes beyond those explained by habitat age and season. 5. Quantification of relative strengths of intrinsic and extrinsic effects on succession in dipteran and other ephemeral communities enables us to disentangle processes that must be understood for predicting changes in community composition.

1. Given sexual size dimorphism, differential mortality owing to body size can lead to sex-biased mortality, proximately biasing sex ratios. This mechanism may apply to mountain pine beetles, Hopkins, which typically have female-biased adult populations (2 : 1) with females larger than males. Smaller males could be more susceptible to stresses than larger females as developing beetles overwinter and populations experience high mortality. 2. Survival of naturally-established mountain pine beetles during the juvenile stage and the resulting adult sex ratios and body sizes (volume) were studied. Three treatments were applied to vary survival in logs cut from trees containing broods of mountain pine beetles. Logs were removed from the forest either in early winter, or in spring after overwintering below snow or after overwintering above snow. Upon removal, logs were placed at room temperature to allow beetles to complete development under similar conditions. 3. Compared with beetles from logs removed in early winter, mortality was higher and the sex ratio was more female-biased in overwintering logs. The bias increased with overwinter mortality. However, sex ratios were female-biased even in early winter, so additional mechanisms, other than overwintering mortality, contributed to the sex-ratio bias. Body volume varied little relative to sex-biased mortality, suggesting other size-independent causes of male-biased mortality. 4. Overwintering mortality is considered a major determinant of mountain pine beetle population dynamics. The disproportionate survival of females, who initiate colonisation of live pine trees, may affect population dynamics in ways that have not been previously considered.

1. Pea aphids ( Harris; Hemiptera: Aphididae) exhibit transgenerational wing polyphenism, in which unwinged females produce genetically identical winged offspring in response to environmental cues such as overcrowding and predation risk that indicate poor habitat quality. 2. Laboratory experiments were carried out to explore the intensity of the wing polyphenic response of pea aphids exposed to cues from ladybird predators and crowding, and their response was compared to pea aphids that were not exposed to any cues (control). 3. The study used cues from two different ladybird species: L. (Coleoptera: Coccinellidae) and Guérin-Méneville (Coleoptera: Coccinellidae) to investigate whether the wing polyphenic response of pea aphids to predator cues can be generalized 4. The intensity of the wing polyphenic response of pea aphids to crowding was found to be much stronger than their response to predator cues. There was no response to cues and the response to cues was mixed.