A novel measure for long-term sediment reduction inspired by dragonfly wings.

The sediment accumulation in drainage pipes has long been recognized as a significant concern in the environmental field. This study addresses sediment accumulation in drainage pipes by introducing an innovative bioinspired approach using various shapes and angles of plates for long-term sediment reduction. Through experiments and numerical simulations, the velocity field, the turbulent kinetic energy, the head loss, and the dynamic pressure distribution in the pipeline with plates are analyzed. Results demonstrate significant increases in local velocity, dynamic pressure, and turbulence energy due to the presence of plates. The sediment reduction performance shows a positive correlation with the angle for folded plates and a non-linear relation with curvature for curved plates. Notably, the superior performance of folded plates is attributed to their exceptional ability to induce vortex formation. The head loss due to sediment reduction measures increases linearly as the angle and the curvature increase. Furthermore, the intentional induction of strong eddies and high shear flow using the undulating topography created by the locally installed folding plates in the pipeline was the main cause of sediment reduction. This novel approach holds promise for more efficient and sustainable sediment reduction in drainage systems.

Semi-field evaluation of aquatic predators for the control of Anopheles funestus in rural south-eastern Tanzania.

BACKGROUND: Biological control is a promising alternative or complementary approach for controlling vector populations in response to the spread of insecticide resistance in malaria vectors. This study evaluated the efficacy of three selected potential predators on the density and fitness parameters of Anopheles funestus larvae in rural Tanzania. METHODS: Common predator families Aeshnidae (dragonflies), Coenagrionidae (damselflies), and Notonectidae (backswimmers) and An. funestus group larvae were collected from natural aquatic habitats in rural south-eastern Tanzania. Predators were starved for 12-h while An. funestus larvae were given fish food before starting the experiment. Anopheles funestus larvae were placed into artificial habitats containing predators, exposing them to potential predation. The number of surviving An. funestus larvae were counted every 24-h. An emergence traps were placed at the top of artificial habitats to capture emerging mosquitoes. Emerged mosquitoes were monitored until they died. Female wings were measured and used as a proxy for body size. Generalized linear mixed models (GLMM) with binomial variates at 95% CI and Cox proportional hazard models were used to assess the proportion of dead mosquitoes and the daily survival determined. RESULTS: There were significant differences in the number of emerged mosquitoes between the treatment and control groups (P < 0.001). Thus, all predator species played a significant role in reducing the density of An. funestus mosquitoes (P < 0.001). Furthermore, these predators had notable effects on the fitness parameters and survival of emerged mosquitoes (P < 0.001). Among the three predators studied, Coenagrionidae (damselflies) were most efficient followed by Notonectidae (backswimmers), with Aeshnidae (dragonflies) being the least efficient. CONCLUSION: Selected aquatic predators have the potential to reduce the survival and density of An. funestus larvae. They might eventually be included within an integrated malaria vector control strategy, ultimately leading to a reduction in malaria transmission.

Bimodal response strategy in Daphnia to ambush predation risk.

Predation's consequences can manifest through either consumptive or nonconsumptive effects, but the prey response may also vary depending on the predator hunting strategy. Considerable attention has been paid to coursing predators, whereas less information is available regarding responses to ambush predators. To remedy this paucity, we utilized a three-dimensional tracking platform to record groups of Daphnia magna under predation risk from the ambush invertebrate predator red-eyed damselfly, Erythromma najas. This design allowed us to test individual antipredator responses in multiple metrics of swimming behaviors. We demonstrate that predation risk was greatest for those that swam at 85% of the available depth and averaged 8.1 mm/s. Examining the swimming behavior of each individual separately showed that predation risk did not affect any of the prey response metrics. Interestingly, however, Daphnia did conform to one of two strategies while under predation risk: either swim fast high up in the water column or swim slowly close to the bottom. Hence, this dichotomous behavior is driven by strategies combining speed and depth in different constellations. In a broader context, our findings highlight the importance of considering both the spatial and temporal dimensions of predation events in order to correctly detect antipredator responses.

Flow sensing on dragonfly wings.

One feature of animal wings is their embedded mechanosensory system that can support flight control. Insect wings are particularly interesting as they are highly deformable yet the actuation is limited to the wing base. It is established that strain sensors on insect wings can directly mediate reflexive control; however, little is known about airflow sensing by insect wings. What information can flow sensors capture and how can flow sensing benefit flight control? Here, we use the dragonfly (Sympetrum striolatum) as a model to explore the function of wing sensory bristles in the context of flight control. Combining our detailed anatomical reconstructions of both the sensor microstructures and wing architecture, we used computational fluid dynamics simulations to ask the following questions. (1) Are there strategic locations on wings that sample flow for estimating aerodynamically relevant parameters such as the local effective angle of attack? (2) Is the sensory bristle distribution on dragonfly wings optimal for flow sensing? (3) What is the aerodynamic effect of microstructures found near the sensory bristles on dragonfly wings? We discuss the benefits of flow sensing for flexible wings and how the evolved sensor placement affects information encoding.

Human-modified habitats imperil ornamented dragonflies less than their non-ornamented counterparts at local, regional, and continental scales.

Biologists have long wondered how sexual ornamentation influences a species' risk of extinction. Because the evolution of condition-dependent ornamentation can reduce intersexual conflict and accelerate the fixation of advantageous alleles, some theory predicts that ornamented taxa can be buffered against extinction in novel and/or stressful environments. Nevertheless, evidence from the wild remains limited. Here, we show that ornamented dragonflies are less vulnerable to extinction across multiple spatial scales. Population-occupancy models across the Western United States reveal that ornamented species have become more common relative to non-ornamented species over >100 years. Phylogenetic analyses indicate that ornamented species exhibit lower continent-wide extinction risk than non-ornamented species. Finally, spatial analyses of local dragonfly assemblages suggest that ornamented species possess advantages over non-ornamented taxa at living in habitats that have been converted to farms and cities. Together, these findings suggest that ornamented taxa are buffered against contemporary extinction at local, regional, and continental scales.

A review of applications and limitations of using aquatic macroinvertebrate predators for biocontrol of the African malaria mosquito, Anopheles gambiae sensu lato.

Macroinvertebrate predators such as backswimmers (Heteroptera: Notonectidae), dragonflies (Odonata: Aeshnidae), and predatory diving beetles (Coleoptera: Dytiscidae) naturally inhabit aquatic ecosystems. Some aquatic ecosystems inhabited by these macroinvertebrate predator taxa equally form malaria vector larval habitats. The presence of these predators in malaria vector larval habitats can negatively impact on development, adult body size, fecundity, and longevity of the malaria vectors, which form important determinants of their fitness and future vectorial capacity. These potential negative impacts caused by aquatic macroinvertebrate predators on malaria vectors warrant their consideration as biocontrol agents in an integrated program to combat malaria. However, the use of these macroinvertebrate predators in malaria biocontrol is currently constrained by technical bottlenecks linked to their generalist predatory tendencies and often long life cycles, demanding complex rearing systems. We reviewed the literature on the use of aquatic macroinvertebrate predators for biocontrol of malaria vectors from the An. gambiae s.l. complex. The available information from laboratory and semi-field studies has shown that aquatic macroinvertebrates have the potential to consume large numbers of mosquito larvae and could thus offer an additional approaches in integrated malaria vector management strategies. The growing number of semi-field structures available in East and West Africa provides an opportunity to conduct ecological experimental studies to reconsider the potential of using aquatic macroinvertebrate predators as a biocontrol tool. To achieve a more sustainable approach to controlling malaria vector populations, additional, non-chemical interventions could provide a more sustainable approach, in comparison with the failing chemical control tools, and should be urgently considered for integration with the current mosquito vector control campaigns.

How to train your dragon: absolute conditioning in larval dragonflies.

Insects, despite possessing relatively small brains, exhibit noteworthy adaptive behaviors, making them intriguing subjects for understanding learning mechanisms. This study explores the learning capabilities of dragonfly larvae (Anisoptera: Aeshnidae) in conditioning experiments, shedding light on the cognitive processes that underpin their remarkable abilities. As apex predators, dragonflies play a crucial role in ecosystems, necessitating a diverse range of learning behaviors for survival and reproductive success. We addressed whether dragonfly larvae can differentiate between different colored stimuli and associate color with prey. Our experimental design demonstrated that dragonfly larvae are able to recognize conditioning stimuli. The findings contribute valuable insights into the cognitive abilities of dragonflies, suggesting that these insects can learn and discriminate colors of stimuli. Overall, this research broadens our understanding of insect learning and cognition, contributing to the broader field of animal behavior and memory.

On the wings of dragons: Wing morphometric differences in the sexually dichromatic common whitetail skimmer dragonfly, Plathemis lydia (Odonata: Libellulidae).

Sexual dimorphism is common throughout the animal kingdom, leading to sex-specific phenotypic differences. The common whitetail skimmer dragonfly, Plathemis lydia (Drury, 1773), is sexually dichromatic, where males of this species display a conspicuous white abdomen and females display a dark brown abdomen. Differences in abdomen conspicuousness between male and female P. lydia are likely attributed to differences in selective pressure where males use their white conspicuous abdomen during male-male territorial chases. We hypothesized that male P. lydia would exhibit wing morphology adaptations to better offset the costs of predation and territoriality and that these adaptations would differ from females. We used field-collected images to quantify differences in body length, wing length, wing area, wing shape, and wing loading between male and female P. lydia. Our results show that male P. lydia have significantly shorter fore and hind wings relative to body size with a higher wing loading when compared to females. We also found that male P. lydia have narrower and pointier fore and hind wings compared to females. These results are consistent with the idea that males are adapted for faster flight, specifically higher acceleration capacity, and higher agility whereas females are adapted for higher maneuverability.

Research of biomimetic corrugation on the blade flutter suppression in large-scale wind turbine systems.

Aiming at the blade flutter of large horizontal-axis wind turbines, a method by utilizing biomimetic corrugation to suppress blade flutter is first proposed. By extracting the dragonfly wing corrugation, the biomimetic corrugation airfoil is constructed, finding that mapping corrugation to the airfoil pressure side has better aerodynamic performance. The influence of corrugation type, amplitudeλ, and intensity on airfoil flutter is analyzed using orthogonal experiment, which determines that theλhas the greatest influence on airfoil flutter. Based on the fluctuation range of the moment coefficient ΔCm, the optimal airfoil flutter suppression effect is obtained when the type is III,λ= 0.6, and intensity is denser (n= 13). The effective corrugation layout area in the chord direction is determined to be the leading edge, and the ΔCmof corrugation airfoil is reduced by 7.405%, compared to the original airfoil. The application of this corrugation to NREL 15 MW wind turbine 3D blades is studied, and the influence of corrugation layout length in the blade span direction on the suppressive effect is analyzed by fluid-structure interaction. It is found that when the layout length is 0.85 R, the safety marginSfreaches a maximum value of 0.3431 Hz, which is increased 2.940%. The results show that the biomimetic corrugated structure proposed in this paper can not only improve the aerodynamic performance by changing the local flow field on the surface of the blade, but also increase the structural stiffness of the blade itself, and achieve the effect of flutter suppression.

Sum or mean in calculation of qualitative scoring methods using the Dragonfly Biotic Index, and an alternative approach facilitating conservation prioritization.

Qualitative scoring methods are tools for rapid freshwater health assessments. Total score is often calculated as the sum or mean of the values of the species involved, with minor nuances in interpretation, but with significant implications. We used the Dragonfly Biotic Index (DBI) calculated on Central European odonate species to demonstrate these implications. Each species within a community has a score ranging from 0 (widespread generalists) to 9 (sensitive specialists). A total score is calculated as the sum of the scores of all species (DBIsum) or is calculated by dividing by species richness (DBImean). Despite this duality, there has been little debate on either approach. Using simulated scenarios (high vs low richness, presence or absence of high- or low-scoring species), we tested the implications of DBIsum and DBImean and suggested a total score calculation for conservation prioritization based on permutation. This algorithm shows the percentile of a community compared to a set of randomly assembled communities of the same species richness. We also present the 'dragDBI' package for the statistical software R, a tool for more automated DBI-based environmental health assessments. Our permutational calculation is applicable to other macroinvertebrate-based scoring methods, such as the Biological Monitoring Working Party and the Average Score Per Taxon.

Sub-lethal effects of the insecticide, imidacloprid, on the responses of damselfly larvae to chemosensory cues indicating predation risk.

Insecticides, including the widely used neonicotinoids, can affect both pest and non-target species. In addition to lethal effects, these insecticides at sub-lethal levels may cause disruption to sensory perception and processing leading to behavioural impairments. In this laboratory experiment, we investigated the effects of a 10-day exposure to the neonicotinoid insecticide, imidacloprid, on the behaviour of larvae of the damselfly, Lestes congener. In tests of baseline activity, imidacloprid concentrations of 1.0 and 10.0 µg/L caused significant reductions in foraging behaviour. Moreover, in response to chemical cues that indicate a potential risk to the larvae, imidacloprid caused the loss of an appropriate antipredator response (reduced foraging) depending on the concentration and duration of exposure. Imidacloprid at 0.1 µg/L caused the loss of responses toward the odour of a beetle (Dytiscus spp.) predator after 10 days of exposure, whereas 1.0 µg/L caused lost responses toward both the predator odour and injured conspecific cues (i.e., alarm cues) and after only 2 days of exposure. However, at 10.0 µg/L, larvae responded appropriately to both cues throughout the duration of the study, suggesting compensatory responses to imidacloprid at higher concentrations. Hence, the lack of appropriate responses at 1.0 µg/L likely resulted from a cognitive impairment rather than chemical alteration of these important chemosensory cues. In the natural environment, such effects will likely cause decreased survivorship in predator encounters. Hence, imidacloprid exposure, even at low concentrations, could have adverse consequences for chemosensory ecology of this damselfly species.

Microgeographic differentiation in thermal and antipredator responses and their carry-over effects across life stages in a damselfly.

Global warming and invasive species, separately or combined, can impose a large impact on the condition of native species. However, we know relatively little about how these two factors, individually and in combination, shape phenotypes in ectotherms across life stages and how this can differ between populations. We investigated the non-consumptive predator effects (NCEs) imposed by native (perch) and invasive (signal crayfish) predators experienced only during the egg stage or during both the egg and larval stages in combination with warming on adult life history traits of the damselfly Ischnura elegans. To explore microgeographic differentiation, we compared two nearby populations differing in thermal conditions and predator history. In the absence of predator cues, warming positively affected damselfly survival, possibly because the warmer temperature was closer to the optimal temperature. In the presence of predator cues, warming decreased survival, indicating a synergistic effect of these two variables on survival. In one population, predator cues from perch led to increased survival, especially under the current temperature, likely because of predator stress acclimation phenomena. While warming decreased, predator cues increased larval development time with a proportionally stronger effect of signal crayfish cues experienced during the egg stage, indicating a negative carry-over effect from egg to larva. Warming and predator cues increased mass at emergence, with the predator effect driven mainly by exposure to signal crayfish cues during the egg stage, indicating a positive carry-over effect from egg to adult. Notably, warming and predator effects were not consistent across the two studied populations, suggesting a phenotypic signal of adaptation at a microgeographic scale to thermal conditions and predator history. We also observed pronounced shifts during ontogeny from synergistic (egg and early larval stage) toward additive (late larval stage up to emergence) effects between warming and predator stress. The results point out that population- and life-stage-specific responses in life-history traits to NCEs are needed to predict fitness consequences of exposure to native and invasive predators and warming in prey at a microgeographic scale.

Warming and top-down control of stage-structured prey: Linking theory to patterns in natural systems.

Warming has broad and often nonlinear impacts on organismal physiology and traits, allowing it to impact species interactions like predation through a variety of pathways that may be difficult to predict. Predictions are commonly based on short-term experiments and models, and these studies often yield conflicting results depending on the environmental context, spatiotemporal scale, and the predator and prey species considered. Thus, the accuracy of predicted changes in interaction strength, and their importance to the broader ecosystems they take place in, remain unclear. Here, we attempted to link one such set of predictions generated using theory, modeling, and controlled experiments to patterns in the natural abundance of prey across a broad thermal gradient. To do so, we first predicted how warming would impact a stage-structured predator-prey interaction in riverine rock pools between Pantala spp. dragonfly nymph predators and Aedes atropalpus mosquito larval prey. We then described temperature variation across a set of hundreds of riverine rock pools (n = 775) and leveraged this natural gradient to look for evidence for or against our model's predictions. Our model's predictions suggested that warming should weaken predator control of mosquito larval prey by accelerating their development and shrinking the window of time during which aquatic dragonfly nymphs could consume them. This was consistent with data collected in rock pool ecosystems, where the negative effects of dragonfly nymph predators on mosquito larval abundance were weaker in warmer pools. Our findings provide additional evidence to substantiate our model-derived predictions while emphasizing the importance of assessing similar predictions using natural gradients of temperature whenever possible.

Readjusting observational grids in dragonfly field guides.

Wildlife field guide books present salient features of species, from colour and form to behaviour, and give their readers a vocabulary to express what these features look like. Such structures for observation, or observational grids, allow users to identify wildlife species through what Law and Lynch have called 'the difference that makes the difference'. In this article, we show how these grids, and the characteristics that distinguish species, change over time in response to wider concerns in the community that use and make the field guides. We use the development of Dutch field guides for dragonflies to show how the ethics of observing wildlife, the recreational value of dragonfly observation, the affordances of observational tools, and biodiversity monitoring and conservation goals all have repercussions for how dragonflies are to be identified. Ultimately, this affects not only how dragonflies are to be observed and identified, but also what is taken to be 'out there'. The article is based on a transdisciplinary cooperation between a dragonfly enthusiast with emic knowledge and access, and an STS researcher. We hope the articulation of our approach might inspire analyses of other observational practices and communities.

Egg Morphology of Two Neotropical Dragonflies: Micrathyria hesperis and Miathyria simplex (Odonata: Libellulidae).

Odonata species live through their immature development in aquatic habitats, where the eggs are adapted to survive the initial stages. The survival of the eggs is linked to the layers that make up the eggshell. These layers are common to all odonate species, but show variations among them, depending on the oviposition method. For example, endophytic eggs (laid in plant tissues) and exophytic eggs (laid on the water surface) have different morphologies, which may reveal egg adaptations to environmental conditions and stresses. The differences in the egg morphologies could be shown in the length of the egg layers, their histochemical composition, and in the ultrastructure. The histochemical composition can reveal several features that could lead us to a better understanding of its function, for example, how glycoproteins and total proteins could regulate the humidity and water retention in the eggs according to the environments in which the eggs are placed.

Relatively large wings facilitate life at higher elevations among Nearctic dragonflies.

Determining which traits allow species to live at higher elevations is essential to understanding the forces that shape montane biodiversity. For the many animals that rely on flight for locomotion, a long-standing hypothesis is that species with relatively large wings should better persist in high-elevation environments because wings that are large relative to the body generate more lift and decrease the aerobic costs of remaining aloft. Although these biomechanical and physiological predictions have received some support in birds, other flying taxa often possess smaller wings at high elevations or no wings at all. To test if predictions about the requirements for relative wing size at high elevations are generalizable beyond birds, we conducted macroecological analyses on the altitudinal characteristics of 302 Nearctic dragonfly species. Consistent with the biomechanical and aerobic hypotheses, species with relatively larger wings live at higher elevations and have wider elevation breadths-even after controlling for a species' body size, mean thermal conditions, and range size. Moreover, a species' relative wing size had nearly as large of an impact on its maximum elevation as being adapted to the cold. Relatively large wings may be essential to high-elevation life in species that completely depend on flight for locomotion, like dragonflies or birds. With climate change forcing taxa to disperse upslope, our findings further suggest that relatively large wings could be a requirement for completely volant taxa to persist in montane habitats.

A switchable joint in the head of dragonfly larvae (Insecta: Odonata) as key to the multifunctionality of the prehensile labial mask.

Dragonfly and damselfly larvae (Insecta: Odonata) capture prey by rapid protraction of a raptorial mouthpart, based on a modified labium. Yet, in insects with biting-chewing mouthparts, the labium has an essential role in food handling. These two distinct functions -prey capturing and handling-lead to a mechanical problem in Odonata larvae: while the labium is always protracted in a straight line during prey capture, food handling requires more dexterity. In this study, we investigate the role of the labium in the feeding process and analyse the mechanics of the labial joints in the dragonfly larva Anax imperator. Our results show that the labium features a multiaxial joint connecting the basal segment (postmentum) and the head. During feeding, a combination of rotations around different axes is used to handle and orient prey, which is unique among biting-chewing mouthparts. Furthermore, we identified structures at the joint which likely restrict lateral motion during the predatory strike. Our results provide a further understanding of the unique prey-capturing apparatus of odonate larvae capable of controlling a 'switchable' multiaxial to a restricted monoaxial joint. This concept highlights the evolution of a highly modified raptorial mouthpart appendage where the degrees of freedom can be actively restricted to allow for the respectively needed functionality.

Thermal and latitudinal patterns in pace-of-life traits are partly mediated by the gut microbiome.

The integration of life-history, physiological and behavioural traits into the pace-of-life generates a powerful framework to understand trait variation in nature both along environmental gradients and in response to environmental stressors. While the gut microbiome has been hypothesized as a candidate mechanism to underlie differentiation in the pace-of-life, this has been rarely studied. We investigated the role of the gut microbiome in contributing to the differentiation in pace-of-life and in thermal adaptation between populations of Ischnura elegans damselfly larvae inhabiting warmer low latitudes and colder high latitudes. We carried out a common-garden experiment, whereby we manipulated the exposure of the damselfly larvae to two key global warming factors: 4 °C warming and a 30 °C heat wave. Comparing the bacterial composition of the food source and the bacterioplankton indicated that damselfly larvae differentially take up bacteria from the surrounding environment and have a resident and functionally relevant microbiome. The gut microbiome differed between larvae of both latitudes, and this was associated with the host's latitudinal differentiation in activity, a key pace-of-life trait. Under heat wave exposure, the gut microbial community composition of high-latitude larvae converged towards that of the low-latitude larvae, with an increase in bacteria that likely are important in providing energy to cope with the heat wave. This suggests an adaptive latitude-specific shift in the gut microbiota matching the better ability of low-latitude hosts to deal with heat extremes. In general, our study provides evidence for the gut microbiome contributing to latitudinal differentiation in both the pace-of-life and in heat adaptation in natural populations.

Effects of predation risk on the sensory asymmetries and defensive strategies of Bufotes balearicus tadpoles.

Lateralization consists of the differential use of bilateral organs or limbs and is well described in many taxa and in several contexts. Common ecological frameworks where it can be observed are foraging and predatory ones, with benefits related to both visual and auditory lateralization such as faster response or increasing neural processing ability. Anuran amphibians are considered relevant models for investigating lateralization, due to their great ecological variety and the possibility of easily being raised under laboratory conditions. By adopting the "rotational preference test", we used Balearic green toad tadpoles to test the effects of behavioural defensive responses triggered by different predator types (native vs alien, i.e. dragonfly larvae Aeshna cyanea and adult red swamp crayfish Procambarus clarkii) and diets (fasted vs. tadpole-fed predators) on their lateralization. We recorded tadpoles' responses to five different chemical cues: clean water (control treatment), fasted dragonfly larvae and crayfish, and tadpole-fed dragonfly larvae and crayfish. Green toad tadpoles did not show a bias in a predominant direction, although lateralization occurred at the individual level, as shown by the intensity index (LA). Perceived predation risk was the highest in tadpoles exposed to the combined chemical cues of conspecific prey and native predators, which elicited both changes in the intensity of lateralization and a marked reduction in tadpoles' activity level. Our results suggest that contextual predation threat may induce very rapid changes in the expression of asymmetries at the individual level, and might play a role as part of the complex defensive strategies adopted by prey in the attempt to escape predators.

Warming and predation risk only weakly shape size-mediated priority effects in a cannibalistic damselfly.

Differences in hatching dates can shape intraspecific interactions through size-mediated priority effects (SMPE), a phenomenon where bigger, early hatched individuals gain advantage over smaller, late hatched ones. However, it remains unclear to what extent and how SMPE are affected by key environmental factors such as warming and predation risk imposed by top predators. We studied effects of warming (low and high temperature) and predation risk (presence and absence of predator cues of perch) on SMPE in life history and physiological traits in the cannibalistic damselfly Ischnura elegans. We induced SMPE in the laboratory by manipulating hatching dates, creating following groups: early and late hatchlings reared in separate containers, and mixed phenology groups where early and late hatchlings shared the same containers. We found strong SMPE for survival and emergence success, with the highest values in early larvae of mixed phenology groups and the lowest values in late larvae of mixed phenology groups. Neither temperature nor predator cues affected SMPE for these two traits. The other life history traits (development rate and mass at emergence) did not show SMPE, but were affected by temperature and predator cues. A tendency for SMPE was found for protein content, in the high temperature treatment. The other physiological traits (phenoloxidase activity and fat content) showed fixed expressions across treatments, indicating decoupling between physiology and life history. The results underline that SMPEs are trait-dependent, and only weakly or not affected by temperature and predation risk.