The innate immune system in insects is composed of a large variety of specific and non-specific responses that are activated in response to the presence of foreign agents. One important element in such responses is the enzyme phenoloxidase (PO). Here, we review recent progress in PO research and discuss new applications in the emerging field of ecological immunology. Phenoloxidase produces indole groups, which are subsequently polymerized to melanin. The enzymatic reactions in turn produce a set of intermediate products such as quinones, diphenols, superoxide, hydrogen peroxide, and reactive nitrogen intermediates, which are important during defense against bacterial (gram+ and )), fungal, and viral agents. Phenoloxidase requires a complex system of activation and inhibition that involves various cell types, PO zymogens, inhibitor enzymes, and signaling molecules. Finally, research in evolutionary ecology has studied the costs of PO in terms of resource use and pleiotropic relations with other key traits and functions. These studies indicate that PO is a costly trait, whose production and maintenance have fitness costs for hosts. Phenoloxidase does not seem to be an indicator of resistance but rather of host condition. Finally, we put forward some basic directions for future investigation of PO aimed at explaining its activating system, its substrates, its coordination with other immune components to fight off pathogens, and variation in PO in relation to gender, life stages, seasonality, and across different host species.

Most insects have a complex life cycle with ecologically different larval and adult stages. We present an ontogenetic perspective to analyze and summarize the complex life cycle of Odonata within an evolutionary ecology framework. Morphological, physiological, and behavioral pathways that generate carry-over effects across the aquatic egg and larval stages and the terrestrial adult stage are identified. We also highlight several mechanisms that can decouple life stages including compensatory mechanisms at the larval and adult stages, stressful and stochastic events during metamorphosis, and stressful environmental conditions at the adult stage that may overrule effects of environmental conditions in the preceding stage. We consider the implications of these findings for the evolution, selection, and fitness of odonates; underline the role of the identified numerical and carry-over effects in shaping population and metapopulation dynamics and the community structure across habitat boundaries; and discuss implications for applied conservation issues.

Surviving changing climate conditions is particularly difficult for organisms such as insects that depend on environmental temperature to regulate their physiological functions. Insects are extremely threatened by global warming, since many do not have enough physiological tolerance even to survive continuous exposure to the current maximum temperatures experienced in their habitats. Here, we review literature on the physiological mechanisms that regulate responses to heat and provide heat tolerance in insects: (i) neuronal mechanisms to detect and respond to heat; (ii) metabolic responses to heat; (iii) thermoregulation; (iv) stress responses to tolerate heat; and (v) hormones that coordinate developmental and behavioural responses at warm temperatures. Our review shows that, apart from the stress response mediated by heat shock proteins, the physiological mechanisms of heat tolerance in insects remain poorly studied. Based on life-history theory, we discuss the costs of heat tolerance and the potential evolutionary mechanisms driving insect adaptations to high temperatures. Some insects may deal with ongoing global warming by the joint action of phenotypic plasticity and genetic adaptation. Plastic responses are limited and may not be by themselves enough to withstand ongoing warming trends. Although the evidence is still scarce and deserves further research in different insect taxa, genetic adaptation to high temperatures may result from rapid evolution. Finally, we emphasize the importance of incorporating physiological information for modelling species distributions and ecological interactions under global warming scenarios. This review identifies several open questions to improve our understanding of how insects respond physiologically to heat and the evolutionary and ecological consequences of those responses. Further lines of research are suggested at the species, order and class levels, with experimental and analytical approaches such as artificial selection, quantitative genetics and comparative analyses.

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An explanation for courting traits is that they convey information about the bearer's condition to conspecifics, more specifically immune ability. Here we test a series of immune-based assumptions in the territorial damselfly Hetaerina americana, whose males bear wing pigmentation patterns, which are maintained via male-male competition. H. americana males emerge and take some time to mature sexually, after which, depending on their fat reserves, may start defending territories where females arrive at for copulation. Territorial males are eventually defeated and lose their territories. This loss is a consequence of a reduction in muscular fat reserves. We tested whether: (a) territorial males had more pigmented wings, more intense melaninebased immune response (encapsulation response to a nylon filament implant) and higher fat reserves than nonterritorial males; (b) pigmentation is related to immunity and fat reserves; (c) the immune response held constant in two different episodes (3 days between each) in the same male during territorial tenure; and (d) immune response and fat reserves decreased after experimentally simulated fighting event. Our results agree with current views of immune ability and courting traits: (1) territorial males had more wing pigmentation, higher immune responses and fat reserves than non-territorial males; (2) pigmentation was also correlated with immunity and fat reserves; and (3) immune response was similarly intense in the two episodes during territorial tenure. However, this response and fat reserves were considerably lower after fighting compared to that of territorial males and non-territorial males. Our work points out a link between fat reserves and immune ability which agree with previous studies in insects. Given, however, that in this species the use of wing pigmentation via male-male competition is more likely to provide information about current fat reserves than immunity, it is suggested that immune ability is only indirectly selected and may not be the information that pigmentation would convey to conspecifics.

Male damsel£ies possess very specialized genitalia. Females mate multiply and store sperm in two sperm storage organs, the bursa copulatrix and the spermatheca. During copulation, males physically remove the sperm stored in these organs using their genitalia. I document a novel mechanism by which males gain access to the spermatheca in Calopteryx haemorrhoidalis asturica. The mechanism is based on male stimulation of the female sensory system that controls egg fertilization and laying. During copulation, the aedeagus (a male genitalic structure indirectly involved in sperm transfer) distorts the cuticular plates in the female genital tract that bear mechanoreceptive sensilla. This stimulation results in sperm ejection from the spermatheca. Aedeagus width is positively correlated with the amount of sperm ejected. I propose that males have exploited a pre-existing female sensory bias to gain access to otherwise physically unreachable sperm. These results shed light on the issue of the origin of female preferences in current models of sexual selection and on the evolution of genitalia via sexual selection. It is postulated that females might use this process as a form of post-copulatory sexual selection on the basis of males' genitalia.

Odonates (dragonflies) are well known for the ability of the males to displace sperm stored in the female's spermstorage organs during copulation. By this means, copulating males are able to increase their fertilization success. This ability has been used as an example to illustrate a conflict of interests between the sexes in which males have evolved sperm-displacement mechanisms whilst females have presumably evolved means to avoid sperm displacement. The present review has four aims: (1) to describe the copulatory mechanisms used during sperm displacement; (2) to analyse the causes of sperm usage patterns; (3) to discuss this information using current hypotheses on conflict between the sexes; (4) to illuminate topics for further research. Four copulatory mechanisms are described: sperm removal (physical withdrawal of stored sperm), sperm repositioning ('pushing' of rival sperm to sites where its use will be least likely), female sensory stimulation to induce sperm ejection, and sperm flushing (displacement of sperm using the copulating male's sperm). Sperm-precedence studies in Odonata are scarce and their values vary considerably between species. In those species in which sperm displacement is incomplete, the last copulating male obtains a high but variable short-term fertilization success which decreases with time. Some male and female factors affecting sperm precedence patterns are mentioned: (1) male variation in genital morphology; (2) duration of copulation influenced by the male (the longer the copulation, the more stored sperm displaced); (3) adaptations of the sperm-storage organs that allow the female to manipulate the sperm she has received (i.e. avoiding sperm displacement, re-distributing sperm masses, favouring sperm located in certain sites and ejecting sperm after copulation). We suggest that male and female odonates have co-evolved at the level of genital function with the control of stored sperm as the focus of the conflict. The benefits for males in this co-evolution lie in maximizing their fertilization success. However, it is not clear what females obtain from storing sperm and making it unreachable during sperm displacement. Two hypothetical benefits that females may obtain for which some evidence has been gathered are genetic diversity and viability genes. It is finally suggested that odonates can become excellent subjects of study for testing current ideas related to sexual conflict and speciation processes through sexual selection.

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