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In nature, most parasitoids attack more than one host species, and nearly all hosts are attacked by several species of parasitoids. This opens many potential opportunities for interactions of invasive species with native parasitoid-host association networks in invaded communities. Despite this, few studies have examined the direct and indirect impacts of biological invasion on parasitoid-host associations. This review examines what is known of these relationships from the most recent literature and suggests future research priorities. We conclude that parasitoid-host association networks in invaded communities are complex, dynamic, and subject to trophic intrusions from invasive plants, herbivores, plant pathogens, parasitoids, and hyperparasitoids. Future studies should take a holistic systems approach to understanding the impact of biological invasion and its consequences in shaping community structure through altering existing native, coevolved parasitoid-host association networks., Competing Interests: Declaration of Competing Interest The authors declare no competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. No artifical intellegence assistance was used in writting, reviewing, and editing the manuscript., (Published by Elsevier Inc.)

Anthropogenic climate change, including temperature extremes, is having a major impact on insect physiology, phenology, behavior, populations, and communities. Hyperparasitoids (insects whose offspring develop in, or on, the body of a primary parasitoid host) are expected to be especially impacted by such effects due to their typical life history traits (e.g. low fecundity and slow development), small populations (being high on the food chain), and cascading effects mediated via lower trophic levels. We review evidence for direct and indirect temperature and climate-related effects mediated via plants, herbivores, and the primary parasitoid host species on hyperparasitoid populations, focusing on higher temperatures. We discuss how hyperparasitoid responses may feed back to the community and affect biological control programs. We conclude that despite their great importance, very little is known about the potential effects of climate change on hyperparasitoids and make a plea for additional studies exploring such responses., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Inc. All rights reserved.)

The most important aim of the integrated management of forest insect pests remains the prevention of insect outbreaks, which are a consequence of the interaction of many factors in forest ecosystems, including species composition, age and health of the forest, soil type, the presence of natural enemies, and climatic factors. Integrated pest management until now has been achieved using measures aimed at shaping the functioning of stands in a changing environment. The aim of this review is to summarize research on the use of entomopathogens (microorganisms and nematodes) in the management of forest insect pests and to identify the principal knowledge gaps. We briefly describe the main research directions on the use of pathogens and nematodes to control insect pests and discuss limitations affecting their implementation. Research on entomopathogens for the biocontrol of forest insects has provided a wealth of knowledge that can be used effectively to reduce insect populations. Despite this, few entomopathogens are currently used in integrated pest management in forestry. They are applied in inoculation or inundation biocontrol strategies. While the use of entomopathogens in forest pest management shows great promise, practical implementation remains a distant goal. Consequently, sustainable reduction of forest pests, mainly native species, will be largely based on conservation biological control, which aims to modify the environment to favor the activity of natural enemies that regulate pest populations. This type of biocontrol can be supported by a range of silvicultural measures to increase the resilience of stands to insect infestations. © 2023 Society of Chemical Industry., (© 2023 Society of Chemical Industry.)

Trypanosoma cruzi is a digenetic unicellular parasite that alternates between a blood-sucking insect and a mammalian, host causing Chagas disease or American trypanosomiasis. In the insect gut, the parasite differentiates from the non-replicative trypomastigote forms that arrive upon blood ingestion to the non-infective replicative epimastigote forms. Epimastigotes develop into infective non-replicative metacyclic trypomastigotes in the rectum and are delivered via the feces. In addition to these parasite stages, transitional forms have been reported. The insect-feeding behavior, characterized by few meals of large blood amounts followed by long periods of starvation, impacts the parasite population density and differentiation, increasing the transitional forms while diminishing both epimastigotes and metacyclic trypomastigotes. To understand the molecular changes caused by nutritional restrictions in the insect host, mid-exponentially growing axenic epimastigotes were cultured for more than 30 days without nutrient supplementation (prolonged starvation). We found that the parasite population in the stationary phase maintains a long period characterized by a total RNA content three times smaller than that of exponentially growing epimastigotes and a distinctive transcriptomic profile. Among the transcriptomic changes induced by nutrient restriction, we found differentially expressed genes related to managing protein quality or content, the reported switch from glucose to amino acid consumption, redox challenge, and surface proteins. The contractile vacuole and reservosomes appeared as cellular components enriched when ontology term overrepresentation analysis was carried out, highlighting the roles of these organelles in starving conditions possibly related to their functions in regulating cell volume and osmoregulation as well as metabolic homeostasis. Consistent with the quiescent status derived from nutrient restriction, genes related to DNA metabolism are regulated during the stationary phase. In addition, we observed differentially expressed genes related to the unique parasite mitochondria. Finally, our study identifies gene expression changes that characterize transitional parasite forms enriched by nutrient restriction. The analysis of the here-disclosed regulated genes and metabolic pathways aims to contribute to the understanding of the molecular changes that this unicellular parasite undergoes in the insect vector., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2023 Smircich, Pérez-Díaz, Hernández, Duhagon and Garat.)

Background: Trypanosomatid parasites are widely distributed in nature and can have a monoxenous or dixenous life-cycle. These parasites thrive in a wide number of insect orders, some of which have an important economic and environmental value, such as bees. The objective of this study was to develop a robust and sensitive real-time quantitative PCR (qPCR) assay for detecting trypanosomatid parasites in any type of parasitized insect sample., Methods: A TaqMan qPCR assay based on a trypanosomatid-conserved region of the α-tubulin gene was standardized and evaluated. The limits of detection, sensitivity and versatility of the α-tubulin TaqMan assay were tested and validated using field samples of honeybee workers, wild bees, bumblebees and grasshoppers, as well as in the human infective trypanosomatid Leishmania major., Results: The assay showed a detection limit of 1 parasite equivalent/µl and successfully detected trypanosomatids in 10 different hosts belonging to the insect orders Hymenoptera and Orthoptera. The methodology was also tested using honeybee samples from four apiaries (n = 224 worker honeybees) located in the Alpujarra region (Granada, Spain). Trypanosomatids were detected in 2.7% of the honeybees, with an intra-colony prevalence of 0% to 13%. Parasite loads in the four different classes of insects ranged from 40.6 up to 1.1 × 10 8 cell equivalents per host., Conclusions: These results show that the α-tubulin TaqMan qPCR assay described here is a versatile diagnostic tool for the accurate detection and quantification of trypanosomatids in a wide range of environmental settings., (© 2023. The Author(s).)

Parasite proliferations within/on the host form the basis of the outcome of all infectious diseases. However, within-host dynamics are difficult to study in vertebrates, as it requires regularly following pathogen proliferation from the start of the infection and at the organismal level. Invertebrate models allow for this monitoring under controlled conditions using population approaches. These approaches offer the possibility to describe many parameters of the within-host dynamics, such as rate of proliferation, probability to control the infection, and average time at which the pathogen is controlled. New parameters such as the Pathogen Load Upon Death and the Set-Point Pathogen Load have emerged to characterize within-host dynamics and better understand disease outcome. While contextualizing the potential of studying within-host dynamics in insects to build fundamental knowledge, we review what we know about within-host dynamics using insect models, and what it can offer to our knowledge of infectious diseases., (Copyright © 2022 Elsevier Inc. All rights reserved.)

Recent advances in insect-pathogen interactions have started to reveal the role of insect tissues and organs as natural infection routes for parasites and microbial pathogens. Here we summarize this information highlighting the micro- and macro-parasites that enter insects through distinct infection routes and link them to innate immune activity. We also examine whether the infection route determines the insect immune response and if the resulting immunological and physiological processes underpinning these different routes of infection are clearly distinct. Understanding how the infection route is associated with the robustness in insect host defense will help us identify conserved evolutionary and ecological patterns in order to design novel strategies for the management of destructive agricultural pests and disease vectors., (Copyright © 2022 European Federation of Immunological Societies. Published by Elsevier B.V. All rights reserved.)

Insect-associated microbes, including pathogens, parasites, and symbionts, influence the interactions of herbivorous insects and pollinators with their host plants. Moreover, herbivory-induced changes in plant resource allocation and defensive chemistry can influence pollinator behavior. This suggests that the outcomes of interactions between herbivores, their microbes and host plants could have implications for pollinators. As epizootic diseases occur at high population densities, pathogen and parasite-mediated effects on plants could have landscape-level impacts on foraging pollinators. The goal of this minireview is to highlight the potential for an herbivore's multitrophic interactions to trigger plant-mediated effects on the immunity and health of pollinators. We highlight the importance of plant quality and gut microbiomes in bee health, and how caterpillars as model herbivores interact with pathogens, parasites, and symbionts to affect plant quality, which forms the centerpiece of multitrophic interactions between herbivores and pollinators. We also discuss the impacts of other herbivore-associated factors, such as agricultural inputs aimed at decreasing herbivorous pests, on pollinator microbiomes., (Copyright © 2021 Elsevier Inc. All rights reserved.)

The complex nature of climate change-mediated multitrophic interaction is an underexplored area, but has the potential to dramatically shift transmission and distribution of many insects and their pathogens, placing some populations closer to the brink of extinction. However, for individual insect-pathogen interactions climate change will have complicated hard-to-anticipate impacts. Thus, both pathogen virulence and insect host immunity are intrinsically linked with generalized stress responses, and in both pathogen and host have extensive trade-offs with nutrition (e.g., host plant quality), growth and reproduction. Potentially alleviating or exasperating these impacts, some pathogens and hosts respond genetically and rapidly to environmental shifts. This review identifies many areas for future research including a particular need to identify how altered global warming interacts with other environmental changes and stressors, and how consistent these impacts are across pathogens and hosts. With that achieved we would be closer to producing an overarching framework to integrate knowledge on all environmental interplay and infectious disease events., (Copyright © 2021 Elsevier Inc. All rights reserved.)

Entomopathogenic nematodes (EPNs) have been successfully applied as biological control agents against above ground and soil stages of insect pests. However, for commercial application, it is crucial to mass culture these nematodes using in vitro liquid culture technology, as it is not attainable when using susceptible insects as hosts. Lobesia vanillana (Lepidoptera: Tortricidae) is regarded a sporadic pest of wine grapes in South Africa. The in vivo- and in vitro-cultured South African EPNs, Steinernema yirgalemense and Steinernema jeffreyense (Rhabditida: Steinernematidae), were evaluated against larvae and pupae of L. vanillana in laboratory bioassays. For larvae, high mortality was observed for all treatments: In vitro-cultured S. yirgalemense (98%) performed better than S. jeffreyense (73%), while within in vivo cultures, there was no difference between nematode species (both 83%). No significant difference was detected between in vivo- and in vitro cultures of the same nematode species. The LD50 of the in vitro-cultured S. yirgalemense, was 7.33 nematodes per larva. Mortality by infection was established by dissecting L. vanillana cadavers and confirming the presence of nematodes, which was > 90% for all treatments. Within in vitro cultures, both S. yirgalemense and S. jeffreyense were able to produce a new cohort of infective juveniles from L. vanillana larvae. Pupae, however, were found to be considerably less susceptible to EPN infection. This is the first study on the use of EPNs to control L. vanillana. The relative success of in vitro-cultured EPN species in laboratory assays, without any loss in pathogenicity, is encouraging for further research and development of this technology., Competing Interests: The authors have declared that no competing interests exist.

Taeniosis/cysticercosis and echinococcosis are neglected zoonotic helminth infections with high disease burden caused by tapeworms which circulate between definitive and intermediate host reflecting a predator-prey interaction. Taeniid eggs can remain vital for months, allowing arthropods to mechanically transport them to intermediate hosts. However, the multiple routes that arthropods provide as carriers of taeniid eggs are still often unregarded or not considered. This review focuses on the prevalence and importance of arthropods as carriers and spreaders of taeniid eggs in the epidemiology of taeniosis/cysticercosis and echinococcosis. Current scientific knowledge showed a relevant role of houseflies (Muscidae), blowflies (Calliphoridae), dung beetles (Scarabaeoidea), darkling beetles (Tenebrionidae), ground beetles (Carabidae) and skin beetles (Dermestidae) in the spread of taeniid eggs in the environment, which may favor the infection of new hosts through the direct ingestion of an insect or of contaminated food and water. At last, key research challenges are highlighted, illustrating that further knowledge on the topic is needed to develop and improve guidelines and actions to prevent taeniid infections worldwide., (Copyright © 2021 Elsevier B.V. All rights reserved.)

In this review, we summarize the current data on development of monoxenous trypanosomatids and phytomonads in various insects. Of these, Diptera and Hemiptera are the main host groups, and, consequently, most available information concerns their parasites. Within the insect body, the midgut and hindgut are the predominant colonization sites; in addition, some trypanosomatids can invade the foregut, Malpighian tubules, hemolymph, and/or salivary glands. Differences in the intestinal structure and biology of the host determine the variety of parasites' developmental and transmission strategies. Meanwhile, similar mechanisms are used by unrelated trypanosomatids, reflecting the limited range of options to achieve the same goal., Competing Interests: Declaration of Interests The authors declare no competing interests., (Copyright © 2021 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Insect parasitoids have evolved symbiotic interactions with several viruses and thousands of parasitoid species have established mutualistic associations with polydnaviruses (PDVs). While PDVs have often been described as virulence factors allowing development of immature parasitoids inside their herbivore hosts, there is increasing awareness that PDVs can affect plant-insect interactions. We review recent literature showing that PDVs alter not only host physiology, but also feeding patterns and composition of herbivore's oral secretions. In turn PDV-induced changes in herbivore phenotype affect plant responses to herbivory with consequences ranging from differential expression of plant defense-related genes to wider ecological effects across multiple trophic levels. In this opinion paper we also highlight important missing gaps to fully understand the role of PDVs and other parasitoid-associated viral symbionts in a plant-insect interaction perspective. Because PDVs negatively impact performance and survival of herbivore pests, we conclude arguing that PDV genomes offer potential opportunities for biological control., (Copyright © 2021 Elsevier Inc. All rights reserved.)

Xenorhabdus nematophila bacteria are mutualists of Steinernema carpocapsae nematodes and pathogens of insects. Xenorhabdus nematophila exhibits phenotypic variation between insect virulence (V) and the mutualistic (M) support of nematode reproduction and colonization initiation in the infective juvenile (IJ) stage nematode that carries X. nematophila between insect hosts. The V and M phenotypes occur reciprocally depending on levels of the transcription factor Lrp: high-Lrp expressors are M+V- while low-Lrp expressors are V+M-. We report here that variable (wild type) or fixed high-Lrp expressors also are optimized, relative to low- or no-Lrp expressors, for colonization of additional nematode stages: juvenile, adult and pre-transmission infective juvenile (IJ). In contrast, we found that after the bacterial population had undergone outgrowth in mature IJs, the advantage for colonization shifted to low-Lrp expressors: fixed low-Lrp expressors (M-V+) and wild type (M+V+) exhibited higher average bacterial CFU per IJ than did high-Lrp (M+V-) or no-Lrp (M-V-) strains. Further, the bacterial population becomes increasingly low-Lrp expressing, based on expression of an Lrp-dependent fluorescent reporter, as IJs age. These data support a model that virulent X. nematophila have a selective advantage and accumulate in aging IJs in advance of exposure to insect hosts in which this phenotype is necessary., (© 2020 Society for Applied Microbiology and John Wiley & Sons Ltd.)

The global spread of a parasitic mite ( Varroa destructor) has resulted in Deformed wing virus (DWV), a previously rare pathogen, now dominating the viromes in honey bees and contributing to large-scale honey bee colony losses. DWV can be found in diverse insect taxa and has been implicated in spilling over from honey bees into associated ("apiary") and other ("non-apiary") insects. Here we generated next generation sequence data from 127 insect samples belonging to diverse taxa collected from Hawaiian islands with and without Varroa to identify whether the mite has indirectly affected the viral landscapes of key insect taxa across bees, wasps, flies and ants. Our data showed that, while Varroa was associated with a dramatic increase in abundance of (predominantly recombinant) DWV in honey bees (and no other honey bee-associated RNA virus), this change was not seen in any other taxa sampled. Honey bees share their environment with other insect populations and exist as a homogenous group, frequently sharing common viruses, albeit at low levels. Our data suggest that the threat of Varroa to increase viral load in an apiary does not automatically translate to an increase in virus load in other insects living in the wider community.

O -Acetylation of polysaccharides predominantly modifies plant cell walls by changing the physicochemical properties and, consequently, the structure and function of the cell wall. Expression regulation and specific function of cell wall-acetylating enzymes remain to be fully understood. In this report, we cloned a previously identified stunted growth mutant named sucrose uncoupled1 ( sun1 ) in Arabidopsis ( Arabidopsis thaliana ). SUN1 encodes a member of the TRICHOME BIREFRINGEN-LIKE family, AtTBL37 AtTBL37 is highly expressed in fast-growing plant tissues and encodes a Golgi apparatus-localized protein that regulates secondary cell wall thickening and acetylation. In sun1 , jasmonate signaling and expression of downstream chemical defense genes, including VEGETATIVE STORAGE PROTEIN1 and BRANCHED-CHAIN AMINOTRANSFERASE4 , are increased but, unexpectedly, sun1 is more susceptible to insect feeding. The central transcription factor in jasmonate signaling, MYC2, binds to and induces AtTBL37 expression. MYC2 also promotes the expression of many other TBLs Moreover, MYC activity enhances cell wall acetylation. Overexpression of AtTBL37 in the myc2-2 background reduces herbivore feeding. Our study highlights the role of O- acetylation in controlling plant cell wall properties, plant development, and herbivore defense., (© 2020 American Society of Plant Biologists. All Rights Reserved.)

Endosymbiotic bacteria that obligately associate with entomopathogenic nematodes as a complex are a unique model system to study competition. These nematodes seek an insect host and provide entry for their endosymbionts. Through their natural products, the endosymbionts nurture their nematodes by eliminating secondary infection, providing nutrients through bioconversion of the insect cadaver, and facilitating reproduction. On one hand, they cooperatively colonize the insect host and neutralize other opportunistic biotic threats. On the other hand, inside the insect cadaver as a fighting pit, they fiercely compete for the fittest partnership that will grant them the reproductive dominance. Here, we review the protective and nurturing nature of endosymbiotic bacteria for their nematodes and how their selective preference shapes the superior nematode-endosymbiont pairs as we know today., Competing Interests: Declaration of Competing Interest The authors declare no competing financial interest., (Copyright © 2020 Elsevier B.V. All rights reserved.)

Different model systems have, over the years, contributed to our current understanding of the molecular mechanisms underpinning the various types of interaction between bacteria and their animal hosts. The genus Photorhabdus comprises Gram-negative insect pathogenic bacteria that are normally found as symbionts that colonize the gut of the infective juvenile stage of soil-dwelling nematodes from the family Heterorhabditis . The nematodes infect susceptible insects and release the bacteria into the insect haemolymph where the bacteria grow, resulting in the death of the insect. At this stage the nematodes feed on the bacterial biomass and, following several rounds of reproduction, the nematodes develop into infective juveniles that leave the insect cadaver in search of new hosts. Therefore Photorhabdus has three distinct and obligate roles to play during this life-cycle: (1) Photorhabdus must kill the insect host; (2) Photorhabdus must be capable of supporting nematode growth and development; and (3) Photorhabdus must be able to colonize the gut of the next generation of infective juveniles before they leave the insect cadaver. In this review I will discuss how genetic analysis has identified key genes involved in mediating, and regulating, the interaction between Photorhabdus and each of its invertebrate hosts. These studies have resulted in the characterization of several new families of toxins and a novel inter-kingdom signalling molecule and have also uncovered an important role for phase variation in the regulation of these different roles.

Parasitoids depend on other insects for the development of their offspring. Their eggs are laid in or on a host insect that is consumed during juvenile development. Parasitoids harbor a diversity of microbial symbionts including viruses, bacteria, and fungi. In contrast to symbionts of herbivorous and hematophagous insects, parasitoid symbionts do not provide nutrients. Instead, they are involved in parasitoid reproduction, suppression of host immune responses, and manipulation of the behavior of herbivorous hosts. Moreover, recent research has shown that parasitoid symbionts such as polydnaviruses may also influence plant-mediated interactions among members of plant-associated communities at different trophic levels, such as herbivores, parasitoids, and hyperparasitoids. This implies that these symbionts have a much more extended phenotype than previously thought. This review focuses on the effects of parasitoid symbionts on direct and indirect species interactions and the consequences for community ecology.

Monoxenous (one host) trypanosomatids from insects and other invertebrates can be introduced into axenic culture relatively easily and efficiently, allowing for their transfer from the field into the laboratory. Here we describe simple methods and alternative cultivation protocols, the wider application of which will allow substantial expansion of trypanosomatids available for research.

The extreme biological diversity of Oceanian archipelagos has long stimulated research in ecology and evolution. However, parasitic protists in this geographic area remained neglected and no molecular analyses have been carried out to understand the evolutionary patterns and relationships with their hosts. Papua New Guinea (PNG) is a biodiversity hotspot containing over 5% of the world's biodiversity in less than 0.5% of the total land area. In the current work, we examined insect heteropteran hosts collected in PNG for the presence of trypanosomatid parasites. The diversity of insect flagellates was analysed, to our knowledge for the first time, east of Wallace's Line, one of the most distinct biogeographic boundaries of the world. Out of 907 investigated specimens from 138 species and 23 families of the true bugs collected in eight localities, 135 (15%) were infected by at least one trypanosomatid species. High species diversity of captured hosts correlated with high diversity of detected trypanosomatids. Of 46 trypanosomatid Typing Units documented in PNG, only eight were known from other geographic locations, while 38 TUs (~83%) have not been previously encountered. The widespread trypanosomatid TUs were found in both widely distributed and endemic/sub-endemic insects. Approximately one-third of the endemic trypanosomatid TUs were found in widely distributed hosts, while the remaining species were confined to endemic and sub-endemic insects. The TUs from PNG form clades with conspicuous host-parasite coevolutionary patterns, as well as those with a remarkable lack of this trait. In addition, our analysis revealed new members of the subfamilies Leishmaniinae and Strigomonadinae, potentially representing new genera of trypanosomatids., (Copyright © 2019 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved.)

Mermithids (phylum Nematoda) and hairworms (phylum Nematomorpha) somehow drive their arthropod hosts into water, which is essential for the worms' survival after egression. The mechanisms behind this behavioural change have been investigated in hairworms, but not in mermithids. Establishing a similar mechanistic basis for host behavioural change between these two distantly related parasitic groups would provide strong convergent evidence for adaptive manipulation and insight into how these parasites modify and/or create behaviour. Here, we search for this convergence, and also contrast changes in physiology between hosts infected with immature and mature mermithids to provide the first ontogenetic evidence for adaptive manipulation by disentangling host response and pathology from the parasite's apparent manipulative effects. We used SWATH-mass spectrometry on brains of Forficula auricularia (earwig) and Bellorchestia quoyana (sandhopper), infected with the mermithids Mermis nigrescens and Thaumamermis zealandica , respectively, at both immature and mature stages of infection, to quantify proteomic changes resulting from mermithid infection. Across both hosts (and hairworm-infected hosts, from earlier studies), the general function of dysregulated proteins was conserved. Proteins involved in energy generation/mobilization were dysregulated, corroborating reports of erratic/hyperactive behaviour in infected hosts. Dysregulated proteins involved in axon/dendrite and synapse modulation were also common to all hosts, suggesting neuronal manipulation is involved in inducing positive hydrotaxis. Furthermore, downregulation of CamKII and associated proteins suggest manipulation of memory also contributes to the behavioural shift.

The defence reactions of insects to parasitic invaders are both varied and complex. Melanisation of pathogens is often an important step in insect immunity and can play a key role in isolating parasites. Within samples collected from a subalpine stream in New Zealand during two consecutive seasons (i.e., winter and spring), we observed and categorised different levels of melanotic encapsulation by aquatic insect larvae to dormant Gordius sp. hairworm (Phylum Nematomorpha) cysts, a relatively obscure group of parasites. Some of these insect species act as intermediate transport hosts in the complex life cycle of hairworms. Based on these new observations, we calculated the melanisation response for an abundant species of caddisfly larvae (Olinga sp.) using the proportion of non-melanised cysts per individual host. We tested the hypothesis that season and total number of cysts in an infected host impact its melanisation response. Also, we explored the effect of host body size on the total number of cysts it carries. We found that the total number of cysts does not affect the melanisation response of the host. Season did have an impact on the melanisation response in Olinga sp., with lower levels observed in the spring. Additionally, larger caddisfly larvae harboured more cysts than smaller ones. Since little is known about the cryptic interactions between hairworms and their intermediate hosts, this new information adds some complexity to this poorly understood group of parasites., (Copyright © 2019 Elsevier Inc. All rights reserved.)

Certain species of parasites have the apparent ability to alter the behaviour of their host in order to facilitate the completion of their own life cycle. While documented in hairworms (phylum Nematomorpha), the ability for mermithid parasites (from the sister phylum Nematoda) to force hosts to enter water remains more enigmatic. Here, we present the first experimental evidence in a laboratory setting that an insect which normally never enters open water (the European earwig Forficula auricularia) will readily enter the water when infected with a mermithid nematode (Mermis nigrescens). Only adult mermithids appear capable of inducing this polarising shift in behaviour, with mermithid length being a very strong predictor of whether their host enters water. However, mermithid length was only weakly associated with how long it took an earwig to enter water following the beginning of a trial. Considering the evidence presented here and its alignment with a proteomic investigation on the same host-parasite system, this study provides strong evidence for adaptive behavioural manipulation and a foundational system for further behavioural and mechanistic exploration.

Many animals depend on microbial symbionts to provide nutrition, defence or other services. Holometabolous insects, as well as other animals that undergo metamorphosis, face unique constraints on symbiont maintenance. Microbes present in larvae encounter a radical transformation of their habitat and may also need to withstand chemical and immunological challenges. Metamorphosis also provides an opportunity, in that symbiotic associations can be decoupled over development. For example, some holometabolous insects maintain the same symbiont as larvae and adults, but house it in different tissues; in other species, larvae and adults may harbour entirely different types or numbers of microbes, in accordance with shifts in host diet or habitat. Such flexibility may provide an advantage over hemimetabolous insects, in which selection on adult-stage microbial associations may be constrained by its negative effects on immature stages, and vice versa. Additionally, metamorphosis itself can be directly influenced by symbionts. Across disparate insect taxa, microbes protect hosts from pathogen infection, supply nutrients essential for rebuilding the adult body and provide cues regulating pupation. However, microbial associations remain completely unstudied for many families and even orders of Holometabola, and future research will undoubtedly reveal more links between metamorphosis and microbiota, two widespread features of animal life. This article is part of the theme issue 'The evolution of complete metamorphosis'.

Insect phenology is affected by climate change and main responses are driven by phenotypic plasticity and evolutionary changes. Any modification in seasonal activity in one species can have consequences on interacting species, within and among trophic levels. In this overview, we focus on synchronisation mismatches that can occur between tightly interacting species such as hosts and parasitoids or preys and predators. Asynchronies happen because species from different trophic levels can have different response rates to climate change. We show that insect species alter their seasonal activities by modifying their life-cycle through change in voltinism or by altering their development rate. We expect strong bottom-up effects for phenology adjustments rather than top-down effects within food-webs. Extremely complex outcomes arise from such trophic mismatches, which make consequences at the community or ecosystem levels tricky to predict in a climate change context. We explore a set of potential consequences on population dynamics, conservation of species interactions, with a particular focus on the provision of ecosystem services by predators and parasitoids, such as biological pest control., (Copyright © 2019. Published by Elsevier Inc.)

From 1 January 2018 came into force Regulation (EU) 2015/2238 of the European Parliament and of the Council of 25 November 2015, introducing the concept of "novel foods", including insects and their parts. One of the most commonly used species of insects are: mealworms (Tenebrio molitor), house crickets (Acheta domesticus), cockroaches (Blattodea) and migratory locusts (Locusta migrans). In this context, the unfathomable issue is the role of edible insects in transmitting parasitic diseases that can cause significant losses in their breeding and may pose a threat to humans and animals. The aim of this study was to identify and evaluate the developmental forms of parasites colonizing edible insects in household farms and pet stores in Central Europe and to determine the potential risk of parasitic infections for humans and animals. The experimental material comprised samples of live insects (imagines) from 300 household farms and pet stores, including 75 mealworm farms, 75 house cricket farms, 75 Madagascar hissing cockroach farms and 75 migrating locust farms. Parasites were detected in 244 (81.33%) out of 300 (100%) examined insect farms. In 206 (68.67%) of the cases, the identified parasites were pathogenic for insects only; in 106 (35.33%) cases, parasites were potentially parasitic for animals; and in 91 (30.33%) cases, parasites were potentially pathogenic for humans. Edible insects are an underestimated reservoir of human and animal parasites. Our research indicates the important role of these insects in the epidemiology of parasites pathogenic to vertebrates. Conducted parasitological examination suggests that edible insects may be the most important parasite vector for domestic insectivorous animals. According to our studies the future research should focus on the need for constant monitoring of studied insect farms for pathogens, thus increasing food and feed safety., Competing Interests: The authors have declared that no competing interests exist.

This study aimed to inventory the herbivore insects associated with Enterolobium contortisiliquum (Vell.) Morong (Fabaceae) fruits and seeds and their primary and secondary parasitoids. Six samples collected between May and October 2013 yielded 210 fruits, from which 326 insects of six orders emerged: Coleoptera, Hymenoptera, Lepidoptera, Diptera, Thysanoptera and Psocoptera. Coleoptera (five families) was represented by the seed consumers Merobruchus bicoloripes Pic, Stator sp. Bridwell (Chrysomelidae, Bruchinae), two species of Silvanidae, one species of Scolytinae (Curculionidae), one species of Nitidulidae and one species of Cerambycidae. The cerambycid was also observed forming galleries on fruit mesocarp. Immature individuals of Lepidoptera were observed consuming the fruits and seeds. From the seven Hymenoptera families, only two species were associated with Coleoptera, being Horismenus Walker sp. (Eulophidae) as parasitoid of M. bicoloripes, and Neoheterospilus falcatus (Marsh) (Braconidae) as parasitoid of Scolytinae. The Lepidoptera parasitoids represented four genera: Pseudophanerotoma Zetel, Chelonus Panzer (Braconidae), Orgilus Nees (Braconidae) and Goniozus Forster (Bethylidae). The host associations for the reared parasitoids Bracon Fabricius (Braconidae), Pimplinae sp. (Ichneumonidae) and Perilampus Forster (Perilampidae) were not confirmed. We obtained a single representative of Diptera (Tachinidae) associated with Lepidoptera hosts in this food web.

Some parasitoids modify the behavior of their hosts, benefiting themselves at the host's expense. This phenomenon is called 'manipulation', and current research on parasitoid manipulation of host behavior tends to fall into one of three categories. First, the frequency of manipulation and the magnitude of its benefits to the parasitoid remains unclear. Basic documentation of manipulations is thus a major research focus, with especially valuable recent data coming from spiders manipulated by Polysphincta wasps. Second, for a handful of systems, we now have sufficient phylogenetic and behavioral data to begin asking questions about how manipulation evolved. Finally, the field continues to probe the mechanisms through which parasitoids manipulate host behavior, and now examines the role of parasitoid symbionts in this interaction., (Copyright © 2019 Elsevier Inc. All rights reserved.)

Virus-insect interactions are primarily parasitic, yet diverse mutualistic interactions, some of which are symbiogenic, also occur. These viruses can modify insect physiology and behavior so that hosts can gain resistance against various biotic challenges like pathogen and parasites. In the recent past, many insect mutualistic viruses have been reported. Viruses can show symbiogenic interactions with some insects, which have been explored at the molecular level. However, understanding about molecular mechanisms for many of the mutualistic viruses is still enigmatic. Exploration of these interactions and its mechanism can shed light on phenomenon of virus mediated biotic stress resistance in insects., (Copyright © 2019 Elsevier Inc. All rights reserved.)

Insect populations are declining even in protected areas, but the underlying causes are unclear. Here, I consider whether the factors driving the loss of insect diversity include invasive and/or introduced insects transmitting pathogens to less-resistant native species. The introduction of insects into new areas for biocontrol, to promote pollination, or for mass rearing in insect farms, threatens the health and diversity of indigenous insects by the co-introduction of entomopathogens whose spillover is difficult to control. Even less virulent pathogens or covert infections can become lethal if environmental stressors weaken the resistance of indigenous host species in an additive, potentiating or synergistic manner. More research is needed to develop effective strategies that protect the health and diversity of native insects., (Copyright © 2019 The Author. Published by Elsevier Inc. All rights reserved.)

Studies have demonstrated that medication behaviors by insects are much more common than previously thought. Bees, ants, flies, and butterflies can use a wide range of toxic and nutritional compounds to medicate themselves or their genetic kin. Medication occurs either in response to active infection (therapy) or high infection risk (prophylaxis), and can be used to increase resistance or tolerance to infection. While much progress has been made over the last few years, there are also key areas that require in-depth investigation. These include quantifying the costs of medication, especially at the colony level of social insects, and formulating theoretical models that can predict the role of infection risk in driving micro-evolutionary and macro-evolutionary patterns of animal medication behaviors., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Some parasites manipulate host behavior by exploiting the host's behavioral control networks. This review explores two examples of this approach using parasites from opposite ends of the size spectrum, that is, viruses and parasitic insects. The first example explores the use of the gene (egt) by some baculoviruses to deactivate the hormone 20-hydroxyecdysone. Suppressing this chemical signal prevents the expression of behaviors that could reduce viral transmission. The second example explores how a parasitic wasp uses the host's immune/neural communication system to control host behavior. When a host's manipulated behavior requires complex neural coordination, exploitation of host behavioral control systems is likely to be involved. Simpler host behaviors can be induced by damage to host tissues., (Copyright © 2019 Elsevier Inc. All rights reserved.)

In insects, bacterial endosymbionts are known to influence the ecology of their hosts by modifying interactions with natural enemies such as parasitoids. Symbionts can modulate both parasitoid behavioral and/or physiological traits as well as host behaviors and life-history traits. Together these suggest that endosymbionts may impact the host range of parasitoids. For example, endosymbionts may narrow parasitoid host range through first, reducing parasitoid ability to locate hosts and/or larval survival, second, affecting fitness traits of the emerging adult parasitoid and/or third, modulating the outcome of interference and exploitative competition between parasitoid species. From both a fundamental and applied point of view, these symbiotic effects would influence the ecology and evolution of parasitoids and associated population-level processes and ecosystem services (e.g. biocontrol)., (Copyright © 2018. Published by Elsevier Inc.)

Divergence and escalation in defences promote chemical diversity in plants, and consequently the diversity of insect herbivores. This diversification cascades to insect parasitoids through direct effects on host herbivore susceptibility, changes in herbivore community composition, or disparity in plant volatiles. Large tropical plant genera represent an ideal model for studying these trends due to the high diversity of sympatric species and their insects. Novel measures of chemical structural similarity should be used to analyse evolutionary trends in both direct and indirect defences. Host chemical data need to be combined with detailed herbivore and parasitoid data. This will help to identify truly active compounds. Furthermore, resolved genomic phylogenies for plants and insects should be included to assign directionality in the processes., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Insects typically forage in complex habitats in which their resources are surrounded by non-resources. For herbivores, pollinators, parasitoids, and higher level predators research has focused on how specific trophic levels filter and integrate information from cues in their habitat to locate resources. However, these insights frequently build specific theory per trophic level and seldom across trophic levels. Here, we synthesize advances in understanding of insect foraging behavior in complex habitats by comparing trophic levels in specialist host-parasitoid-hyperparasitoid systems. We argue that resources may become less apparent to foraging insects when they are member of higher trophic levels and hypothesize that higher trophic level organisms require a larger number of steps in their foraging decisions. We identify important knowledge gaps of information integration strategies by insects that belong to higher trophic levels., (Copyright © 2018 The Authors. Published by Elsevier Inc. All rights reserved.)

Entomopathogenic nematodes in the genus Steinernema (Nematoda: Steinernematidae) have a mutualistic relationship with Xenorhabdus bacteria (Gram-negative Enterobacteriaceae). This partnership however, is pathogenic to a wide range of insect species. Because of their potent insecticidal ability, they have successfully been implemented in biological control and integrated pest management programs worldwide. Steinernema-Xenorhabdus-insect partnerships are extremely diverse and represent a model system in ecology and evolution to investigate symbioses between invertebrates and microbes. The reproductive fitness of the nematode-bacterium partnership is tightly associated, and maintenance of their virulence is critical to the conversion of the insect host as a suitable environment where this partnership can be perpetuated., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Hamilton's theory of inclusive fitness provides a framework for understanding the evolution of social behaviour between kin, including parental and alloparental care. Brood parasitism is a reproductive tactic in which parasites exploit the care of other individuals of the same species (conspecific parasitism) or different species (interspecific parasitism) to rear their brood. Here, drawing from examples in birds and social insects, we identify two insights into brood parasitism that stem from inclusive fitness theory. First, the kin structure within nests, or between neighbouring nests, can create a niche space favouring the evolution of conspecific parasitism. For example, low average relatedness within social insect nests can increase selection for reproductive cheats. Likewise, high average relatedness between adjacent nests of some birds can increase a female's tolerance of parasitism by her neighbour. Second, intrabrood conflict will be high in parasitized broods, from the perspective of both parasite and host young, relative to unparasitized broods. We also discuss offspring recognition by hosts as an example of discrimination in a kin-selected social behaviour. We conclude that the inclusive fitness framework is instructive for understanding aspects of brood parasite and host evolution. In turn, brood parasites present some unique opportunities to test the predictions of inclusive fitness theory. This article is part of the theme issue 'The coevolutionary biology of brood parasitism: from mechanism to pattern'.

Parasitic interactions are so ubiquitous that all multicellular organisms have evolved a system of defences to reduce their costs, whether the parasites they encounter are the classic parasites which feed on the individual, or brood parasites which usurp parental care. Many parallels have been drawn between defences deployed against both types of parasite, but typically, while defences against classic parasites have been selected to protect survival, those against brood parasites have been selected to protect the parent's inclusive fitness, suggesting that the selection pressures they impose are fundamentally different. However, there is another class of defences against classic parasites that have specifically been selected to protect an individual's inclusive fitness, known as social immunity. Social immune responses include the anti-parasite defences typically provided for others in kin-structured groups, such as the antifungal secretions produced by termite workers to protect the brood. Defences against brood parasites, therefore, are more closely aligned with social immune responses. Much like social immunity, host defences against brood parasitism are employed by a donor (a parent) for the benefit of one or more recipients (typically kin), and as with social defences against classic parasites, defences have therefore evolved to protect the donor's inclusive fitness, not the survival or ultimately the fitness of individual recipients This can lead to severe conflicts between the different parties, whose interests are not always aligned. Here, we consider defences against brood parasitism in the light of social immunity, at different stages of parasite encounter, addressing where conflicts occur and how they might be resolved. We finish with considering how this approach could help us to address longstanding questions in our understanding of brood parasitism. This article is part of the theme issue 'The coevolutionary biology of brood parasitism: from mechanism to pattern'.

Defensive endosymbionts are now understood to be widespread among insects, targeting many different threats, including predators, parasites and disease. The effects on natural enemies can be significant, resulting in dramatic changes in the outcome of interactions between insects and their attackers. Evidence is now emerging from laboratory and field work that defensive symbionts can have important effects on the surrounding insect community, as well as on vulnerable enemy species; for example, by reducing prey available for the trophic level above the enemy. However, there is a need for more experimental work across a greater taxonomic range of species in order to understand the different ways in which defensive symbionts influence insect communities., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Insects commonly possess heritable microbial symbionts that increase their resistance to particular parasites. A diverse community of defensive symbionts may thus provide hosts with effective and specific protection against multiple parasites, although costs might constrain the accumulation of many symbionts. In parallel to the allelic diversity in the MHC complex of the vertebrate immune system, parasite diversity could be the driving force behind symbiont diversity. There is indeed evidence that parasites have the ability to drive frequencies of defensive symbionts in their hosts, and that these symbionts influence parasite communities, but direct evidence that parasite diversity can promote symbiont diversity is still lacking. We provide suggestions to investigate this potential link., (Copyright © 2018 Elsevier Inc. All rights reserved.)

The main modes of action of insect parasitoids are considered to be killing their hosts with egg laying followed by offspring development (reproductive mortality), and adults feeding on hosts directly (host feeding). However, parasitoids can also negatively affect their hosts in ways that do not contribute to current or future parasitoid reproduction (nonreproductive effects). Outcomes of nonreproductive effects for hosts can include death, altered behavior, altered reproduction, and altered development. On the basis of these outcomes and the variety of associated mechanisms, we categorize nonreproductive effects into ( a) nonconsumptive effects, ( b) mutilation, ( c) pseudoparasitism, ( d) immune defense costs, and ( e) aborted parasitism. These effects are widespread and can cause greater impacts on host populations than successful parasitism or host feeding. Nonreproductive effects constitute a hidden dimension of host-parasitoid trophic networks, with theoretical implications for community ecology as well as applied importance for the evaluation of ecosystem services provided by parasitoid biological control agents.

Immunology textbooks teach us about the ways hosts can recognize and kill microbes but leave out something important: the mechanisms used to survive infections. Survival depends on more than simply detecting and eliminating microbes; it requires that we prevent and repair the damage caused by pathogens and the immune response. Recent work in insects is helping to build our understanding of this aspect of pathology, called disease tolerance. Here we discuss papers that explore disease tolerance using theoretical, population genetics, and mechanistic approaches., (Copyright © 2018. Published by Elsevier Inc.)

Insect societies face many social parasites that exploit their altruistic behaviours or their resources. Due to the fitness costs these social parasites incur, hosts have evolved various behavioural, chemical, architectural and morphological defence traits. Similar to bacteria infecting multicellular hosts, social parasites have to successfully go through several steps to exploit their hosts. Here, we review how social insects try to interrupt this sequence of events. They can avoid parasite contact by choosing to nest in parasite-free locales or evade attacks by adapting their colony structure. Once social parasites attack, hosts attempt to detect them, which can be facilitated by adjustments in colony odour. If social parasites enter the nest, hosts can either aggressively defend their colony or take their young and flee. Nest structures are often shaped to prevent social parasite invasion or to safeguard host resources. Finally, if social parasites successfully establish themselves in host nests, hosts can rebel by killing the parasite brood or by reproducing in the parasites' presence. Hosts of social parasites can therefore develop multiple traits, leading to the evolution of complex defence portfolios of co-dependent traits. Social parasites can respond to these multi-level defences with counter-adaptations, potentially leading to geographical mosaics of coevolution.This article is part of the Theo Murphy meeting issue 'Evolution of pathogen and parasite avoidance behaviours'., (© 2018 The Author(s).)