Scientific Publications

Andres, J.A., Cordero, R.A., 2000. Copulation duration and fertilization success in a damselfly: an example of cryptic female choice? Anim Behav. 59, 695-703.
Abstract: Copulation duration is highly variable (0.5-3 h) in the damselfly, Ceriagrion tenellum (Coenagrionidae). Using laboratory experiments, we tested four adaptive hypotheses to explain this variation: the effect of time constraints, in-copula mate guarding, sperm displacement and cryptic female choice. Copulation duration was negatively correlated with time of day, as predicted by the first two hypotheses, and positively correlated with male density, as predicted by the mate-guarding hypothesis. Males prolonged copulation in response to the volume of sperm stored by females, suggesting they were able to detect and quantify the amount of sperm stored. This behaviour is not explained by mate guarding or time constraint effects. Males removed all the sperm from the bursa copulatrix in just 10 min. Our results also suggest that, because the duct is too narrow to allow male genitalia to enter, males do not remove spermathecal sperm. Therefore, direct sperm removal could not explain long copulations. Prolonged copulations could also have evolved as a result of cryptic female choice if they increase male fertilization success by female-mediated processes. Our results support this idea: male fertilization success was greater after long copulations. Apparently, male copulatory behaviour elicits female responses that increase male fertilization success. Copyright 2000 The Association for the Study of Animal Behaviour

Bagheri ZM, Cazzolato BS, Grainger S, O'Carroll DC, Wiederman SD. 2017. An autonomous robot inspired by insect neurophysiology pursues moving features in natural environments. J Neural Eng 14:046030.
Abstract: OBJECTIVE: Many computer vision and robotic applications require the implementation of robust and efficient target-tracking algorithms on a moving platform. However, deployment of a real-time system is challenging, even with the computational power of modern hardware. Lightweight and low-powered flying insects, such as dragonflies, track prey or conspecifics within cluttered natural environments, illustrating an efficient biological solution to the target-tracking problem. APPROACH: We used our recent recordings from 'small target motion detector' neurons in the dragonfly brain to inspire the development of a closed-loop target detection and tracking algorithm. This model exploits facilitation, a slow build-up of response to targets which move along long, continuous trajectories, as seen in our electrophysiological data. To test performance in real-world conditions, we implemented this model on a robotic platform that uses active pursuit strategies based on insect behaviour. MAIN RESULTS: Our robot performs robustly in closed-loop pursuit of targets, despite a range of challenging conditions used in our experiments; low contrast targets, heavily cluttered environments and the presence of distracters. We show that the facilitation stage boosts responses to targets moving along continuous trajectories, improving contrast sensitivity and detection of small moving targets against textured backgrounds. Moreover, the temporal properties of facilitation play a useful role in handling vibration of the robotic platform. We also show that the adoption of feed-forward models which predict the sensory consequences of self-movement can significantly improve target detection during saccadic movements. SIGNIFICANCE: Our results provide insight into the neuronal mechanisms that underlie biological target detection and selection (from a moving platform), as well as highlight the effectiveness of our bio-inspired algorithm in an artificial visual system

Ball-Damerow, J.E., Oboyski, P.T., Resh, V.H., 2015. California dragonfly and damselfly (Odonata) database: temporal and spatial distribution of species records collected over the past century. Zookeys. 67-89.
Abstract: The recently completed Odonata database for California consists of specimen records from the major entomology collections of the state, large Odonata collections outside of the state, previous literature, historical and recent field surveys, and from enthusiast group observations. The database includes 32,025 total records and 19,000 unique records for 106 species of dragonflies and damselflies, with records spanning 1879-2013. Records have been geographically referenced using the point-radius method to assign coordinates and an uncertainty radius to specimen locations. In addition to describing techniques used in data acquisition, georeferencing, and quality control, we present assessments of the temporal, spatial, and taxonomic distribution of records. We use this information to identify biases in the data, and to determine changes in species prevalence, latitudinal ranges, and elevation ranges when comparing records before 1976 and after 1979. The average latitude of where records occurred increased by 78 km over these time periods. While average elevation did not change significantly, the average minimum elevation across species declined by 108 m. Odonata distribution may be generally shifting northwards as temperature warms and to lower minimum elevations in response to increased summer water availability in low-elevation agricultural regions. The unexpected decline in elevation may also be partially the result of bias in recent collections towards centers of human population, which tend to occur at lower elevations. This study emphasizes the need to address temporal, spatial, and taxonomic biases in museum and observational records in order to produce reliable conclusions from such data

Barnard AA, Fincke OM, McPeek MA, Masly JP. 2017. Mechanical and tactile incompatibilities cause reproductive isolation between two young damselfly species. Evolution.
Abstract: External male reproductive structures have received considerable attention as a cause of reproductive isolation (RI), because the morphology of these structures often evolves rapidly between populations. This rapid evolution presents the potential for mechanical incompatibilities with heterospecific female structures during mating and could thus prevent interbreeding between nascent species. Although such mechanical incompatibilities have received little empirical support as a common cause of RI, the potential for mismatch of reproductive structures to cause RI due to incompatible species-specific tactile cues has not been tested. We tested the importance of mechanical and tactile incompatibilities in RI between Enallagma anna and E. carunculatum, two damselfly species that diverged within the past approximately 250,000 years and currently hybridize in a sympatric region. We quantified 19 prezygotic and postzygotic RI barriers using both naturally occurring and laboratory-reared damselflies. We found incomplete mechanical isolation between the two pure species and between hybrid males and pure species females. Interestingly, in mating pairs for which mechanical isolation was incomplete, females showed greater resistance and refusal to mate with hybrid or heterospecific males compared to conspecific males. This observation suggests that tactile incompatibilities involving male reproductive structures can influence female mating decisions and form a strong barrier to gene flow in early stages of speciation

Berry, R., van, K.J., Stange, G., 2007. The mapping of visual space by dragonfly lateral ocelli. J. Comp Physiol A Neuroethol. Sens. Neural Behav. Physiol 193, 495-513.
Abstract: We study the extent to which the lateral ocelli of dragonflies are able to resolve and map spatial information, following the recent finding that the median ocellus is adapted for spatial resolution around the horizon. Physiological optics are investigated by the hanging-drop technique and related to morphology as determined by sectioning and three-dimensional reconstruction. L-neuron morphology and physiology are investigated by intracellular electrophysiology, white noise analysis and iontophoretic dye injection. The lateral ocellar lens consists of a strongly curved outer surface, and two distinct inner surfaces that separate the retina into dorsal and ventral components. The focal plane lies within the dorsal retina but proximal to the ventral retina. Three identified L-neurons innervate the dorsal retina and extend the one-dimensional mapping arrangement of median ocellar L-neurons, with fields of view that are directed at the horizon. One further L-neuron innervates the ventral retina and is adapted for wide-field intensity summation. In both median and lateral ocelli, a distinct subclass of descending L-neuron carries multi-sensory information via graded and regenerative potentials. Dragonfly ocelli are adapted for high sensitivity as well as a modicum of resolution, especially in elevation, suggesting a role for attitude stabilisation by localization of the horizon

Berry, R.P., Stange, G., Warrant, E.J., 2007. Form vision in the insect dorsal ocelli: an anatomical and optical analysis of the dragonfly median ocellus. Vision Res. 47, 1394-1409.
Abstract: Previous work has suggested that dragonfly ocelli are specifically adapted to resolve horizontally extended features of the world, such as the horizon. We investigate the optical and anatomical properties of the median ocellus of Hemicordulia tau and Aeshna mixta to determine the extent to which the findings support this conclusion. Dragonfly median ocelli are shown to possess a number of remarkable properties: astigmatism arising from the elliptical shape of the lens is cancelled by the bilobed shape of the inner lens surface, interference microscopy reveals complex gradients of refractive index within the lens, the morphology of the retina results in zones of high acuity, and the eye has an exceedingly high sensitivity for a diurnal terrestrial invertebrate. It is concluded that dragonfly ocelli employ a number of simple, yet elegant, anatomical and optical strategies to ensure high sensitivity, fast transduction speed, wide fields of views and a modicum of spatial resolving power

Bick, G.H., Bick, J.C., 1965. Sperm transfer in damselflies (Odonata: Zygoptera). Ann. Entomol. Soc. Am. 58, 592.

Bode-Oke AT, Zeyghami S, Dong H. 2017. Aerodynamics and flow features of a damselfly in takeoff flight. Bioinspir Biomim 12:056006.
Abstract: Flight initiation is fundamental for survival, escape from predators and lifting payload from one place to another in biological fliers and can be broadly classified into jumping and non-jumping takeoffs. During jumping takeoffs, the legs generate most of the initial impulse. Whereas the wings generate most of the forces in non-jumping takeoffs, which are usually voluntary, slow, and stable. It is of great interest to understand how these non-jumping takeoffs occur and what strategies insects use to generate large amount of forces required for this highly demanding flight initiation mode. Here, for the first time, we report accurate wing and body kinematics measurements of a damselfly during a non-jumping takeoff. Furthermore, using a high fidelity computational fluid dynamics simulation, we identify the 3D flow features and compute the wing aerodynamics forces to unravel the key mechanisms responsible for generating large flight forces. Our numerical results show that a damselfly generates about three times its body weight during the first half-stroke for liftoff. In generating these forces, the wings flap through a steeply inclined stroke plane with respect to the horizon, slicing through the air at high angles of attack (45 degrees -50 degrees ). Consequently, a leading edge vortex (LEV) is formed during both the downstroke and upstroke on all the four wings. The formation of the LEV, however, is inhibited in the subsequent upstrokes following takeoff. Accordingly, we observe a drastic reduction in the magnitude of the aerodynamic force, signifying the importance of LEV in augmenting force production. Our analysis also shows that forewing-hindwing interaction plays a favorable role in enhancing both lift and thrust production during takeoff

Bomphrey, R.J., Nakata, T., Henningsson, P., Lin, H.T., 2016. Flight of the dragonflies and damselflies. Philos. Trans. R. Soc. Lond. B Biol. Sci. 371.
Abstract: This work is a synthesis of our current understanding of the mechanics, aerodynamics and visually mediated control of dragonfly and damselfly flight, with the addition of new experimental and computational data in several key areas. These are: the diversity of dragonfly wing morphologies, the aerodynamics of gliding flight, force generation in flapping flight, aerodynamic efficiency, comparative flight performance and pursuit strategies during predatory and territorial flights. New data are set in context by brief reviews covering anatomy at several scales, insect aerodynamics, neuromechanics and behaviour. We achieve a new perspective by means of a diverse range of techniques, including laser-line mapping of wing topographies, computational fluid dynamics simulations of finely detailed wing geometries, quantitative imaging using particle image velocimetry of on-wing and wake flow patterns, classical aerodynamic theory, photography in the field, infrared motion capture and multi-camera optical tracking of free flight trajectories in laboratory environments. Our comprehensive approach enables a novel synthesis of datasets and subfields that integrates many aspects of flight from the neurobiology of the compound eye, through the aeromechanical interface with the surrounding fluid, to flight performance under cruising and higher-energy behavioural modes.This article is part of the themed issue 'Moving in a moving medium: new perspectives on flight'

Braune, P., Rolff, J., 2001. Parasitism and survival in a damselfly: does host sex matter? Proc. Biol. Sci. 268, 1133-1137.
Abstract: We present experimental data on the survivorship of damselflies infested by parasitic water mites from a population in field cages. In addition, we show correlative laboratory data under simulated severe weather conditions. In the manipulative experiment, parasitized females' individual condition, which was measured as weight at emergence, was an important determinant of survival under field conditions. In contrast, such a relationship did not occur in males and unparasitized females. It was found in the laboratory experiment that water mites as well as weight at emergence both contributed significantly to the reduced survivorship of male and female damselflies. It was concluded that the impact of parasitism depends on environmental conditions and that host sexes differ in their responses to parasitism. This is discussed in the light of immunocompetence in invertebrates

Bush, A.A., Nipperess, D.A., Duursma, D.E., Theischinger, G., Turak, E., Hughes, L., 2014. Continental-scale assessment of risk to the Australian odonata from climate change. PLoS. One. 9, e88958.
Abstract: Climate change is expected to have substantial impacts on the composition of freshwater communities, and many species are threatened by the loss of climatically suitable habitat. In this study we identify Australian Odonata (dragonflies and damselflies) vulnerable to the effects of climate change on the basis of exposure, sensitivity and pressure to disperse in the future. We used an ensemble of species distribution models to predict the distribution of 270 (85%) species of Australian Odonata, continent-wide at the subcatchment scale, and for both current and future climates using two emissions scenarios each for 2055 and 2085. Exposure was scored according to the departure of temperature, precipitation and hydrology from current conditions. Sensitivity accounted for change in the area and suitability of projected climatic habitat, and pressure to disperse combined measurements of average habitat shifts and the loss experienced with lower dispersal rates. Streams and rivers important to future conservation efforts were identified based on the sensitivity-weighted sum of habitat suitability for the most vulnerable species. The overall extent of suitable habitat declined for 56-69% of the species modelled by 2085 depending on emissions scenario. The proportion of species at risk across all components (exposure, sensitivity, pressure to disperse) varied between 7 and 17% from 2055 to 2085 and a further 3-17% of species were also projected to be at high risk due to declines that did not require range shifts. If dispersal to Tasmania was limited, many south-eastern species are at significantly increased risk. Conservation efforts will need to focus on creating and preserving freshwater refugia as part of a broader conservation strategy that improves connectivity and promotes adaptive range shifts. The significant predicted shifts in suitable habitat could potentially exceed the dispersal capacity of Odonata and highlights the challenge faced by other freshwater species

Bybee, S., Cordoba-Aguilar, A., Duryea, M.C., Futahashi, R., Hansson, B., Lorenzo-Carballa, M.O., Schilder, R., Stoks, R., Suvorov, A., Svensson, E.I., Swaegers, J., Takahashi, Y., Watts, P.C., Wellenreuther, M., 2016. Odonata (dragonflies and damselflies) as a bridge between ecology and evolutionary genomics. Front Zool. 13, 46.
Abstract: Odonata (dragonflies and damselflies) present an unparalleled insect model to integrate evolutionary genomics with ecology for the study of insect evolution. Key features of Odonata include their ancient phylogenetic position, extensive phenotypic and ecological diversity, several unique evolutionary innovations, ease of study in the wild and usefulness as bioindicators for freshwater ecosystems worldwide. In this review, we synthesize studies on the evolution, ecology and physiology of odonates, highlighting those areas where the integration of ecology with genomics would yield significant insights into the evolutionary processes that would not be gained easily by working on other animal groups. We argue that the unique features of this group combined with their complex life cycle, flight behaviour, diversity in ecological niches and their sensitivity to anthropogenic change make odonates a promising and fruitful taxon for genomics focused research. Future areas of research that deserve increased attention are also briefly outlined

Callahan, M.S., McPeek, M.A., 2016. Multi-locus phylogeny and divergence time estimates of Enallagma damselflies (Odonata: Coenagrionidae). Mol. Phylogenet. Evol. 94, 182-195.
Abstract: Reconstructing evolutionary patterns of species and populations provides a framework for asking questions about the impacts of climate change. Here we use a multilocus dataset to estimate gene trees under maximum likelihood and Bayesian models to obtain a robust estimate of relationships for a genus of North American damselflies, Enallagma. Using a relaxed molecular clock, we estimate the divergence times for this group. Furthermore, to account for the fact that gene tree analyses can overestimate ages of population divergences, we use a multi-population coalescent model to gain a more accurate estimate of divergence times. We also infer diversification rates using a method that allows for variation in diversification rate through time and among lineages. Our results reveal a complex evolutionary history of Enallagma, in which divergence events both predate and occur during Pleistocene climate fluctuations. There is also evidence of diversification rate heterogeneity across the tree. These divergence time estimates provide a foundation for addressing the relative significance of historical climatic events in the diversification of this genus

Chauhan, P., Wellenreuther, M., Hansson, B., 2016. Transcriptome profiling in the damselfly Ischnura elegans identifies genes with sex-biased expression. BMC. Genomics. 17, 985.
Abstract: BACKGROUND: Sexual dimorphism occurs widely across the animal kingdom and has profound effects on evolutionary trajectories. Here, we investigate sex-specific gene expression in Ischnura elegans (Odonata: dragonflies and damselflies), a species with pronounced sexual differences including a female-limited colour polymorphism with two female-like gynochrome morphs and one male-mimicking, androchrome morph. Whole-organism transcriptome profiling and sex-biased gene expression analysis was conducted on adults of both sexes (pooling all females as well as separating the three morphs) to gain insights into genes and pathways potentially associated with sexual development and sexual conflict. RESULTS: The de novo transcriptome assembly was of high quality and completeness (54 k transcripts; 99.6% CEGMA score; 55% annotated). We identified transcripts of several relevant pathways, including transcripts involved in sex determination, hormone biosynthesis, pigmentation and innate immune signalling. A total of 1,683 genes were differentially expressed (DE) between males and all females (1,173 were female-biased; 510 male-biased). The DE genes were associated with sex-specific physiological and reproductive processes, olfaction, pigmentation (ommochrome and melanin), hormone (ecdysone) biosynthesis and innate immunity signalling pathways. Comparisons between males and each female morph category showed that the gynochromes differed more from males than the androchrome morph. CONCLUSIONS: This is the first study to characterize sex-biased gene expression in odonates, one of the most ancient extant insect orders. Comparison between I. elegans sexes revealed expression differences in several genes related to sexual differences in behaviour and development as well as morphology. The differential expression of several olfactory genes suggests interesting sexual components in the detection of odours, pheromones and environmental volatiles. Up-regulation of pigmentation pathways in females indicates a prominent role of ommochrome pigments in the formation of the genetically controlled female colour polymorphism. Finally, the female-biased expression of several immunity genes suggests a stronger immune response in females, possibly related to the high levels of male mating harassment and recurrent matings in this species, both of which have been shown to injure females and expose them to sexually transmitted diseases and toxins contained in seminal fluids

Combes, S.A., Crall, J.D., Mukherjee, S., 2010. Dynamics of animal movement in an ecological context: dragonfly wing damage reduces flight performance and predation success. Biol. Lett. 6, 426-429.
Abstract: Much of our understanding of the control and dynamics of animal movement derives from controlled laboratory experiments. While many aspects of animal movement can be probed only in these settings, a more complete understanding of animal locomotion may be gained by linking experiments on relatively simple motions in the laboratory to studies of more complex behaviours in natural settings. To demonstrate the utility of this approach, we examined the effects of wing damage on dragonfly flight performance in both a laboratory drop-escape response and the more natural context of aerial predation. The laboratory experiment shows that hindwing area loss reduces vertical acceleration and average flight velocity, and the predation experiment demonstrates that this type of wing damage results in a significant decline in capture success. Taken together, these results suggest that wing damage may take a serious toll on wild dragonflies, potentially reducing both reproductive success and survival

Combes, S.A., Salcedo, M.K., Pandit, M.M., Iwasaki, J.M., 2013. Capture success and efficiency of dragonflies pursuing different types of prey. Integr. Comp Biol. 53, 787-798.
Abstract: The dynamics of predator-prey interactions vary enormously, due both to the heterogeneity of natural environments and to wide variability in the sensorimotor systems of predator and prey. In addition, most predators pursue a range of different types of prey, and most organisms are preyed upon by a variety of predators. We do not yet know whether predators employ a general kinematic and behavioral strategy, or whether they tailor their pursuits to each type of prey; nor do we know how widely prey differ in their survival strategies and sensorimotor capabilities. To gain insight into these questions, we compared aerial predation in 4 species of libelluid dragonflies pursuing 4 types of dipteran prey, spanning a range of sizes. We quantified the proportion of predation attempts that were successful (capture success), as well as the total time spent and the distance flown in pursuit of prey (capture efficiency). Our results show that dragonfly prey-capture success and efficiency both decrease with increasing size of prey, and that average prey velocity generally increases with size. However, it is not clear that the greater distances and times required for capturing larger prey are due solely to the flight performance (e.g., speed or evasiveness) of the prey, as predicted. Dragonflies initiated pursuits of large prey when they were located farther away, on average, as compared to small prey, and the total distance flown in pursuit was correlated with initial distance to the prey. The greater initial distances observed during pursuits of larger prey may arise from constraints on dragonflies' visual perception; dragonflies typically pursued prey subtending a visual angle of 1 degrees , and rarely pursued prey at visual angles greater than 3 degrees . Thus, dragonflies may be unable to perceive large prey flying very close to their perch (subtending a visual angle greater than 3-4 degrees ) as a distinct target. In comparing the performance of different dragonfly species that co-occur in the same habitat, we found significant differences that are not explained by body size, suggesting that some dragonflies may be specialized for pursuing particular types of prey. Our results underscore the importance of performing comparative studies of predator-prey interactions with freely behaving subjects in natural settings, to provide insight into how the behavior of both participants influences the dynamics of the interaction. In addition, it is clear that gaining a full understanding of predator-prey interactions requires detailed knowledge not only of locomotory mechanics and behavior, but also of the sensory capabilities and constraints of both predator and prey

Combes, S.A., 2015. Neuroscience: Dragonflies predict and plan their hunts. Nature 517, 279-280.

Cooper, I.A., Brown, J.M., Getty, T., 2016. A role for ecology in the evolution of colour variation and sexual dimorphism in Hawaiian damselflies. J. Evol. Biol. 29, 418-427.
Abstract: Variation in traits that are sexually dimorphic is usually attributed to sexual selection, in part because the influence of ecological differences between sexes can be difficult to identify. Sex-limited dimorphisms, however, provide an opportunity to test ecological selection disentangled from reproductive differences between the sexes. Here, we test the hypothesis that ecological differences play a role in the evolution of body colour variation within and between sexes in a radiation of endemic Hawaiian damselflies. We analysed 17 Megalagrion damselflies species in a phylogenetic linear regression, including three newly discovered cases of species with female-limited dimorphism. We find that rapid colour evolution during the radiation has resulted in no phylogenetic signal for most colour and habitat traits. However, a single ecological variable, exposure to solar radiation (as measured by canopy cover) significantly predicts body colour variation within sexes (female-limited dimorphism), between sexes (sexual dimorphism), and among populations and species. Surprisingly, the degree of sexual dimorphism in body colour is also positively correlated with the degree of habitat differences between sexes. Specifically, redder colouration is associated with more exposure to solar radiation, both within and between species. We discuss potential functions of the pigmentation, including antioxidant properties that would explain the association with light (specifically UV) exposure, and consider alternative mechanisms that may drive these patterns of sexual dimorphism and colour variation

Cordero-Rivera, A., 2016. Sperm removal during copulation confirmed in the oldest extant damselfly, Hemiphlebia mirabilis. PeerJ. 4, e2077.
Abstract: Postcopulatory sexual selection may favour mechanisms to reduce sperm competition, like physical sperm removal by males. To investigate the origin of sperm removal, I studied the reproductive behaviour and mechanisms of sperm competition in the only living member of the oldest damselfly family, Hemiphlebia mirabilis, one species that was considered extinct in the 1980s. This species displays scramble competition behaviour. Males search for females with short flights and both sexes exhibit a conspicuous "abdominal flicking". This behaviour is used by males during an elaborate precopulatory courtship, unique among Odonata. Females use a similar display to reject male attempts to form tandem, but eventually signal receptivity by a particular body position. Males immobilise females during courtship using their legs, which, contrarily to other damselflies, never autotomise. Copulation is short (range 4.1-18.7 min), and occurs in two sequential stages. In the first stage, males remove part of the stored sperm, and inseminate during the second stage, at the end of mating. The male genital ligula matches the size and form of female genitalia, and ends by two horns covered by back-oriented spines. The volume of sperm in females before copulation was 2.7 times larger than the volume stored in females whose copulation was interrupted at the end of stage I, indicative of a significant sperm removal. These results point out that sperm removal is an old character in the evolution of odonates, possibly dating back to the Permian

Cordero-Rivera A. 2017. Sexual conflict and the evolution of genitalia: male damselflies remove more sperm when mating with a heterospecific female. Sci Rep 7:7844.
Abstract: In Calopteryx damselflies, males remove rivals' sperm stored by the female, thereby reducing sperm competition. This behaviour may create a sexual conflict, because females could lose the sperm stored in the spermatheca, used for long-term storage. Comparative evidence suggested antagonistic coevolution between sexes, which might prompt the evolution of narrow spermathecal ducts, or longer spermathecae, hindering sperm removal. Calopteryx haemorrhoidalis and C. splendens coexist and sometimes hybridize. Therefore, here I predicted that if females coevolve with conspecific males, heterospecific males should have an advantage when interspecific matings occur because females will show less resistance to them than to conspecific males. By hand-pairing females to males of both species, I found that in intraspecific and interspecific matings, sperm was almost completely removed from the bursa (97-100%), but only partially from the spermathecae, with more spermathecal removal in interspecific (63-71%) than intraspecific matings (14-33%). This suggests that heterospecific males are more efficient in sperm removal as predicted by a sexually-antagonistic coevolutionary scenario. Furthermore, in most cases, only the left spermatheca was emptied, suggesting that the evolution of more than one spermatheca might also be a female counter-adaptation to regain control over fertilization

Cordoba-Aguilar, A., Contreras-Garduno, J., Peralta-Vazquez, H., Luna-Gonzalez, A., Campa-Cordova, A.I., Ascencio, F., 2006. Sexual comparisons in immune ability, survival and parasite intensity in two damselfly species. J. Insect Physiol 52, 861-869.
Abstract: Recent evolutionary studies have suggested that females have a more robust immune system than males. Using two damselfly species (Hetaerina americana and Argia tezpi), we tested if females produced higher immune responses (as phenoloxidase and hydrolytic enzymes), had a higher survival (using a nylon implant inserted in the abdomen and measuring survival after 24h) and fewer parasites (gregarines and water mites) than males. We also tested whether immune differences should emerge in different body areas (thorax vs. abdomen) within each sex with the prediction that only females will differ with the abdomen having a higher immune response than their thorax since the former area, for ecological and physiological reasons, may be a target zone for increased immune investment. Animals were adults of approximately the same age. In both species, females were more immunocompetent than males, but only in H. americana females were immune responses greater in the abdomen than in the thorax. However, there were no differences in survival and parasite intensity or the probability of being parasitised between the sexes in either of the two species. Thus, this study lends partial support to the principle that females are better at defending than males despite the null difference in parasitism and survival

Cordoba-Aguilar, A., Munguia-Steyer, R., 2013. The sicker sex: understanding male biases in parasitic infection, resource allocation and fitness. PLoS. One. 8, e76246.
Abstract: The "sicker sex" idea summarizes our knowledge of sex biases in parasite burden and immune ability whereby males fare worse than females. The theoretical basis of this is that because males invest more on mating effort than females, the former pay the costs by having a weaker immune system and thus being more susceptible to parasites. Females, conversely, have a greater parental investment. Here we tested the following: a) whether both sexes differ in their ability to defend against parasites using a natural host-parasite system; b) the differences in resource allocation conflict between mating effort and parental investment traits between sexes; and, c) effect of parasitism on survival for both sexes. We used a number of insect damselfly species as study subjects. For (a), we quantified gregarine and mite parasites, and experimentally manipulated gregarine levels in both sexes during adult ontogeny. For (b), first, we manipulated food during adult ontogeny and recorded thoracic fat gain (a proxy of mating effort) and abdominal weight (a proxy of parental investment) in both sexes. Secondly for (b), we manipulated food and gregarine levels in both sexes when adults were about to become sexually mature, and recorded gregarine number. For (c), we infected male and female adults of different ages and measured their survival. Males consistently showed more parasites than females apparently due to an increased resource allocation to fat production in males. Conversely, females invested more on abdominal weight. These differences were independent of how much food/infecting parasites were provided. The cost of this was that males had more parasites and reduced survival than females. Our results provide a resource allocation mechanism for understanding sexual differences in parasite defense as well as survival consequences for each sex

Corser, J.D., White, E.L., Schlesinger, M.D., 2015. Adult activity and temperature preference drives region-wide damselfly (Zygoptera) distributions under a warming climate. Biol. Lett. 11, 20150001.

Abstract: We analysed a recently completed statewide odonate Atlas using multivariate linear models. Within a phylogenetically explicit framework, we developed a suite of data-derived traits to assess the mechanistic distributional drivers of 59 species of damselflies in New York State (NYS). We found that length of the flight season (adult breeding activity period) mediated by thermal preference drives regional distributions at broad (10(5) km(2)) scales. Species that had longer adult flight periods, in conjunction with longer growing seasons, had significantly wider distributions. These intrinsic traits shape species' responses to changing climates and the mechanisms behind such range shifts are fitness-based metapopulation processes that adjust phenology to the prevailing habitat and climate regime through a photoperiod filter

Culler, L.E., McPeek, M.A., Ayres, M.P., 2014. Predation risk shapes thermal physiology of a predaceous damselfly. Oecologia. 176, 653-660.
Abstract: Predation risk has strong effects on organismal physiology that can cascade to impact ecosystem structure and function. Physiological processes in general are sensitive to temperature. Thus, the temperature at which predators and prey interact may shape physiological response to predation risk. We measured and evaluated how temperature and predation risk affected growth rates of predaceous damselfly nymphs (Enallagma vesperum, Odonata: Coenagrionidae). First, we conducted growth trials at five temperatures crossed with two levels of predation risk (fish predator present versus absent) and measured growth rates, consumption rates, assimilation efficiencies, and production efficiencies of 107 individual damselflies. Second, we used a model to evaluate if and how component physiological responses to predation risk affected growth rates across temperatures. In the absence of mortality threat, growth rates of damselflies increased with warming until about 23.5 degrees C and then began to decline, a typical unimodal response to changes in temperature. Under predation risk, growth rates were lower and the shape of the thermal response was less apparent. Higher metabolic and survival costs induced by predation risk were only partially offset by changes in consumption rates and assimilation efficiencies and the magnitude of non-consumptive effects varied as a function of temperature. Furthermore, we documented that thermal physiology was mediated by predation risk, a known driver of organismal physiology that occurs in the context of species interactions. A general understanding of climatic impacts on ectothermic populations requires consideration of the community context of thermal physiology, including non-consumptive effects of predators

Daniel, K.W., Pattemore, D.E., Hagen, M., 2014. Challenges and prospects in the telemetry of insects. Biol. Rev. Camb. Philos. Soc. 89 , 511-530.
Abstract: Radio telemetry has been widely used to study the space use and movement behaviour of vertebrates, but transmitter sizes have only recently become small enough to allow tracking of insects under natural field conditions. Here, we review the available literature on insect telemetry using active (battery-powered) radio transmitters and compare this technology to harmonic radar and radio frequency identification (RFID) which use passive tags (i.e. without a battery). The first radio telemetry studies with insects were published in the late 1980s, and subsequent studies have addressed aspects of insect ecology, behaviour and evolution. Most insect telemetry studies have focused on habitat use and movement, including quantification of movement paths, home range sizes, habitat selection, and movement distances. Fewer studies have addressed foraging behaviour, activity patterns, migratory strategies, or evolutionary aspects. The majority of radio telemetry studies have been conducted outside the tropics, usually with beetles (Coleoptera) and crickets (Orthoptera), but bees (Hymenoptera), dobsonflies (Megaloptera), and dragonflies (Odonata) have also been radio-tracked. In contrast to the active transmitters used in radio telemetry, the much lower weight of harmonic radar and RFID tags allows them to be used with a broader range of insect taxa. However, the fixed detection zone of a stationary radar unit (< 1 km diameter) and the restricted detection distance of RFID tags (usually < 1-5 m) constitute major constraints of these technologies compared to radio telemetry. Most of the active transmitters in radio telemetry have been applied to insects with a body mass exceeding 1 g, but smaller species in the range 0.2-0.5 g (e.g. bumblebees and orchid bees) have now also been tracked. Current challenges of radio-tracking insects in the field are related to the constraints of a small transmitter, including short battery life (7-21 days), limited tracking range on the ground (100-500 m), and a transmitter weight that sometimes approaches the weight of a given insect (the ratio of tag mass to body mass varies from 2 to 100%). The attachment of radio transmitters may constrain insect behaviour and incur significant energetic costs, but few studies have addressed this in detail. Future radio telemetry studies should address (i) a larger number of species from different insect families and functional groups, (ii) a better coverage of tropical regions, (iii) intraspecific variability between sexes, ages, castes, and individuals, and (iv) a larger tracking range via aerial surveys with helicopters and aeroplanes equipped with external antennae. Furthermore, field and laboratory studies, including observational and experimental approaches as well as theoretical modelling, could help to clarify the behavioural and energetic consequences of transmitter attachment. Finally, the development of commercially available systems for automated tracking and potential future options of insect telemetry from space will provide exciting new avenues for quantifying movement and space use of insects from local to global spatial scales

Danko, M.J., Danko, A., Golab, M.J., Stoks, R., Sniegula, S., 2017. Latitudinal and age-specific patterns of larval mortality in the damselfly Lestes sponsa: Senescence before maturity? Exp. Gerontol. 95, 107-115.
Abstract: Latitudinal differences in life history traits driven by differences in seasonal time constraints have been widely documented. Yet, latitudinal patterns in (age-specific) mortality rates have been poorly studied. Here, we studied latitudinal differences in pre-adult age-specific mortality patterns in the strictly univoltine damselfly Lestes sponsa. We compared individuals from three latitudes reared from the egg stage in the laboratory at temperatures and photoperiods simulating those at the latitude of origin (main experiment) and under common-garden conditions at a fixed temperature and photoperiod (supplementary experiment). Results from the main experiment showed that the high-latitude population exhibited higher mortality rates than the central and southern populations, likely reflecting a cost of their faster development. Age-specific mortality patterns, also indicated higher ageing rates in the high-latitude compared to the low-latitude population, which likely had a genetic basis. The strong within-population variation in hatching dates in the low-latitude population caused variation in mortality rates; individuals that hatched later showed higher mortality rates presumably due to their shorter development times compared to larvae that hatched earlier. In both experiments, larvae from all three latitudes showed accelerated mortality rates with age, which is consistent with a pattern of senescence before adulthood.

Davidovich, H., Ribak, G., 2016. Flying with eight wings: inter-sex differences in wingbeat kinematics and aerodynamics during the copulatory flight of damselflies (Ischnura elegans). Naturwissenschaften. 103, 65.
Abstract: Copulation in the blue-tailed damselfly, Ischnura elegans, can last over 5 hours, during which the pair may fly from place to place in the so-called "wheel position". We filmed copulatory free-flight and analyzed the wingbeat kinematics of males and females in order to understand the contribution of the two sexes to this cooperative flight form. Both sexes flapped their wings but at different flapping frequencies resulting in a lack of synchronization between the flapping of the two insects. Despite their unusual body posture, females flapped their wings in a stroke-plane not significantly different to that of the males (repeated-measures ANOVA, F1,7 = 0.154, p = 0.71). However, their flapping amplitudes were smaller by 42 +/- 17 %, compared to their male mates (t test, t 7 = 9.298, p < 0.001). This was mostly due to shortening of the amplitude at the ventral stroke reversal point. Compared to solitary flight, males flying in copula increased flapping frequency by 19 %, while females decreased flapping amplitude by 27 %. These findings suggest that although both sexes contribute to copulatory flight, females reduce their effort, while males increase their aerodynamic output in order to carry both their own weight and some of the female's weight. This increased investment by the male is amplified due to male I. elegans being typically smaller than females. The need by smaller males to fly while carrying some of the weight of their larger mates may pose a constraint on the ability of mating pairs to evade predators or counter interference from competing solitary males

Dickinson, M.H., 2015. Motor control: how dragonflies catch their prey. Curr. Biol. 25, R232-R234.
Abstract: Detailed measurements of head and body motion have revealed previously unknown complexity in the predatory behavior of dragonflies. The new evidence suggests that the brains of these agile predators compute internal models of their own actions and those of their prey

Dinh, K.V., Janssens, L., Therry, L., Gyulavari, H.A., Bervoets, L., Stoks, R., 2016. Rapid evolution of increased vulnerability to an insecticide at the expansion front in a poleward-moving damselfly. Evol. Appl. 9, 450-461.
Abstract: Many species are too slow to track their poleward-moving climate niche under global warming. Pesticide exposure may contribute to this by reducing population growth and impairing flight ability. Moreover, edge populations at the moving range front may be more vulnerable to pesticides because of the rapid evolution of traits to enhance their rate of spread that shunt energy away from detoxification and repair. We exposed replicated edge and core populations of the poleward-moving damselfly Coenagrion scitulum to the pesticide esfenvalerate at low and high densities. Exposure to esfenvalerate had strong negative effects on survival, growth rate, and development time in the larval stage and negatively affected flight-related adult traits (mass at emergence, flight muscle mass, and fat content) across metamorphosis. Pesticide effects did not differ between edge and core populations, except that at the high concentration the pesticide-induced mortality was 17% stronger in edge populations. Pesticide exposure may therefore slow down the range expansion by lowering population growth rates, especially because edge populations suffered a higher mortality, and by negatively affecting dispersal ability by impairing flight-related traits. These results emphasize the need for direct conservation efforts toward leading-edge populations for facilitating future range shifts under global warming

Docile, T.N., Figueiro, R., Portela, C., Nessimian, J.L., 2016. Macroinvertebrate diversity loss in urban streams from tropical forests. Environ. Monit. Assess. 188, 237.
Abstract: The increase of human activities in recent years has significantly interfered and affected aquatic ecosystems. In this present study, we investigate the effects of urbanization in the community structure of aquatic macroinvertebrates from Atlantic Forest streams. The sampling was conducted in the mountainous region of the State of Rio de Janeiro, Brazil in 10 urban and 10 preserved streams during the dry season (August-September) of 2012. The streams were characterized for its environmental integrity conditions and physico-chemical properties of water. The macroinvertebrates were sampled on rocky substrates with a kicknet. A total of 5370 individuals were collected from all streams and were distributed among Ephemeroptera, Odonata, Plecoptera, Hemiptera, Megaloptera, Coleoptera, Trichoptera, Lepidoptera, and Diptera. In urban sites, all those orders were found, except Megaloptera, while only Mollusca was not found in preserved streams. We performed a non-metric multidimensional scaling (NMDS) analysis that separated two groups distributed among sites in urban communities and another group outside this area. The dominance was significantly higher at urban sites, while the alpha diversity and equitability were greater in preserved sites. A canonical correspondence analysis (CCA) was also performed, indicating that most taxa associated with high values of the Habitat Integrity Index (HII) and a few genus of the order Diptera with the high values of ammonia, total nitrogen, associated to streams in urban sites. Urban and preserved streams differ by physical-chemical variables and aquatic macroinvertebrates. In urban streams, there is most dominance, while alpha diversity and equitability are higher in preserved streams

Doi, H., 2008. Delayed phenological timing of dragonfly emergence in Japan over five decades. Biol. Lett. 4, 388-391.
Abstract: Recent increases in air temperature have affected species phenology, resulting in the earlier onset of spring life-cycle events. Trends in the first appearance of adult dragonflies across Japan were analysed using a dataset consisting of observations from 1953 to 2005. Dynamic factor analysis was used to evaluate underlying common trends in a set of 48 time series. The appearance of the first adult dragonfly has significantly shifted to later in the spring in the past five decades. Generalized linear mixing models suggested that this is probably the result of increased air temperatures. Increased summer and autumn temperatures may provide longer bivoltine periods and a faster growth rate; thus, the second generation, which previously hatched in summer, can emerge in the autumn causing the size of the population of dragonflies that emerge in spring to decrease. It is also possible that reduced dragonfly populations along with human development are responsible for a delay in the first observed dragonflies in the spring. However, human population density did not appear to strongly affect the appearance date. This study provides the first evidence of a delay in insect phenological events over recent decades

Dolny, A., Harabis, F., Mizicova, H., 2014. Home range, movement, and distribution patterns of the threatened dragonfly Sympetrum depressiusculum (Odonata: Libellulidae): a thousand times greater territory to protect? PLoS. One. 9, e100408.
Abstract: Dragonflies are good indicators of environmental health and biodiversity. Most studies addressing dragonfly ecology have focused on the importance of aquatic habitats, while the value of surrounding terrestrial habitats has often been overlooked. However, species associated with temporary aquatic habitats must persist in terrestrial environments for long periods. Little is known about the importance of terrestrial habitat patches for dragonflies, or about other factors that initiate or influence dispersal behaviour. The aim of this study was to reveal the relationship between population dynamics of the threatened dragonfly species Sympetrum depressiusculum at its natal site and its dispersal behaviour or routine movements within its terrestrial home range. We used a mark-release-recapture method (marking 2,881 adults) and exuviae collection with the Jolly-Seber model and generalized linear models to analyse seasonal and spatial patterns of routine movement in a heterogeneous Central European landscape. Our results show that utilisation of terrestrial habitat patches by adult dragonflies is not random and may be relatively long term (approximately 3 mo). Adult dragonflies were present only in areas with dense vegetation that provided sufficient resources; the insects were absent from active agricultural patches (p = 0.019). These findings demonstrate that even a species tightly linked to its natal site utilises an area that is several orders of magnitude larger than the natal site. Therefore, negative trends in the occurrence of various dragonfly species may be associated not only with disturbances to their aquatic habitats, but also with changes in the surrounding terrestrial landscape

Drury, J.P., Anderson, C.N., Grether, G.F., 2015. Seasonal polyphenism in wing coloration affects species recognition in rubyspot damselflies (Hetaerina spp.). J. Evol. Biol. 28, 1439-1452.
Abstract: Understanding how phenotypic plasticity evolves and in turn affects the course of evolution is a major challenge in modern biology. By definition, biological species are reproductively isolated, but many animals fail to distinguish between conspecifics and closely related heterospecifics. In some cases, phenotypic plasticity may interfere with species recognition. Here, we document a seasonal polyphenism in the degree of dark wing pigmentation in smoky rubyspot damselflies (Hetaerina titia) - a shift so pronounced that it led early researchers to classify different forms of H. titia as separate species. We further show how the seasonal colour shift impacts species recognition with the sympatric congener Hetaerina occisa. Interspecific aggression (territorial fights) and reproductive interference (mating attempts) are much more frequent early in the year, when H. titia more closely resembles H. occisa, compared to later in the year when the dark phase of H. titia predominates. Using wing colour manipulations of tethered damselflies, we show that the seasonal changes in interspecific interactions are caused not only by the seasonal colour shift but also by shifts in discriminatory behaviour in both species. We also experimentally tested and rejected the hypothesis that learning underlies the behavioural shifts in H. occisa. An alternative hypothesis, which remains to be tested, is that the seasonal polyphenism in H. titia wing coloration has resulted in the evolution of a corresponding seasonal polyphenism in species recognition in H. occisa. This study illustrates one of the many possible ways that plasticity in species recognition cues may influence the evolution of interspecific interactions

Duong, T.M., McCauley, S.J., 2016. Predation risk increases immune response in a larval dragonfly (Leucorrhinia intacta). Ecology 97, 1605-1610.
Abstract: Predators often negatively affect prey performance through indirect, non-consumptive effects. We investigated the potential relationship between predator-induced stress and prey immune response. To test this, we administered a synthetic immune challenge into dragonfly larvae (Leucorrhinia intacta) and assessed a key immune response (level of encapsulation) in the presence and absence of a caged predator (Anax junius) at two temperatures (22 degrees C and 26 degrees C). We hypothesized that immune response would be lowered when predators were present due to lowered allocation of resources to immune function and leading to reduced encapsulation of the synthetic immune challenge. Contrary to our expectations, larvae exposed to caged predators had encapsulated monofilaments significantly more than larvae not exposed to caged predators. Levels of encapsulation did not differ across temperatures, nor interact with predator exposure. Our results suggest that the previously observed increase in mortality of L. intacta exposed to caged predators is not driven by immune suppression. In situations of increased predation risk, the exposure to predator cues may induce higher levels of melanin production, which could lead to physiological damage and high energetic costs. However, the costs and risks of increased allocations to immune responses and interactions with predation stress remain unknown

Duong TM, Gomez AB, Sherratt TN. 2017. Response of adult dragonflies to artificial prey of different size and colour. PLoS One 12:e0179483.
Abstract: Aposematism is an evolved, cross-species association between a preys' unprofitability and the presence of conspicuous signals. Avian predators have been widely employed to understand the evolution of these warning signals However, insect predators are abundant, diverse, and highly visual foragers that have been shown to be capable of learned aversion. Therefore, it is likely that their behaviour also shapes the nature of anti-predator traits. In this study, we evaluated the rates of attack of a community (13 species) of mature adult dragonflies (Odonata) on artificial prey of varying size (2.5-31 mm lengthwise) and colour pattern (black, black/yellow striped). The relative attack rates of dragonflies on prey increased as prey size decreased, but there was no evidence that the attack rates by dragonflies were affected by prey colour pattern and no evidence for an interaction between colour pattern and size. To investigate prey selection by specific predator species under field conditions, we compared the time to attack distributions of black-painted prey presented to two common dragonflies: Leucorrhinia intacta and the larger, Libellula pulchella. We found that the two dragonfly species, as well as the two sexes, had different foraging responses. L. pulchella was more likely to attack larger prey, and females of both species more likely to attack prey than males. Collectively, our results indicate that dragonflies are highly size selective. However, while the nature of this selectivity varies among dragonfly species, there is little evidence that classic black/yellow warning signals deter attack by these aerial invertebrate predators

Edgehouse, M., Brown, C.P., 2014. Predatory luring behavior of odonates. J. Insect Sci. 14, 146.
Abstract: Organisms in the order Odonata are highly predatory insects that have a wide distribution globally. To date, there has been zero evidence that odonates employ luring as a means of prey acquisition. However, in this study, we show that Aeshna palmata larvae use abdominal movements to lure larval Argia vivida, subsequently consuming the lured organism. We also present findings of a similar behavior from larval Ar. vivida in an attempt to lure larval A. palmata within striking distan

Fincke, O.M., 2015. Trade-offs in female signal apparency to males offer alternative anti-harassment strategies for colour polymorphic females. J. Evol. Biol. 28, 931-943.
Abstract: Colour polymorphisms are known to influence receiver behaviour, but how they affect a receiver's ability to detect and recognize individuals in nature is usually unknown. I hypothesized that polymorphic female damselflies represent an evolutionary stable strategy, maintained by trade-offs between the relative apparency of morphs to male receivers. Using field experiments on Enallagma hageni and focal studies of E. hageni and Enallagma boreale, I tested for the first time the predictions that (i) green heteromorphs and blue andromorphs gain differential protection from sexual harassment via background crypsis and sexual mimicry, respectively, and (ii) female morphs behaviourally optimize their signal apparency to mate-searching males. First, based on male reactions elicited by females, against a high-contrast background, the two morphs did not differ in being detected by males, and once detected, they did not differ in being recognized (eliciting sexual reactions). However, on green ferns, heteromorphs were less likely to be detected (elicited only fly-bys) than andromorphs, but once detected, the morphs did not differ in being recognized. In contrast, when perched on a dowel with two male signal distractors, andromorphs were detected less often, and once detected, they were recognized less often than heteromorphs. Second, in fields where females foraged, andromorphs perched higher on vegetation than heteromorphs and were more often in the vicinity of males. Neither harassment rates nor evasive behaviours differed between morphs. Males aggregated in high density near shore where solitary females were rare. Equilibrium frequencies of these and other colour morphs should reflect the relative ease with which receivers detect and recognize them in the context where they are encountered

Fitt, R.N., Lancaster, L.T., 2017. Range shifting species reduce phylogenetic diversity in high latitude communities via competition. J. Anim Ecol. 86, 543-555.
Abstract: Under anthropogenic climate change, many species are expanding their ranges to higher latitudes and altitudes, resulting in novel species interactions. The consequences of these range shifts for native species, patterns of local biodiversity and community structure in high latitude ecosystems are largely unknown but critical to understand in light of widespread poleward expansions by many warm-adapted generalists. Using niche modelling, phylogenetic methods, and field and laboratory studies, we investigated how colonization of Scotland by a range expanding damselfly, Ischnura elegans, influences patterns of competition and niche shifts in native damselfly species, and changes in phylogenetic community structure. Colonization by I. elegans was associated with reduced population density and niche shifts in the resident species least related to I. elegans (Lestes sponsa), reflecting enhanced competition. Furthermore, communities colonized by I. elegans exhibited phylogenetic underdispersion, reflecting patterns of relatedness and competition. Our results provide a novel example of a potentially general mechanism whereby climate change-mediated range shifts can reduce phylogenetic diversity within high latitude communities, if colonizing species are typically competitively superior to members of native communities that are least-closely related to the colonizer.

Fitzstephens, D.M., Getty, T., 2000. Colour, fat and social status in male damselflies, Calopteryx maculata. Anim Behav. 60, 851-855.
Abstract: In the black-winged damselfly, Calopteryx maculata, younger males challenge and displace older males from mating territories. Fatter males tend to win fights. These fights were initially interpreted as wars of attrition based on fat reserves, but the distributions of fat at the end of fights suggests at least some assessment of the opponent's condition. Alternatively, new models have been developed that show how the observed pattern could result without assessment. We show that there is a subtle but reliable cue to fat reserves: colour. Females are a relatively drab brown-black. Males are a strikingly iridescent blue-green colour, resulting from a multilayer constructive interference reflector system in the epicuticle. In fatter males the lamellae are more compressed and the peak reflectance is at shorter wavelengths (blue). Leaner, greener males have greater spacing between lamellae and reflect longer wavelengths. The peak reflectance is as predicted from transmission electron micrograph measurements of the lamellar spacing. The rate of change in spacing over time can be manipulated experimentally by manipulating the diet. Individuals on a higher food diet remained blue longer and at the end of the experiment were fatter and bluer. In our studies, colour is a better predictor of territorial status than fat. Copyright 2000 The Association for the Study of Animal Behaviour

Frati, F., Piersanti, S., Conti, E., Rebora, M., Salerno, G., 2015. Scent of a dragonfly: sex recognition in a polymorphic Coenagrionid. PLoS. One. 10, e0136697.
Abstract: In polymorphic damselflies discrimination of females from males is complex owing to the presence of androchrome and gynochrome females. To date there is no evidence that damselflies use sensory modalities other than vision (and tactile stimuli) in mate searching and sex recognition. The results of the present behavioural and electrophysiological investigations on Ischnura elegans, a polymorphic damselfly, support our hypothesis that chemical cues could be involved in Odonata sex recognition. The bioassays demonstrate that males in laboratory prefer female to male odour, while no significant difference was present in male behavior between stimuli from males and control. The bioassays suggest also some ability of males to distinguish between the two female morphs using chemical stimuli. The ability of male antennae to perceive odours from females has been confirmed by electrophysiological recordings. These findings are important not only to get insight into the chemical ecology of Odonata, and to shed light into the problem of olfaction in Paleoptera, but could be useful to clarify the controversial aspects of the mating behavior of polymorphic coenagrionids. Behavioural studies in the field are necessary to investigate further these aspects

Frati, F., Piersanti, S., Rebora, M., Salerno, G., 2016. Volatile cues can drive the oviposition behavior in Odonata. J. Insect. Physiol 91-92, 34-38.
Abstract: Selection for the oviposition site represents the criterion for the behavioral process of habitat selection for the next generation. It is well known that in Odonata the most general cues are detected visually, but laboratory investigations on the coenagrionid Ischnura elegans showed through behavioral and electrophysiological assays that adults were attracted by olfactory cues emitted by prey and that males of the same species are attracted by female odor. The results of the present behavioral and electrophysiological investigations on I. elegans suggest the involvement of antennal olfactory sensilla in oviposition behavior. In particular, I. elegans females laid in the laboratory significantly more eggs in water from larval rearing aquaria than in distilled or tap water. Moreover, the lack of preference between rearing water and tap water with plankton suggests a role of volatiles related to conspecific and plankton presence in the oviposition site choice. I. elegans may rely on food odor for oviposition site selection, thus supporting the predictions of the "mother knows best" theory. These behavioral data are partially supported by electroantennographic responses. These findings confirm a possible role of olfaction in crucial aspects of Odonata biology

Futahashi, R., Kurita, R., Mano, H., Fukatsu, T., 2012. Redox alters yellow dragonflies into red. Proc. Natl. Acad. Sci. U. S. A 109, 12626-12631.

Abstract: Body color change associated with sexual maturation--so-called nuptial coloration--is commonly found in diverse vertebrates and invertebrates, and plays important roles for their reproductive success. In some dragonflies, whereas females and young males are yellowish in color, aged males turn vivid red upon sexual maturation. The male-specific coloration plays pivotal roles in, for example, mating and territoriality, but molecular basis of the sex-related transition in body coloration of the dragonflies has been poorly understood. Here we demonstrate that yellow/red color changes in the dragonflies are regulated by redox states of epidermal ommochrome pigments. Ratios of reduced-form pigments to oxidized-form pigments were significantly higher in red mature males than yellow females and immature males. The ommochrome pigments extracted from the dragonflies changed color according to redox conditions in vitro: from red to yellow in the presence of oxidant and from yellow to red in the presence of reductant. By injecting the reductant solution into live insects, the yellow-to-red color change was experimentally reproduced in vivo in immature males and mature females. Discontinuous yellow/red mosaicism was observed in body coloration of gynandromorphic dragonflies, suggesting a cell-autonomous regulation over the redox states of the ommochrome pigments. Our finding extends the mechanical repertoire of pigment-based body color change in animals, and highlights an impressively simple molecular mechanism that regulates an ecologically important color trait

Futahashi, R., Kawahara-Miki, R., Kinoshita, M., Yoshitake, K., Yajima, S., Arikawa, K., Fukatsu, T., 2015. Extraordinary diversity of visual opsin genes in dragonflies. Proc. Natl. Acad. Sci. U. S. A 112, E1247-E1256.
Abstract: Dragonflies are colorful and large-eyed animals strongly dependent on color vision. Here we report an extraordinary large number of opsin genes in dragonflies and their characteristic spatiotemporal expression patterns. Exhaustive transcriptomic and genomic surveys of three dragonflies of the family Libellulidae consistently identified 20 opsin genes, consisting of 4 nonvisual opsin genes and 16 visual opsin genes of 1 UV, 5 short-wavelength (SW), and 10 long-wavelength (LW) type. Comprehensive transcriptomic survey of the other dragonflies representing an additional 10 families also identified as many as 15-33 opsin genes. Molecular phylogenetic analysis revealed dynamic multiplications and losses of the opsin genes in the course of evolution. In contrast to many SW and LW genes expressed in adults, only one SW gene and several LW genes were expressed in larvae, reflecting less visual dependence and LW-skewed light conditions for their lifestyle under water. In this context, notably, the sand-burrowing or pit-dwelling species tended to lack SW gene expression in larvae. In adult visual organs: (i) many SW genes and a few LW genes were expressed in the dorsal region of compound eyes, presumably for processing SW-skewed light from the sky; (ii) a few SW genes and many LW genes were expressed in the ventral region of compound eyes, probably for perceiving terrestrial objects; and (iii) expression of a specific LW gene was associated with ocelli. Our findings suggest that the stage- and region-specific expressions of the diverse opsin genes underlie the behavior, ecology, and adaptation of dragonflies

Futahashi, R., 2016. Color vision and color formation in dragonflies. Curr. Opin. Insect. Sci. 17, 32-39.
Abstract: Dragonflies including damselflies are colorful and large-eyed insects, which show remarkable sexual dimorphism, color transition, and color polymorphism. Recent comprehensive visual transcriptomics has unveiled an extraordinary diversity of opsin genes within the lineage of dragonflies. These opsin genes are differentially expressed between aquatic larvae and terrestrial adults, as well as between dorsal and ventral regions of adult compound eyes. Recent topics of color formation in dragonflies are also outlined. Non-iridescent blue color is caused by coherent light scattering from the quasiordered nanostructures, whereas iridescent color is produced by multilayer structures. Wrinkles or wax crystals sometimes enhances multilayer structural colors. Sex-specific and stage-specific color differences in red dragonflies is attributed to redox states of ommochrome pigments

Gering, E.J., 2017. Male-mimicking females increase male-male interactions, and decrease male survival and condition in a female-polymorphic damselfly. Evolution. 71, 1390-1396.
Abstract: Biologists are still discovering diverse and powerful ways sexual conflicts shape biodiversity. The present study examines how the proportion of females in a population that exhibit male mimicry, a mating resistance trait, influences conspecific males' behavior, condition, and survival. Like most female-polymorphic damselflies, Ischnura ramburii harbors both "andromorph" females, which closely resemble males, and sexually dimorphic "gynomorph" counterparts. There is evidence that male mimicry helps andromorphs evade detection and harassment, but males can also learn to target locally prevalent morph(s) via prior mate encounters. I hypothesized that the presence of male mimics could therefore predispose males to mate recognition errors, and thereby increase rates of costly male-male interactions. Consistent with this hypothesis, male-male interaction rates were highest in mesocosms containing more andromorph (vs. gynomorph) females. Males in andromorph-biased mesocosms also had lower final body mass and higher mortality than males assigned to gynomorph-majority treatments. Male survival and body mass were each negatively affected by mesocosm density, and mortality data revealed a marginally significant interaction between andromorph frequency and population density. These findings suggest that, under sufficiently crowded conditions, female mating resistance traits such as male mimicry could have pronounced indirect effects on male behavior, condition, and survival.

Gonzalez-Tokman, D.M., Munguia-Steyer, R., Gonzalez-Santoyo, I., Baena-Diaz, F.S., Cordoba-Aguilar, A., 2012. Support for the immunocompetence handicap hypothesis in the wild: hormonal manipulation decreases survival in sick damselflies. Evolution 66, 3294-3301.
Abstract: The immunocompetence handicap hypothesis (ICHH) states that hormones enhance sexual trait expression but impair immunity. Previous tests of the ICHH have been hampered by experimental design problems. Here, we report on an experimental test of the ICHH that includes manipulations of both hormones and infections in males of the territorial damselfly, Hetaerina americana, with accurate survival measurements. We conducted a fully factorial experiment subjecting each individual to one of three topical treatments: methoprene (a juvenile hormone analog), acetone, or control, and one of three injection treatments: bacteria, PBS, or control. We measured survival of manipulated males in both the wild and in captivity. As predicted, survival was most heavily impaired in methoprene-bacteria males than in the other groups in the wild, and no survival differences emerged in captive animals. This result confirms that survival is one cost an animal pays for increased hormonal levels. This corroborates theoretical predictions of the ICHH

Guillermo-Ferreira R, Appel E, Urban P, Bispo PC, Gorb SN. 2017. The unusual tracheal system within the wing membrane of a dragonfly. Biol Lett 13.
Abstract: Some consider that the first winged insects had living tissue inside the wing membrane, resembling larval gills or developing wing pads. However, throughout the developmental process of the wing membrane of modern insects, cells and tracheoles in the lumen between dorsal and ventral cuticle disappear and both cuticles become fused. This process results in the rather thin rigid stable structure of the membrane. The herewith described remarkable case of the dragonfly Zenithoptera lanei shows that in some highly specialized wings, the membrane can still be supplemented by tracheae. Such a characteristic of the wing membrane presumably represents a strong specialization for the synthesis of melanin-filled nanolayers of the cuticle, nanospheres inside the wing membrane and complex arrangement of wax 

Haber, W.A., Wagner, D.L., Rosa, C.L., 2015. A new species of Erythrodiplax breeding in bromeliads in Costa Rica (Odonata: Libellulidae). Zootaxa. 3947, 386-396.
Abstract: We describe a new species, Erythrodiplax laselva (Libellulidae), that breeds in bromeliads and Cochliostema (Commelinaceae) in the eastern lowlands of Costa Rica. The closest known relative is thought to be E. castanea, widespread in Central and South America, and not E. bromeliicola, which is known to breed in bromeliads in Cuba and Jamaica. The male, female, genitalia, and larva are described and illustrated

Hassall, C., 2014. Continental variation in wing pigmentation in Calopteryx damselflies is related to the presence of heterospecifics. PeerJ. 2, e438.
Abstract: Wing pigmentation in Calopteryx damselflies, caused by the deposition of melanin, is energetically expensive to produce and enhances predation risk. However, patterns of melanisation are used in species identification, greater pigmentation is an accurate signal of male immune function in at least some species, and there may be a role for pigment in thermoregulation. This study tested two potential hypotheses to explain the presence of, and variation in, this pigmentation based on these three potential benefits using 907 male specimens of Calopteryx maculata collected from 49 sites (34 discrete populations) across the geographical range of the species in North America: (i) pigmentation varies with the presence of the closely related species, Calopteryx aequabilis, and (ii) pigment increases at higher latitudes as would be expected if it enhances thermoregulatory capacity. No gradual latitudinal pattern was observed, as might be expected if pigmentation was involved in thermoregulation. However, strong variation was observed between populations that were sympatric or allopatric with C. aequabilis. This variation was characterised by dark wings through allopatry in the south of the range and then a step change to much lighter wings at the southern border of sympatry. Pigmentation then increased further north into the sympatric zone, finally returning to allopatry levels at the northern range margin. These patterns are qualitatively similar to variation in pigmentation in C. aequabilis, meaning that the data are consistent with what would be expected from convergent character displacement. Overall, the results corroborate recent research that has suggested sexual selection as a primary driver behind the evolution of wing pigmentation in this group

Hassall, C., Keat, S., Thompson, D.J., Watts, P.C., 2014. Bergmann's rule is maintained during a rapid range expansion in a damselfly. Glob. Chang Biol. 20, 475-482.
Abstract: Climate-induced range shifts result in the movement of a sample of genotypes from source populations to new regions. The phenotypic consequences of those shifts depend upon the sample characteristics of the dispersive genotypes, which may act to either constrain or promote phenotypic divergence, and the degree to which plasticity influences the genotype-environment interaction. We sampled populations of the damselfly Erythromma viridulum from northern Europe to quantify the phenotypic (latitude-body size relationship based on seven morphological traits) and genetic (variation at microsatellite loci) patterns that occur during a range expansion itself. We find a weak spatial genetic structure that is indicative of high gene flow during a rapid range expansion. Despite the potentially homogenizing effect of high gene flow, however, there is extensive phenotypic variation among samples along the invasion route that manifests as a strong, positive correlation between latitude and body size consistent with Bergmann's rule. This positive correlation cannot be explained by variation in the length of larval development (voltinism). While the adaptive significance of latitudinal variation in body size remains obscure, geographical patterns in body size in odonates are apparently underpinned by phenotypic plasticity and this permits a response to one or more environmental correlates of latitude during a range expansion

Hassall, C., Sherratt, T.N., Watts, P.C., Thompson, D.J., 2015. Live fast, die old: no evidence of reproductive senescence or costs of mating in a damselfly (Odonata: Zygoptera). J. Anim Ecol.
Abstract: Recent examples of actuarial senescence in wild insect populations have challenged the long-held assumption that the brevity of wild insect life spans precludes senescence. We investigate age-related patterns in mating behaviour in adults of a short-lived damselfly, Coenagrion puella and the implications of this mating. Using capture histories for 1033 individuals over two field seasons, we conduct both pooled and stratified analyses of variations in breeding activity. Pooled analyses suggest that there is strong age-related variation in the probability of being present at the mating rendezvous. However, no age-related variation was observed in the probability of mating. Stratified approaches confirmed a general pattern of age-related declines in survival probability, but provided only equivocal evidence of an effect of age on transition between temporary breeding states. Mating males and females showed greater survival than non-mating individuals, possibly as a consequence of higher body condition. Older males that were not currently breeding were less likely to commence breeding on the next day, but showed no patterns in breeding cessation. Overall, transitions between both breeding states declined with age, suggesting that males that breed tend to continue breeding while those that do not breed continue to be unsuccessful. Female mating rates were consistently high across all ages with no age-related decline apparent. While previous research has demonstrated actuarial senescence in this population, as does this study, we find little evidence of either age-related declines in reproductive behaviour or breeding-related declines in survival, which might indicate functional senescence or costs of mating, respectively. Indeed, the greater survival in mating individuals of both sexes suggests that variations in individual quality may mediate both reproductive success and longevity. Contrary to recent studies, we found no compelling evidence for reproductive senescence or a cost of mating in an important and well-studied model odonate. The possible link between condition and ageing suggests that individual quality needs to be taken into account when studying senescence. We recommend the use of multistrata models for the future investigation of these phenomena

Hedstrom, I., Sahlen, G., 2001. A key to the adult Costa Rican "helicopter" damselflies (Odonata: Pseudostigmatidae) with notes on their phenology and life zone preferences. Rev. Biol. Trop. 49, 1037-1056.
Abstract: We present a key to the Costa Rican species of Pseudostigmatidae, comprising three genera with the following species: Megaloprepus caerulatus, Mecistogaster linearis, M. modesta, M. ornata and Pseudostigma aberrans. Pseudostigma accedens, which may occur in the region, is also included. For each species we give a brief account of morphology, phenology and life zone preferences, including distributional maps based on more than 270 records. These are not all of the known specimens from the area, but a high enough number to give a relatively good picture of the distribution and status of the species. We found M. caerulatus to be active during the first half of the year in seasonal, tropical semi-dry lowland forest and tropical moist forest at mid-elevation, but like M. linearis, M. caerulatus was active all year round in non-seasonal, tropical wet lowland forest and tropical moist forest at mid-elevation. Mecistogaster modesta also flew year round in non-seasonal, tropical wet lowland forest and tropical moist evergreen forest at mid-elevation, and likewise in seasonal and non-seasonal, tropical premontane moist forest. Only a few findings, however, have been made of M. modesta in seasonal, tropical semi-dry deciduous forest and seasonal, tropical moist evergreen forest. Mecistogaster ornata was missing entirely from non-seasonal, tropical wet lowland forest and non-seasonal, tropical moist forest at mid-elevation, while this species was active year round in seasonal, tropical dry lowland forest and tropical semi-dry forest, as well as in seasonal, tropical moist evergreen forest and tropical premontane moist forest, both at mid-elevation. Pseudostigma aberrans has so far been found too few times in Costa Rica for any indication of flight time preference

Henry, J.R., Harrison, J.F., 2014. Effects of body size on the oxygen sensitivity of dragonfly flight. J. Exp. Biol. 217,  3447-3456.
Abstract: One hypothesis for the small size of insects relative to vertebrates, and the existence of giant fossil insects, is that atmospheric oxygen levels constrain insect body sizes because oxygen delivery is more challenging in larger insects. This study tested this hypothesis in dragonflies by measuring the oxygen sensitivity of flight metabolic rates and behavior during hovering for 11 species of dragonflies that ranged in mass by an order of magnitude. We measured flight times and flight metabolic rates in seven oxygen concentrations ranging from 30% to 2.5% to assess the sensitivity of their flight to atmospheric oxygen. We also assessed the oxygen sensitivity of flight in low-density air (nitrogen replaced with helium) in order to increase the metabolic demands of hovering flight. Lowered atmospheric densities did induce higher flight metabolic rates. Flight behavior was more sensitive to decreasing oxygen levels than flight metabolic rate. The oxygen sensitivity of flight metabolic rates and behaviors were not correlated with body size, indicating that larger insects are able to maintain an oxygen supply-to-demand balance even during flight

Herzog, R., Hadrys, H., 2017. Long-term genetic monitoring of a riverine dragonfly, Orthetrum coerulescens (Odonata: Libellulidae]: Direct anthropogenic impact versus climate change effects. PLoS. One. 12, e0178014.
Abstract: Modern conservationists call for long term genetic monitoring datasets to evaluate and understand the impact of human activities on natural ecosystems and species on a global but also local scale. However, long-term monitoring datasets are still rare but in high demand to correctly identify, evaluate and respond to environmental changes. In the presented study, a population of the riverine dragonfly, Orthetrum coerulescens (Odonata: Libellulidae), was monitored over a time period from 1989 to 2013. Study site was an artificial irrigation ditch in one of the last European stone steppes and "nature heritage", the Crau in Southern France. This artificial riverine habitat has an unusual high diversity of odonate species, prominent indicators for evaluating freshwater habitats. A clearing of the canal and destruction of the bank vegetation in 1996 was assumed to have great negative impact on the odonate larval and adult populations. Two mitochondrial markers (CO1 & ND1) and a panel of nuclear microsatellite loci were used to assess the genetic diversity. Over time they revealed a dramatic decline in diversity parameters between the years 2004 and 2007, however not between 1996 and 1997. From 2007 onwards the population shows a stabilizing trend but has not reached the amount of genetic variation found at the beginning of this survey. This decline cannot be referred to the clearing of the canal or any other direct anthropogenic impact. Instead, it is most likely that the populations' decay was due to by extreme weather conditions during the specific years. A severe drought was recorded for the summer months of these years, leading to reduced water levels in the canal causing also other water parameters to change, and therefore impacting temperature sensitive riverine habitat specialists like the O. coerulescens in a significant way. The data provide important insights into population genetic dynamics and metrics not always congruent with traditional monitoring data (e.g. abundance); a fact that should be regarded with caution when management plans for developed landscapes are designed.

Hill, M.J., Sayer, C.D., Wood, P.J., 2016. When is the best time to sample aquatic macroinvertebrates in ponds for biodiversity assessment? Environ. Monit. Assess. 188, 194.
Abstract: Ponds are sites of high biodiversity and conservation value, yet there is little or no statutory monitoring of them across most of Europe. There are clear and standardised protocols for sampling aquatic macroinvertebrate communities in ponds, but the most suitable time(s) to undertake the survey(s) remains poorly specified. This paper examined the aquatic macroinvertebrate communities from 95 ponds within different land use types over three seasons (spring, summer and autumn) to determine the most appropriate time to undertake sampling to characterise biodiversity. The combined samples from all three seasons provided the most comprehensive record of the aquatic macroinvertebrate taxa recorded within ponds (alpha and gamma diversity). Samples collected during the autumn survey yielded significantly greater macroinvertebrate richness (76 % of the total diversity) than either spring or summer surveys. Macroinvertebrate diversity was greatest during autumn in meadow and agricultural ponds, but taxon richness among forest and urban ponds did not differ significantly temporally. The autumn survey provided the highest measures of richness for Coleoptera, Hemiptera and Odonata. However, richness of the aquatic insect order Trichoptera was highest in spring and lowest in autumn. The results illustrate that multiple surveys, covering more than one season, provide the most comprehensive representation of macroinvertebrate biodiversity. When sampling can only be undertaken on one occasion, the most appropriate time to undertake surveys to characterise the macroinvertebrate community biodiversity is during autumn, although this may need to be modified if other floral and faunal groups need to be incorporated into the sampling programme

Hobson, K.A., Anderson, R.C., Soto, D.X., Wassenaar, L.I., 2012. Isotopic evidence that dragonflies (Pantala flavescens) migrating through the Maldives come from the northern Indian subcontinent. PLoS. One. 7, e52594.
Abstract: Large numbers of the Globe Skimmer dragonfly (Pantala flavescens) appear in the Maldives every October-December. Since they cannot breed on these largely waterless islands, it has recently been suggested that they are "falling out" during a trans-oceanic flight from India to East Africa. In addition, it has been suggested that this trans-oceanic crossing is just one leg of a multi-generational migratory circuit covering about 14,000-18,000 km. The dragonflies are presumed to accomplish this remarkable feat by riding high-altitude winds associated with the Inter-tropical Convergence Zone (ITCZ). While there is considerable evidence for this migratory circuit, much of that evidence is circumstantial. Recent developments in the application of stable isotope analyses to track migratory dragonflies include the establishment of direct associations between dragonfly wing chitin delta(2)H values with those derived from long-term delta(2)H precipitation isoscapes. We applied this approach by measuring wing chitin delta(2)H values in 49 individual Pantala flavescens from the November-December migration through the Maldives. Using a previously established spatial calibration algorithm for dragonflies, the mean wing delta(2)H value of -117+/-16 per thousand corresponded to a predicted mean natal ambient water source of -81 per thousand, which resulted in a probabilistic origin of northern India, and possibly further north and east. This strongly suggests that the migratory circuit of this species in this region is longer than previously suspected, and could possibly involve a remarkable trans-Himalayan high-altitude traverse

Hou, D., Yin, Y., Zhao, H., Zhong, Z., 2015. Effects of blood in veins of dragonfly wing on the vibration characteristics. Comput. Biol. Med. 58, 14-19.
Abstract: How the blood in veins of dragonfly wing affects its vibration characteristics is investigated. Based on the experimental results of the wing's morphology and microstructures, including the veins, the membranes and the pterostigma, accurate three-dimensional finite element models of the dragonfly forewing are developed. Considering the blood in veins, the total mass, mass distribution and the moments of inertia of the wing are studied. The natural frequencies/modal shapes are analyzed when the veins are filled with and without blood, respectively. The based natural frequency of the model with blood (189 Hz) is much closer to the experimental result. Relative to bending modal shapes, the torsional ones are affected more significantly by the blood. The results in this article reveal the multi-functions of the blood in dragonfly wings and have important implications for the bionic design of flapping-wing micro air vehicles

Ilvonen, J.J., Suhonen, J., 2016. Phylogeny affects host's weight, immune response and parasitism in damselflies and dragonflies. R. Soc. Open. Sci. 3, 160421.
Abstract: Host-parasite interactions are an intriguing part of ecology, and understanding how hosts are able to withstand parasitic attacks, e.g. by allocating resources to immune defence, is important. Damselflies and dragonflies show a variety of parasitism patterns, but large-scale comparative immune defence studies are rare, and it is difficult to say what the interplay is between their immune defence and parasitism. The aim of this study was to find whether there are differences in immune response between different damselfly and dragonfly species and whether these could explain their levels of gregarine and water mite parasitism. Using an artificial pathogen, a piece of nylon filament, we measured the encapsulation response of 22 different damselfly and dragonfly species and found that (i) there are significant encapsulation differences between species, (ii) body mass has a strong association with encapsulation and parasite prevalences, (iii) body mass shows a strong phylogenetic signal, whereas encapsulation response and gregarine and water mite prevalences show weak signals, and (iv) associations between the traits are affected by phylogeny. We do not know what the relationship is between these four traits, but it seems clear that phylogeny plays a role in determining parasitism levels of damselflies and dragonflies

Ioannidis, P., Simao, F.A., Waterhouse, R.M., Manni, M., Seppey, M., Robertson, H.M., Misof, B., Niehuis, O., Zdobnov, E.M., 2017. Genomic features of the damselfly Calopteryx splendens representing a sister clade to most insect orders. Genome. Biol. Evol., in press.
Abstract: Insects comprise the most diverse and successful animal group with over one million described species that are found in almost every terrestrial and limnic habitat, with many being used as important models in genetics, ecology, and evolutionary research. Genome sequencing projects have greatly expanded the sampling of species from many insect orders, but genomic resources for species of certain insect lineages have remained relatively limited to date. To address this paucity, we sequenced the genome of the banded demoiselle, Calopteryx splendens, a damselfly (Odonata: Zygoptera) belonging to Palaeoptera, the clade containing the first winged insects. The 1.6 Gbp C. splendens draft genome assembly is one of the largest insect genomes sequenced to date and encodes a predicted set of 22,523 protein-coding genes. Comparative genomic analyses with other sequenced insects identified a relatively small repertoire of C. splendens detoxification genes, which could explain its previously noted sensitivity to habitat pollution. Intriguingly, this repertoire includes a cytochrome P450 gene not previously described in any insect genome. The C. splendens immune gene repertoire appears relatively complete and features several genes encoding novel multi-domain peptidoglycan recognition proteins. Analysis of chemosensory genes revealed the presence of both gustatory and ionotropic receptors, as well as the insect odorant receptor coreceptor gene (OrCo) and at least four partner odorant receptors (ORs). This represents the oldest known instance of a complete OrCo/OR system in insects, and provides the molecular underpinning for odonate olfaction. The C. splendens genome improves the sampling of insect lineages that diverged before the radiation of Holometabola and offers new opportunities for molecular-level evolutionary, ecological, and behavioral studies

Jeremiason, J.D., Reiser, T.K., Weitz, R.A., Berndt, M.E., Aiken, G.R., 2016. Aeshnid dragonfly larvae as bioindicators of methylmercury contamination in aquatic systems impacted by elevated sulfate loading. Ecotoxicology 25, 456-468.
Abstract: Methylmercury (MeHg) levels in dragonfly larvae and water were measured over two years in aquatic systems impacted to varying degrees by sulfate releases related to iron mining activity. This study examined the impact of elevated sulfate loads on MeHg concentrations and tested the use of MeHg in dragonfly larvae as an indicator of MeHg levels in a range of aquatic systems including 16 river/stream sites and two lakes. MeHg concentrations in aeshnid dragonfly larvae were positively correlated (R 2 = 0.46, p < 0.01) to peak MeHg concentrations in the dissolved phase for the combined years of 2012 and 2013. This relation was strong in 2012 (R 2 = 0.85, p < 0.01), but showed no correlation in 2013 (R 2 = 0.02, p > 0.05). MeHg in dragonfly larvae were not elevated at the highest sulfate sites, but rather the reverse was generally observed. Record rainfall events in 2012 and above average rainfall in 2013 likely delivered the majority of Hg and MeHg to these systems via interflow and activated groundwater flow through reduced sediments. As a result, the impacts of elevated sulfate releases due to mining activities were not apparent in these systems where little of the sulfate is reduced. Lower bioaccumulation factors for MeHg in aeshnid dragonfly larvae were observed with increasing dissolved organic carbon (DOC) concentrations. This finding is consistent with previous studies showing that MeHg in high DOC systems is less bioavailable; an equilibrium model shows that more MeHg being associated with DOC rather than algae at the base of the food chain readily explains the lower bioaccumulation factors

Kallapur, V.L., 1975. Fuel economy during flight of the dragonfly Pantala flavescens (F). Indian J. Exp. Biol. 13, 200-202.

Kassner, Z., Dafni, E., Ribak, G., 2016. Kinematic compensation for wing loss in flying damselflies. J. Insect. Physiol 85, 1-9.
Abstract: Flying insects can tolerate substantial wing wear before their ability to fly is entirely compromised. In order to keep flying with damaged wings, the entire flight apparatus needs to adjust its action to compensate for the reduced aerodynamic force and to balance the asymmetries in area and shape of the damaged wings. While several studies have shown that damaged wings change their flapping kinematics in response to partial loss of wing area, it is unclear how, in insects with four separate wings, the remaining three wings compensate for the loss of a fourth wing. We used high-speed video of flying blue-tailed damselflies (Ischnura elegans) to identify the wingbeat kinematics of the two wing pairs and compared it to the flapping kinematics after one of the hindwings was artificially removed. The insects remained capable of flying and precise maneuvering using only three wings. To compensate for the reduction in lift, they increased flapping frequency by 18+/-15.4% on average. To achieve steady straight flight, the remaining intact hindwing reduced its flapping amplitude while the forewings changed their stroke plane angle so that the forewing of the manipulated side flapped at a shallower stroke plane angle. In addition, the angular position of the stroke reversal points became asymmetrical. When the wingbeat amplitude and frequency of the three wings were used as input in a simple aerodynamic model, the estimation of total aerodynamic force was not significantly different (paired t-test, p=0.73) from the force produced by the four wings during normal flight. Thus, the removal of one wing resulted in adjustments of the motions of the remaining three wings, exemplifying the precision and plasticity of coordination between the operational wings. Such coordination is vital for precise maneuvering during normal flight but it also provides the means to maintain flight when some of the wings are severely damaged

Kaunisto, K.M., Viitaniemi, H.M., Leder, E.H., Suhonen, J., 2013. Association between host's genetic diversity and parasite burden in damselflies. J. Evol. Biol. 26, 1784-1789.
Abstract: Recent research indicates that low genetic variation in individuals can increase susceptibility to parasite infection, yet evidence from natural invertebrate populations remains scarce. Here, we studied the relationship between genetic heterozygosity, measured as AFLP-based inbreeding coefficient fAFLP , and gregarine parasite burden from eleven damselfly, Calopteryx splendens, populations. We found that in the studied populations, 5-92% of males were parasitized by endoparasitic gregarines (Apicomplexa: Actinocephalidae). Number of parasites ranged from none to 47 parasites per male, and parasites were highly aggregated in a few hosts. Mean individual fAFLP did not differ between populations. Moreover, we found a positive association between individual's inbreeding coefficient and parasite burden. In other words, the more homozygous the individual, the more parasites it harbours. Thus, parasites are likely to pose strong selection pressure against inbreeding and homozygosity. Our results support the heterozygosity-fitness correlation hypothesis, which suggests the importance of heterozygosity for an individual's pathogen resistance

Lancaster, L.T., Dudaniec, R.Y., Chauhan, P., Wellenreuther, M., Svensson, E.I., Hansson, B., 2016. Gene expression under thermal stress varies across a geographic range expansion front. Mol. Ecol. 25, 1141-1156.
Abstract: Many ectothermic species are currently expanding their distributions polewards due to anthropogenic global warming. Molecular genetic mechanisms facilitating range expansion under these conditions are largely unknown, but understanding these could help mitigate expanding pests and disease vectors, or help explain why some species fail to track changing climates. Here, using RNA-seq data, we examine genome-wide changes in gene expression under heat and cold stress in the range-expanding damselfly Ischnura elegans in northern Europe. We find that both the number of genes involved and levels of gene expression under heat stress have become attenuated during the expansion, consistent with a previously-reported release from selection on heat tolerances as species move polewards. Genes upregulated under cold stress differed between core and edge populations, corroborating previously-reported rapid adaptation to cooler climates at the expansion front. Expression of sixty-nine genes exhibited a region x treatment effect; these were primarily upregulated in response to heat stress in core populations but in response to cold stress at the range edge, suggesting that some cellular responses originally adapted to heat stress may switch to cold stress functionality upon encountering novel thermal selection regimes during range expansion. Transcriptional responses to thermal stress involving heat shock and neural function genes were largely geographically conserved, while retrotransposon, regulatory, muscle function and defence gene expression patterns were more variable. Flexible mechanisms of cold stress response and the ability of some genes to shift their function between heat and cold stress might be key mechanisms facilitating rapid poleward expansion in insects. This article is protected by copyright. All rights reserved

Laughlin, S., McGinness, S., 1978. The structures of dorsal and ventral regions of a dragonfly retina. Cell Tissue Res. 188, 427-447.
Abstract: The apposition eyes of the corduliid dragonfly Hemicordulia tau are each divided by pigment colour, facet size and facet arrangement into three regions: dorsal, ventral, and a posterior larval strip. Each ommatidium has two primary pigment cells, twenty-five secondary pigment cells, and eight receptor cells, all surrounded by tracheae which probably prevent light passing between ommatidia, and reduce the weight of the eye. Electron microscopy reveals that the receptor cells are of two types: small vestigial cells making virtually no contribution to the rhabdom, and full-size typical cells. The ventral ommatidia have a distal typical cell (oriented either horizontally or vertically), four medial typical cells, two proximal typical cells and one full-length vestigial cell. The dorsal ommatidia have only four full-length typical cells, and one distal and three vestigial full-length cells. The cross-section of dorsal rhabdoms is small and circular distally, but expands to a large three-pointed star medially and proximally. The tiered receptor arrangement in the ventral ommatidia is typical of other Odonata but the dorsal structure has not been fully described in other species. Specialised dorsal eye regions are typical of insects that detect others against the sky

Letsch, H., Gottsberger, B., Ware, J.L., 2016. Not going with the flow: a comprehensive time-calibrated phylogeny of dragonflies (Anisoptera: Odonata: Insecta) provides evidence for the role of lentic habitats on diversification. Mol. Ecol. 25, 1340-1353.
Abstract: Ecological diversification of aquatic insects has long been suspected to have been driven by differences in freshwater habitats, which can be classified into flowing (lotic) waters, and standing (lentic) waters. The contrasting characteristics of lotic and lentic freshwater systems imply different ecological constraints on their inhabitants. The ephemeral and discontinuous character of most lentic water bodies may encourage dispersal by lentic species in turn reducing geographical isolation among populations. Hence, speciation probability would be lower in lentic species. Here, we assess the impact of habitat use on diversification patterns in dragonflies (Anisoptera: Odonata). Based on eight nuclear and mitochondrial genes, we inferred species diversification with a model-based evolutionary framework, to account for rate variation through time and among lineages, and to estimate the impact of larval habitat on the potentially non-random diversification among anisopteran groups. Ancestral state reconstruction revealed lotic fresh water systems as their original primary habitat, while lentic waters have been colonised independently in Aeshnidae, Corduliidae and Libellulidae. Furthermore, our results indicate a positive correlation of speciation and lentic habitat colonisation by dragonflies: speciation rates increased in lentic Aeshnidae and Libellulidae, whereas they remain mostly uniform among lotic groups. This contradicts the hypothesis of inherently lower speciation in lentic groups and suggests species with larger ranges are more likely to diversify, perhaps due to higher probability of larger areas being dissected by geographical barriers. Furthermore, larger range sizes may comprise more habitat types, which could also promote speciation by providing additional niches, allowing the coexistence of emerging species. This article is protected by copyright. All rights reserved

Li, X.J., Zhang, Z.H., Liang, Y.H., Ren, L.Q., Jie, M., Yang, Z.G., 2014. Antifatigue properties of dragonfly Pantala flavescens wings. Microsc. Res. Tech. 77, 356-362.
Abstract: The wing of a dragonfly is thin and light, but can bear high frequent alternating stress and present excellent antifatigue properties. The surface morphology and microstructure of the wings of dragonfly Pantala flavescens were observed using SEM in this study. Based on the biological analysis method, the configuration, morphology, and structure of the vein were studied, and the antifatigue properties of the wings were investigated. The analytical results indicated that the longitudinal veins, cross veins, and membrane of dragonfly wing form a optimized network morphology and spacially truss-like structure which can restrain the formation and propagation of the fatigue cracks. The veins with multilayer structure present high strength, flexibility, and toughness, which are beneficial to bear alternating load during the flight of dragonfly. Through tensile-tensile fatigue failure tests, the results were verified and indicate that the wings of dragonfly P. flavescens have excellent antifatigue properties which are the results of the biological coupling and synergistic effect of morphological and structural factors

Lin, H.T., Leonardo, A., 2017. Heuristic Rules Underlying Dragonfly Prey Selection and Interception. Curr. Biol. 27, 1124-1137.
Abstract: Animals use rules to initiate behaviors. Such rules are often described as triggers that determine when behavior begins. However, although less explored, these selection rules are also an opportunity to establish sensorimotor constraints that influence how the behavior ends. These constraints may be particularly significant in influencing success in prey capture. Here we explore this in dragonfly prey interception. We found that in the moments leading up to takeoff, perched dragonflies employ a series of sensorimotor rules that determine the time of takeoff and increase the probability of successful capture. First, the dragonfly makes a head saccade followed by smooth pursuit movements to orient its direction-of-gaze at potential prey. Second, the dragonfly assesses whether the prey's angular size and speed co-vary within a privileged range. Finally, the dragonfly times the moment of its takeoff to a prediction of when the prey will cross the zenith. Each of these processes serves a purpose. The angular size-speed criteria biases interception flights to catchable prey, while the head movements and the predictive takeoff ensure flights begin with the prey visually fixated and directly overhead-the key parameters that underlie interception steering. Prey that do not elicit takeoff generally fail at least one of the criterion, and the loss of prey fixation or overhead positioning during flight is strongly correlated with terminated flights. Thus from an abundance of potential targets, the dragonfly selects a stereotyped set of takeoff conditions based on the prey and body states most likely to end in successful capture.

Matushkina, N., Lambret, P., Gorb, S., 2016. Keeping the golden mean: plant stiffness and anatomy as proximal factors driving endophytic oviposition site selection in a dragonfly. Zoology (Jena.) 119, 474-480.
Abstract: Oviposition site selection is a crucial component of habitat selection in dragonflies. The presence of appropriate oviposition plants at breeding waters is considered to be one of the key habitat determinants for species laying eggs endophytically. Thus, Lestes macrostigma, a species which is regarded as threatened in Europe because of its highly disjunct distribution, typically prefers to lay eggs in the sea club rush Bolboschoenus maritimus. However, little is known about how the anatomical and mechanical properties of plant tissues determine the choice of L. macrostigma females. We examined green shoots of six plant species used by L. macrostigma for oviposition, either in the field (actual oviposition plants) or under experimental conditions (potential oviposition plants), to analyse anatomical and mechanical properties of shoots in a framework of known preferences regarding plant substrates for oviposition. As expected, the anatomy of shoots differed between representatives of two plant families, Cyperaceae and Juncaceae, most essentially in the distribution of supporting bundles and the presence of large aeriferous cavities that may affect egg placing within a shoot. The force necessary to puncture the tested plant samples ranged from 360 to 3298 mN, and their local stiffness ranged from 777 to 3363N/m. We show that the shoots of B. maritimus, the plant most preferred by L. macrostigma, have intermediate characteristics regarding both the stiffness and specific anatomical characteristics. The bending stiffness of the ovipositor in L. macrostigma was estimated as 1414N/m, one of the highest values recorded for zygopteran dragonflies so far. The ecological and behavioural implications of plant choice mechanisms in L. macrostigma are discussed in the context of the disjunct distribution of this species

May, M., 1995. Dependence of flight behavior and heat production on air temperature in the green darner dragonfly Anax junius (Odonata: Aeshnidae). J. Exp. Biol. 198, 2385-2392.
Abstract: The large, endothermic dragonfly Anax junius regulates the temperatures of its thorax (Tth) and head (Th) during flight. At high ambient temperature (Ta) it is able to dispose of excess heat from the thorax by increasing hemolymph circulation to the abdomen, but recent evidence suggests that heat loss to the abdomen is largely passive at Ta<30 &deg;C. Nevertheless, these insects continue to regulate Tth and Th at least down to 20 &deg;C and probably at much lower values of Ta. As Ta declines, A. junius glide less, probably fly faster when feeding, and increase their wingbeat frequency when patrolling. Presumably as a result of these behavioral changes, heat production, and thus inferred flight metabolic rate, is inversely proportional to Ta. This is the first demonstration based on field data that an insect regulates body temperature while flying by altering heat production

May, M., 1995. Simultaneous control of head and thoracic temperature by the green darner dragonfly Anax junius (Odonata: Aeshnidae). J. Exp. Biol. 198, 2373-2384.
Abstract: Anax junius is a large dragonfly that regulates thoracic temperature (Tth) during flight. This species, like several other intermittently endothermic insects, achieves control of Tth at least in part by increasing circulation of hemolymph to the abdomen at high air temperature (Ta), thus facilitating heat loss from the thorax. In this paper, I demonstrate that heat transfer to the head is also under active control, very probably owing to temperature-sensitive alteration of hemolymph circulation. As a result, head temperature (Th) is strikingly elevated above Ta during endothermic warm-up and flight. Furthermore, during unrestrained flight in the field, Th is regulated actively by increasing hemolymph circulation from the warm thorax at low Ta. Concurrent measurements of abdominal temperature (Tab) confirm that the abdomen is used as a 'thermal window' at Ta>30 &deg;C but apparently not at lower Ta; thus, some additional mechanism(s) must exist for regulation of Tth at low Ta

May ML, Gregoire JA, Gregoire SM, Lubertazzi MA, Matthews JH. 2017. Emergence phenology, uncertainty, and the evolution of migratory behavior in Anax junius (Odonata: Aeshnidae). PLoS One 12:e0183508.
Abstract: Mass migrations by Odonata, although less studied than those of Monarch butterflies and plague locusts, have provoked comment and study for many years. Relatively recently, increasing interest in dragonflies, supported by new technologies, has resulted in more detailed knowledge of the species involved, behavioral mechanisms, and geographic extent. In this paper we examine, in four independent but complementary studies, how larval habitat and emergence phenology interact with climate to shape the evolution of migratory strategy in Anax junius, a common species throughout much of the eastern United States and southern Canada. In brief, we argue that fish predation on larvae, coupled with the need for ample emergent vegetation for oviposition and adult eclosion, dictates that larval development and survival is optimal in ponds that are neither permanent nor extremely ephemeral. Coupled with annual variation in regional weather and winters in much of their range too cold for adult survival, conditions facing newly emerged A. junius may unpredictably favor either local reproduction or long-distance movement to more favorable areas. Both temperature and hydroperiod tend to favor local reproduction early in the adult activity period and migration later, so late emerging adults are more likely to migrate. No single pond is always predictably suitable or unsuitable, however, so ovipositing females also may spread the risk to their offspring by ovipositing at multiple sites that, for migrants, may be distributed over very long distances

McCauley, S.J., Davis, C.J., Werner, E.E., Robeson, M.S., 2014. Dispersal, niche breadth and population extinction: colonization ratios predict range size in North American dragonflies. J. Anim Ecol. 83, 858-865.
Abstract: Species' range sizes are shaped by fundamental differences in species' ecological and evolutionary characteristics, and understanding the mechanisms determining range size can shed light on the factors responsible for generating and structuring biological diversity. Moreover, because geographic range size is associated with a species' risk of extinction and their ability to respond to global changes in climate and land use, understanding these mechanisms has important conservation implications. Despite the hypotheses that dispersal behaviour is a strong determinant of species range areas, few data are available to directly compare the relationship between dispersal behaviour and range size. Here, we overcome this limitation by combining data from a multispecies dispersal experiment with additional species-level trait data that are commonly hypothesized to affect range size (e.g. niche breadth, local abundance and body size.). This enables us to examine the relationship between these species-level traits and range size across North America for fifteen dragonfly species. Ten models based on a priori predictions about the relationship between species traits and range size were evaluated and two models were identified as good predictors of species range size. These models indicated that only two species' level traits, dispersal behaviour and niche breadth were strongly related to range size. The evidence from these two models indicated that dragonfly species that disperse more often and further had larger North American ranges. Extinction and colonization dynamics are expected to be a key linkage between dispersal behaviour and range size in dragonflies. To evaluate how extinction and colonization dynamics among dragonflies were related to range size we used an independent data set of extinction and colonization rates for eleven dragonfly species and assessed the relationship between these populations rates and North American range areas for these species. We found a negative relationship between North American range size and species' extinction-to-colonization ratios. Our results indicate that metapopulation dynamics act to shape the extent of species' continental distributions. These population dynamics are likely to interact with dispersal behaviour, particularly at species range margins, to determine range limits and ultimately species range sizes

McCauley, S.J., Hammond, J.I., Frances, D.N., Mabry, K.E., 2015. Effects of experimental warming on survival, phenology and morphology of an aquatic insect (Odonata). Ecol. Entomol. 40, 211-220.
Abstract: 1. Organisms can respond to changing climatic conditions in multiple ways including changes in phenology, body size or morphology, and range shifts. Understanding how developmental temperatures affect insect life-history timing and morphology is crucial because body size and morphology affect multiple aspects of life history, including dispersal ability, while phenology can shape population performance and community interactions. 2. We experimentally assessed how developmental temperatures experienced by aquatic larvae affected survival, phenology, and adult morphology of dragonflies (Pachydiplax longipennis). Larvae were reared under 3 environmental temperatures: ambient, +2.5 degrees C, and +5 degrees C, corresponding to temperature projections for our study area 50 and 100 years in the future, respectively. Experimental temperature treatments tracked naturally-occurring variation. 3. We found clear effects of temperature in the rearing environment on survival and phenology: dragonflies reared at the highest temperatures had the lowest survival rates, and emerged from the larval stage approximately 3 weeks earlier than animals reared at ambient temperatures. There was no effect of rearing temperature on overall body size. Although neither the relative wing nor thorax size was affected by warming, a non-significant trend towards an interaction between sex and warming in relative thorax size suggests that males may be more sensitive to warming than females, a pattern that should be investigated further. 4. Warming strongly affected survival in the larval stage and the phenology of adult emergence. Understanding how warming in the developmental environment affects later life-history stages is critical to interpreting the consequences of warming for organismal performance 

Mendonca FZ, Bernardy JV, Oliveira CEK, Oliveira PBG, De MP. 2017. Temperature Effect on the Development of Tropical Dragonfly Eggs. Neotrop Entomol .
Abstract: Physiological constraints in insects are related to several large-scale processes such as species distribution and thermal adaptation. Here, we fill an important gap in ecophysiology knowledge by accessing the relationship between temperature and embrionary development time in four dragonfly species. We evaluated two questions (1) what is the effect of temperature on the development time of Odonata eggs, and (2) considering a degree-day relationship, could a simple linear model describe the dependence of embrionary development time on temperature or it is better described by a more complex non-linear relation. Egg development time of Erythrodiplax fusca (Rambur), Micrathyria hesperis Ris, Perithemis mooma Kirby, and Miathyria simplex (Rambur) (Odonata: Libellulidae) were evaluated. We put the eggs at different temperatures (15, 20, 25, and 30 degrees C) and counted the number of hatched larvae daily. A nonlinear response of the development to the temperature was found, differing from the expected pattern for standard degree-day analysis. Furthermore, we observed that there is a similar process in the development time and hatching synchronization between species, with all species presenting faster egg development at high temperatures. Species-specific differences are more evident at lower temperatures (15 degrees C), with no egg development in M. simplex. Only E. fusca was relatively insensitive to temperature changes with similar hatching rates in all treatments

Meyer-Rochow, V.B., 2017. Therapeutic arthropods and other, largely terrestrial, folk-medicinally important invertebrates: a comparative survey and review. J. Ethnobiol. Ethnomed. 13, 9.
Abstract: Traditional healing methods involving hundreds of insect and other invertebrate species are reviewed. Some of the uses are based on the tenet of "similia similibus" (let likes be cured by likes), but not all non-conventional health promoting practices should be dismissed as superstition or wishful thinking, for they have stood the test of time. Two questions are addressed: how can totally different organ systems in a human possibly benefit from extracts, potions, powders, secretions, ashes, etc. of a single species and how can different target organs, e.g. bronchi, lungs, the urinary bladder, kidneys, etc. apparently respond to a range of taxonomically not even closely related species? Even though therapeutically used invertebrates are generally small, they nevertheless possess organs for specific functions, e.g. digestion, gas exchange, reproduction. They have a nervous system, endocrine glands, a heart and muscle tissue and they contain a multitude of different molecules like metabolites, enzymes, hormones, neurotransmitters, secretions, etc. that have come under increased scientific scrutiny for pharmacological properties. Bearing that in mind it seems likely that a single species prepared and used in different ways could have a multitude of uses. But how, for example, can there be remedies for breathing and other problems, involving earthworms, molluscs, termites, beetles, cockroaches, bugs, and dragonflies? Since invertebrates themselves can suffer from infections and cancers, common defence reactions are likely to have evolved in all invertebrates, which is why it would be far more surprising to find that each species had evolved its own unique disease fighting system. To obtain a more comprehensive picture, however, we still need information on folk medicinal uses of insects and other invertebrates from a wider range of regions and ethnic groups, but this task is hampered by western-based medicines becoming increasingly dominant and traditional healers being unable and sometimes even unwilling to transmit their knowledge to the younger generation. However, collecting and uncontrolled uses of therapeutic invertebrates can put undue pressure on certain highly sought after species and this is something that has to be borne in mind as well

Moore, M.P., Martin, R.A., 2016. Intrasexual selection favours an immune-correlated colour ornament in a dragonfly. J. Evol. Biol. 29, 2256-2265.
Abstract: Sexual signalling is predicted to shape the evolution of sex-specific ornamentation, and establishing the costs and benefits of ornamentation and the information that ornamentation provides to receivers is necessary to evaluating this adaptive function. Here, we assessed the adaptive function of a common colour ornament in insects, melanin wing ornamentation, using the dragonfly Pachydiplax longipennis. We hypothesized that greater ornamentation would improve territory-holding success by decreasing aggression that males receive from territorial rivals, but that more ornamented males may have shorter lifespans. Using mark-recapture field observations, we found that more ornamented males had greater territory-holding success and that viability selection did not act on wing melanization. We then compared the aggression of territorial rivals to decoy males before and after experimentally augmenting wing melanization, finding that males significantly reduced aggression following the manipulation. We next hypothesized that wing melanization would signal fighting ability to territorial rivals by reflecting condition via investment in the costly melanin synthesis pathway. We observed a positive relationship between ornamentation and the likelihood of winning territorial disputes, suggesting that wing melanization provides information about fighting ability to rivals. We also found a positive relationship between melanin-based immune defence and ornamentation, supporting a link between the signal and condition. We conclude that wing melanization is a condition-related signal of fighting ability and suggest that this may be a common mechanism promoting the evolution of melanin ornamentation

Mourao, M.A., Peixoto, P.E., 2014. Do morphological and physiological characteristics of males of the dragonfly Macrothemis imitans determine the winner of territorial contests? J. Insect Sci. 14, 89.
Abstract: Males of many animal species show intraspecific disputes for mating territories that range from displays without physical contact to physical fights with risk of injury. This variation motivated the proposition of different models that suggest possible rules used by rivals to decide the contest winner. To evaluate those models, it is necessary to identify how males behave during the fight and the individual attributes that determine their fighting ability (resource holding potential). For this, males of the dragonfly Macrothemis imitans (Karsch) (Odonata: Libellulidae) were used to evaluate two hypotheses conditioned on the occurrence of physical contact during the fight: if the contests occur with physical contact, features related to size should determine male resource holding potential, and if males do not exhibit physical contact during the contests, features that confer greater endurance should determine resource holding potential. To assess these hypotheses, we collected males that had ownership of territories (resident males) and males that occupied the territory after we removed the resident males (substitute males). After the capture, the resident and substitute males were transferred to the laboratory for measurements of wing area, dry weight, thoracic muscle mass, and fat content. The results showed that resident males do not differ in any measured trait from substitutes. Because the fights occur with physical contact, it is intriguing that resident males do not possess higher fighting capacity than intruders. Perhaps physical contact does not incur high costs during the fight, and other asymmetries, such as motivation associated with prior residency of the disputed territory, determine the contest winner

Nasirian H, Irvine KN. 2017. Odonata larvae as a bioindicator of metal contamination in aquatic environments: application to ecologically important wetlands in Iran. Environ Monit Assess 189:436.
Abstract: The objectives of this study were twofold: (i) assess the bioaccumulation characteristics of a suite of metals associated with several different species of Odonata and (ii) examine Odonata species richness as a reflection of ecosystem health in two ecologically important wetlands of southwestern Iran, the Shadegan and Hawr Al Azim wetlands. Levels of arsenic (As), cadmium (Cd), cobalt (Co), chromium (Cr), copper (Cu), iron (Fe), mercury (Hg), manganese (Mn), lead (Pb), and zinc (Zn) were determined using inductively coupled plasma optical emission spectrometry (ICP-OES) in nine different Odonata larva species. Based on these data, biota-sediment accumulation factors (BSAFs) were calculated and generally, it was found that Cr, Cu, Mn, and Zn were being taken up by the Odonata (BSAFs >1). Because of its prevalence in the wetland and its observed ability to take up metals, it is suggested that Ischnura ramburii is an appropriate indicator of ecosystem health for these wetlands with respect to metal contamination. Odonata species richness across all sites was 49, while for the individual sites, the greatest species richness was 26 and the lowest species richness was 13. The species richness value across all sites is quite healthy, given the arid climate of the region

Nilsson-Ortman, V., Stoks, R., De, B.M., Johansson, F., 2012. Generalists and specialists along a latitudinal transect: patterns of thermal adaptation in six species of damselflies. Ecology. 93, 1340-1352.
Abstract: Tropical organisms colonizing temperate environments face reduced average temperatures and dramatic thermal fluctuations. Theoretical models postulate that thermal specialization should be favored either when little environmental variation is experienced within generations or when among-generation variation is small relative to within-generation variation. To test these predictions, we studied six temperate species of damselflies differing in latitudinal distribution. We developed a computer model simulating how organisms experience environmental variation (accounting for diapause and voltinism) and performed a laboratory experiment assaying thermal sensitivities of growth rates. The computer model showed opposing latitudinal trends in among- and within-generation thermal variability: within-generation thermal variability decreased toward higher latitudes, whereas relative levels of among-generation thermal variability peaked at midlatitudes (where a shift in voltinism occurred). The growth experiment showed that low-latitude species were more thermally generalized than mid- and high-latitude species, supporting the prediction that generalists are favored under high levels of within-generation variation. Northern species had steeper, near-exponential reaction norms suggestive of thermal specialization. However, they had strikingly high thermal optima and grew very slowly over most of the thermal range they are expected to experience in the field. This observation is at present difficult to explain. These results highlight the importance of considering interactions between life history and environmental variation when deriving expectations of thermal adaptation

Okamoto, M., Yasuda, K., Azuma, A., 1996. Aerodynamic characteristics of the wings and body of a dragonfly. J. Exp. Biol. 199, 281-294.
Abstract: The aerodynamic characteristics of the wings and body of a dragonfly and of artificial wing models were studied by conducting two types of wind-tunnel tests and a number of free-flight tests of gliders made using dragonfly wings. The results were consistent between these different tests. The effects of camber, thickness, sharpness of the leading edge and surface roughness on the aerodynamic characteristics of the wings were characterized in the flow field with Reynolds numbers (Re) as low as 103 to 104

Olberg, R.M., Worthington, A.H., Venator, K.R., 2000. Prey pursuit and interception in dragonflies. J. Comp Physiol A 186, 155-162.
Abstract: Perching dragonflies (Libellulidae; Odonata) are sit-and-wait predators, which take off and pursue small flying insects. To investigate their prey pursuit strategy, we videotaped 36 prey-capture flights of male dragonflies, Erythemis simplicicollis and Leucorrhinia intacta, for frame-by-frame analysis. We found that dragonflies fly directly toward the point of prey interception by steering to minimize the movement of the prey's image on the retina. This behavior could be guided by target-selective descending interneurons which show directionally selective visual responses to small-object movement. We investigated how dragonflies discriminate distance of potential prey. We found a peak in angular velocity of the prey shortly before take-off which might cue the dragonfly to nearby flying targets. Parallax information from head movements was not required for successful prey pursuit

Olberg, R.M., Worthington, A.H., Fox, J.L., Bessette, C.E., Loosemore, M.P., 2005. Prey size selection and distance estimation in foraging adult dragonflies. J. Comp Physiol A Neuroethol. Sens. Neural Behav. Physiol 191, 791-797.
Abstract: To determine whether perching dragonflies visually assess the distance to potential prey items, we presented artificial prey, glass beads suspended from fine wires, to perching dragonflies in the field. We videotaped the responses of freely foraging dragonflies (Libellula luctuosa and Sympetrum vicinum-Odonata, suborder Anisoptera) to beads ranging from 0.5 mm to 8 mm in diameter, recording whether or not the dragonflies took off after the beads, and if so, at what distance. Our results indicated that dragonflies were highly selective for bead size. Furthermore, the smaller Sympetrum preferred beads of smaller size and the larger Libellula preferred larger beads. Each species rejected beads as large or larger than their heads, even when the beads subtended the same visual angles as the smaller, attractive beads. Since bead size cannot be determined without reference to distance, we conclude that dragonflies are able to estimate the distance to potential prey items. The range over which they estimate distance is about 1 m for the larger Libellula and 70 cm for the smaller Sympetrum. The mechanism of distance estimation is unknown, but it probably includes both stereopsis and the motion parallax produced by head movements

Olberg, R.M., Seaman, R.C., Coats, M.I., Henry, A.F., 2007. Eye movements and target fixation during dragonfly prey-interception flights. J. Comp Physiol A Neuroethol. Sens. Neural Behav. Physiol 193, 685-693.
Abstract: The capture of flying insects by foraging dragonflies is a highly accurate, visually guided behavior. Rather than simply aiming at the prey's position, the dragonfly aims at a point in front of the prey, so that the prey is intercepted with a relatively straight flight trajectory. To better understand the neural mechanisms underlying this behavior, we used high-speed video to quantify the head and body orientation of dragonflies (female Erythemis simplicicollis flying in an outdoor flight cage) relative to an artificial prey object before and during pursuit. The results of our frame-by-frame analysis showed that during prey pursuit, the dragonfly adjusts its head orientation to maintain the image of the prey centered on the "crosshairs" formed by the visual midline and the dorsal fovea, a high acuity streak that crosses midline at right angles about 60 degrees above the horizon. The visual response latencies to drifting of the prey image are remarkably short, ca. 25 ms for the head and 30 ms for the wing responses. Our results imply that the control of the prey-interception flight must include a neural pathway that takes head position into account

Olberg, R.M., 2012. Visual control of prey-capture flight in dragonflies. Curr. Opin. Neurobiol. 22, 267-271.
Abstract: Interacting with a moving object poses a computational problem for an animal's nervous system. This problem has been elegantly solved by the dragonfly, a formidable visual predator on flying insects. The dragonfly computes an interception flight trajectory and steers to maintain it during its prey-pursuit flight. This review summarizes current knowledge about pursuit behavior and neurons thought to control interception in the dragonfly. When understood, this system has the potential for explaining how a small group of neurons can control complex interactions with moving objects

Outomuro, D., Dijkstra, K.D., Johansson, F., 2013. Habitat variation and wing coloration affect wing shape evolution in dragonflies. J. Evol. Biol. 26, 1866-1874.
Abstract: Habitats are spatially and temporally variable, and organisms must be able to track these changes. One potential mechanism for this is dispersal by flight. Therefore, we would expect flying animals to show adaptations in wing shape related to habitat variation. In this work, we explored variation in wing shape in relation to preferred water body (flowing water or standing water with tolerance for temporary conditions) and landscape (forested to open) using 32 species of dragonflies of the genus Trithemis (80% of the known species). We included a potential source of variation linked to sexual selection: the extent of wing coloration on hindwings. We used geometric morphometric methods for studying wing shape. We also explored the phenotypic correlation of wing shape between the sexes. We found that wing shape showed a phylogenetic structure and therefore also ran phylogenetic independent contrasts. After correcting for the phylogenetic effects, we found (i) no significant effect of water body on wing shape; (ii) male forewings and female hindwings differed with regard to landscape, being progressively broader from forested to open habitats; (iii) hindwings showed a wider base in wings with more coloration, especially in males; and (iv) evidence for phenotypic correlation of wing shape between the sexes across species. Hence, our results suggest that natural and sexual selection are acting partially independently on fore- and hindwings and with differences between the sexes, despite evidence for phenotypic correlation of wing shape between males and females

Outomuro, D., Johansson, F., 2015. Bird predation selects for wing shape and coloration in a damselfly. J. Evol. Biol. 28, 791-799.
Abstract: Wing shape is related to flight performance, which is expected to be under selection for improving flight behaviours such as predator avoidance. Moreover, wing conspicuousness, usually involved in sexual selection processes, is also relevant in terms of predation risk. In this study, we examined how predation by a passerine bird, the white wagtail Motacilla alba, selects wing shape and wing colour patch size in males of the banded demoiselle Calopteryx splendens. The wing colour patch is intra- and intersexually selected in the study species. In a field study, we compared wings of live damselflies to wings of predated damselflies which are always discarded after predation. Based on aerodynamic theory and a previous study on wing shape of territorial tactics in damselflies, we predicted an overall short and broad wing, with a concave front margin shape to be selected by predation. This shape would be expected to improve escaping ability. Moreover, we predicted that wing patch size should be negatively selected by predation. We found that selection operated differently on fore- and hindwings. In contrast to our predictions, predation favoured a slender general forewing shape. However, the predicted wing shape was favoured in hindwings. We also found selection favouring a narrower wing colour patch. Our results suggest different roles of fore- and hindwings in flight, as previously suggested for Calopteryx damselflies and shown for butterflies and moths. Forewings would be more involved in sustained flight and hindwings in flight manoeuvrability. Our results differ somehow from a recently published work in the same study system, but using another population, suggesting that selection can fluctuate across space, despite the simplicity of this predator-prey system

Peisker, H., Gorb, S.N., 2010. Always on the bright side of life: anti-adhesive properties of insect ommatidia grating. J. Exp. Biol. 213, 3457-3462.
Abstract: The surface of some insect eyes consists of arrays of cuticular protuberances, which are 50-300 nm in diameter, and are termed corneal nipples or ommatidia gratings. They were widely reported to reduce the reflectance for normally incident light, contributing to camouflage by reducing glare to predators, while furthermore enhancing the intake of light, which is especially important for nocturnal insects. Our preliminary observations suggest a third function: in contrast to the rest of the body, ommatidia of various insects remain clean, even in a heavy contaminated environment. In order to prove such an anti-contamination hypothesis of these structures, we measured the adhesive properties of polymer moulds of insect ommatidia, and compared these data with control surfaces having the same curvature radii but lacking such a nanostructure. A scanning electron microscope (SEM) study and force measurements using an atomic force microscope (AFM) on the eye surfaces of three different insect species, dragonfly Aeshna mixta (Odonata), moth Laothoe populi (Lepidoptera) and fly Volucella pellucens (Diptera), were undertaken. We revealed that adhesion is greatly reduced by corneal grating in L. populi and V. pellucens when compared with their smooth controls. The smooth cornea of A. mixta showed no statistically significant difference to its control. We assume that this anti-adhesive phenomenon is due to a decrease in the real contact area between contaminating particles and the eye's surface. Such a combination of three functions in one nanostructure can be interesting for the development of industrial multifunctional surfaces capable of enhancing light harvesting while reducing light reflection and adhesion

Pfitzner, W.P., Beck, M., Weitzel, T., Becker, N., 2015. The Role of Mosquitoes in the Diet of Adult Dragon and Damselflies (Odonata). J. Am. Mosq. Control Assoc. 31, 187-189.
Abstract: The flood plains of the Upper Rhine Valley provide excellent conditions for the proliferation of mosquitoes as well as for the development of dragon and damselflies. It could be assumed that mosquitoes belong to the diet of the Odonata and that the latter could be harmed by the reduction of the mosquito population with the purpose of diminishing the massive nuisance for the people living there. A total of 41 adult dragonflies and damselflies were examined by immunoblot for remnants of mosquitoes in their guts. A rabbit antiserum against Aedes vexans proteins was used for the immunoblot. Only 3 Aeshna cyanea and 1 Platycnemis pennipes could be shown to have fed on mosquitoes. In specimens of the genus Sympetrum no mosquitoes were detected. It seems very doubtful that mosquitoes are an essential part of the Odonata diet

Piersanti, S., Rebora, M., Almaas, T.J., Salerno, G., Gaino, E., 2011. Electrophysiological identification of thermo- and hygro-sensitive receptor neurons on the antennae of the dragonfly Libellula depressa. J. Insect Physiol 57, 1391-1398.
Abstract: Recent ultrastructural investigations on Odonata antennal flagellum describe two types of sensilla styloconica, T1 and T2. The styloconic sensilla are located in pits, at the bottom of deep cavities, and share common features typical of thermo-hygroreceptors. In order to ascertain if the Odonata antennae are involved in hygroreception and thermoreception, we carried out electrophysiological recordings (single cell recordings, SCR) from adult males and females of Libellula depressa L., 1758. After contact was established, the antenna was stimulated by rapid changes in temperature and humidity. The present research shows the occurrence of a dry (DC), a moist (MC) and a cold (CC) receptor neurons on the antennal flagellum of L. depressa. These data demonstrate for the first time the presence of functional thermo-hygroreceptors on the antennal flagellum of dragonflies. The present results extend our knowledge of the not visual sensory modalities of Odonata, a field of research unexplored so far

Piersanti, S., Frati, F., Conti, E., Rebora, M., Salerno, G., 2014. The sense of smell in Odonata: an electrophysiological screening. J. Insect Physiol 70, 49-58.
Abstract: Volatile chemicals mediate a great range of intra- and interspecific signalling and information in insects. Olfaction has been widely investigated mostly in Neoptera while the knowledge of this sense in most basal insects such as Paleoptera (Odonata and Ephemeroptera) is still poor. In the present study we show the results of an electrophysiological screening on two model species, Libellula depressa (Libellulidae) and Ischnura elegans (Coenagrionidae), representatives of the two Odonata suborders Anisoptera and Zygoptera, with the aim to deep the knowledge on the sense of smell of this insect order. The antennal olfactory sensory neurons (OSNs) of these two species responded to the same 22 compounds (out of 48 chemicals belonging to different functional groups) encompassing mostly amines, carboxylic acids or aldehydes and belonging to green leaf volatiles, vertebrate related volatiles and volatiles emitted by standing waters bacteria. The properties of Odonata OSNs are very similar to those of ionotropic receptors (IRs) expressing OSNs in other insects

Piersanti, S., Frati, F., Conti, E., Gaino, E., Rebora, M., Salerno, G., 2014. First evidence of the use of olfaction in Odonata behaviour. J. Insect Physiol 62, 26-31.
Abstract: Dragonflies and damselflies are among the most ancient winged insects. Adults belonging to this order are visually oriented and are considered anosmic on the basis of neuroanatomical investigations. As a consequence, the chemical ecology of these predatory insects has long been neglected. Morphological and electrophysiological data demonstrated that dragonfly antennae possess olfactory sensilla. Additionally, a neuroanatomical study revealed the presence of spherical knots in the aglomerular antennal lobe that could allow for the perception of odour. However, the biological role of the antennal olfactory sensilla remains unknown, and no bioassay showing the use of olfaction in Odonata has been performed thus far. Here, we demonstrate through behavioural assays that adults of Ischnura elegans are attracted by olfactory cues emitted by prey; furthermore, using electrophysiological single-cell recordings, we prove that the antennal olfactory sensilla of I. elegans respond to prey odour. Our results clearly demonstrate the involvement of antennal olfactory sensilla in Odonata predation, thus showing, for the first time, the use of olfaction in Odonata biology. This finding indicates that the nervous system of Odonata is able to receive and process olfactory information, suggesting that the simple organisation of the antennal lobe does not prevent the use of olfaction in insects

Piersanti, S., Frati, F., Rebora, M., Salerno, G., 2016. Carbon dioxide detection in adult Odonata. Zoology (Jena) 117, 137-142.
Abstract: The present paper shows, by means of single-cell recordings, responses of antennal sensory neurons of the damselfly Ischnura elegans when stimulated by air streams at different CO2 concentrations. Unlike most insects, but similarly to termites, centipedes and ticks, Odonata possess sensory neurons strongly inhibited by CO2, with the magnitude of the off-response depending upon the CO2 concentration. The Odonata antennal sensory neurons responding to CO2 are also sensitive to airborne odors; in particular, the impulse frequency is increased by isoamylamine and decreased by heptanoic and pentanoic acid. Further behavioral investigations are necessary to assign a biological role to carbon dioxide detection in Odonata

Popova, O.N., Haritonov, A.Y., Sushchik, N.N., Makhutova, O.N., Kalachova, G.S., Kolmakova, A.A., Gladyshev, M.I., 2017. Export of aquatic productivity, including highly unsaturated fatty acids, to terrestrial ecosystems via Odonata. Sci. Total. Environ. 581-582, 40-48.
Abstract: Based on 31-year field study of the abundance and biomass of 18 species of odonates in the Barabinsk Forest-Steppe (Western Siberia, Russia), we quantified the contribution of odonates to the export of aquatic productivity to surrounding terrestrial landscape. Emergence varied from 0.8 to 4.9g of wet biomass per m2 of land area per year. Average export of organic carbon was estimated to be 0.30g.m-2.year-1, which is comparable with the average production of herbivorous terrestrial insects in temperate grasslands. Moreover, in contrast to terrestrial insects, emerging odonates contained high quantities of highly unsaturated fatty acids (HUFA), namely eicosapentaenoic acid (20:5n-3, EPA), and docosahexaenoic acid (22:6n-3, DHA), which are known to be essential for many terrestrial animals, especially for birds. The export of EPA+DHA by odonates was found to be 1.92-11.76mg.m-2.year-1, which is equal to an average general estimation of the export of HUFA by emerging aquatic insects. Therefore, odonates appeared to be a quantitatively and qualitatively important conduit of aquatic productivity to forest-steppe ecosystem

Rajabi H, Ghoroubi N, Stamm K, Appel E, Gorb SN. 2017. Dragonfly wing nodus: A one-way hinge contributing to the asymmetric wing deformation. Acta Biomater 60:330-338.
Abstract: Dragonfly wings are highly specialized locomotor systems, which are formed by a combination of several structural components. The wing components, also known as structural elements, are responsible for the various aspects of the wing functionality. Considering the complex interactions between the wing components, modelling of the wings as a whole is only possible with inevitable huge oversimplifications. In order to overcome this difficulty, we have recently proposed a new approach to model individual components of complex wings comparatively. Here, we use this approach to study nodus, a structural element of dragonfly wings which has been less studied to date. Using a combination of several imaging techniques including scanning electron microscopy (SEM), wide-field fluorescence microscopy (WFM), confocal laser scanning microscopy (CLSM) and micro-computed tomography (micro-CT) scanning, we aim to characterize the spatial morphology and material composition of fore- and hindwing nodi of the dragonfly Brachythemis contaminata. The microscopy results show the presence of resilin in the nodi, which is expected to help the deformability of the wings. The computational results based on three-dimensional (3D) structural data suggest that the specific geometry of the nodus restrains its displacements when subjected to pressure on the ventral side. This effect, resulting from an interlocking mechanism, is expected to contribute to the dorso-ventral asymmetry of wing deformation and to provide a higher resistance to aerodynamic forces during the downstroke. Our results provide an important step towards better understanding of the structure-property-function relationship in dragonfly wings. STATEMENT OF SIGNIFICANCE: In this study, we investigate the wing nodus, a specialized wing component in dragonflies. Using a combination of modern imaging techniques, we demonstrate the presence of resilin in the nodus, which is expected to facilitate the wing deformability in flight. The specific geometry of the nodus, however, seems to restrain its displacements when subjected to pressure on the ventral side. This effect, resulting from an interlocking mechanism, is suggested to contribute to dorso-ventral asymmetry of wing deformations and to provide a higher resistance to aerodynamic forces during the downstroke. Our results provide an important step towards better understanding of the structure-property-function relationship in dragonfly wings and might help to design more efficient wings for biomimetic micro-air vehicles

Rajabi H, Schroeter V, Eshghi S, Gorb SN. 2017. The probability of wing damage in the dragonfly Sympetrum vulgatum (Anisoptera: Libellulidae): a field study. Biol Open 6:1290-1293.
Abstract: Dragonfly wings resist millions of cycles of dynamic loading in their lifespan. During their operation, the wings are subjected to relatively high mechanical stresses. They further experience accidental collisions which result from the insects' daily activities, such as foraging, mating and fighting with other individuals. All these factors may lead to irreversible wing damage. Here, for the first time, we collected qualitative and quantitative data to systematically investigate the occurrence of damage in dragonfly wings in nature. The results obtained from the analysis of 119 wings from >30 individual Sympetrum vulgatum (Anisoptera: Libellulidae), collected at the second half of their flight period, indicate a high risk of damage in both fore- and hindwings. Statistical analyses show no significant difference between the extent of damage in fore- and hindwings, or between male and female dragonflies. However, we observe a considerable difference in the probability of damage in different wing regions. The wing damage is found to mainly result from two failure modes: wear and fracture

Sanchez-Guillen RA, Wellenreuther M, Chavez-Rios JR et al. Alternative reproductive strategies and the maintenance of female color polymorphism in damselflies. Ecol Evol 2017;7:5592-5602.
Abstract: Genetic polymorphisms are powerful model systems to study the maintenance of diversity in nature. In some systems, polymorphisms are limited to female coloration; these are thought to have arisen as a consequence of reducing male mating harassment, commonly resulting in negative frequency-dependent selection on female color morphs. One example is the damselfly Ischnura elegans, which shows three female color morphs and strong sexual conflict over mating rates. Here, we present research integrating male tactics, and female evolutionary strategies (female mating behavior and morph-specific female fecundity) in populations with different morph-specific mating frequencies, to obtain an understanding of mating rates in nature that goes beyond the mere measure of color frequencies. We found that female morph behavior differed significantly among but not within morphs (i.e., female morph behavior was fixed). In contrast, male tactics were strongly affected by the female morph frequency in the population. Laboratory work comparing morph-specific female fecundity revealed that androchrome females have lower fecundity than both of the gynochrome female morphs in the short term (3-days), but over a 10-day period one of the gynochrome female morphs became more fecund than either of the other morphs. In summary, our study found sex-specific dynamics in response to different morph frequencies and also highlights the importance of studying morph-specific fecundities across different time frames to gain a better understanding of the role of alternative reproductive strategies in the maintenance of female-limited color polymorphism

Rasmussen, N.L., Rudolf, V.H., 2016. Individual and combined effects of two types of phenological shifts on predator-prey interactions. Ecology 97, 3414-3421.
Abstract: Timing of phenological events varies among years with natural variation in environmental conditions and is also shifting in response to climate change. These phenological shifts likely have many effects on species interactions. Most research on the ecological consequences of phenological shifts has focused on variation in simple metrics such as phenological firsts. However, for a population, a phenological event exhibits a temporal distribution with many attributes that can vary (e.g., mean, variance, skewness), each of which likely has distinct effects on interactions. In this study, we manipulated two attributes of the phenological distribution of a prey species to determine their individual and combined effects on predator-prey interactions. Specifically, we studied how shifts in the mean and variation around the mean (i.e., synchrony) of hatching by tadpoles (Hyla cinerea) affected interactions with predatory dragonfly naiads (Tramea carolina). At the end of larval development, we quantified survival and growth of predator and prey. We found that both types of shifts altered demographic rates of the prey; that the effects of synchrony shifts, though rarely studied, were at least as strong as those due to mean shifts; and that the combined effects of shifts in synchrony and mean were additive rather than synergistic. By dissecting the roles of two types of shifts, this study represents a significant step toward a comprehensive understanding of the complex effects of phenological shifts on species interactions. Embracing this complexity is critical for predicting how climate change will alter community dynamics

Rau, P., 1945. The night habits of the dragonfly, Ajax junius Dru. J. Comp Psychol. 38, 285.

Rebora, M., Salerno, G., Piersanti, S., Dell'otto, A., Gaino, E., 2012. Olfaction in dragonflies: electrophysiological evidence. J. Insect Physiol 58, 270-277.
Abstract: The problem of olfaction in Paleoptera (Odonata, Ephemeroptera) cannot be considered fully elucidated until now. These insects have been traditionally considered anosmic, because their brain lacks glomerular antennal lobes, typically involved in Neoptera odor perception. In order to understand if the presumed coeloconic olfactory receptors described on the antennal flagellum of adult Odonata are really functioning, we performed an electrophysiological investigation with electroantennogram (EAG) and single cell recordings (SCR), using Libellula depressa L. (Odonata, Libellulidae) as a model species. Odors representing different chemical classes such as (Z)-3-hexenyl acetate (acetate ester), (E)-2-hexenal, octanal (aldehydes), (Z)-3-hexen-1-ol (alcohol), propionic acid, butyric acid (carboxylic acids), and 1,4-diaminobutane (amine) were tested. Most of the tested chemicals elicited depolarizing EAG responses in both male and female antennae; SCR show unambiguously for the first time the presence of olfactory neurons in the antennae of L. depressa and strongly support the olfactory function of the coeloconic sensilla located on the antennal flagellum of this species. Electrophysiological activity may not necessarily indicate behavioral activity, and the biological role of olfactory responses in Odonata must be determined in behavioral bioassays. This study represents a starting point for further behavioral, electrophysiological, neuroanatomical and molecular investigation on Odonata olfaction, a research field particularly interesting owing to the basal position of Paleoptera, also for tracing evolutionary trends in insect olfaction

Saito, V.S., Valente-Neto, F., Rodrigues, M.E., de Oliveira, R.F., Siqueira, T., 2016. Phylogenetic clustering among aggressive competitors: evidence from odonate assemblages along a riverine gradient. Oecologia. 182, 219-229.
Abstract: Studies on phylogenetic community ecology usually infer habitat filtering when communities are phylogenetically clustered or competitive exclusion when communities are overdispersed. This logic is based on strong competition and niche similarity among closely related species-a less common phenomenon than previously expected. Dragonflies and damselflies are good models for testing predictions based on this logic because they behave aggressively towards related species due to mistaken identification of conspecifics. This behavior may drive communities toward phylogenetic overdispersion if closely related species frequently exclude each other. However, phylogenetically clustered communities could also be observed if habitat filtering and/or competitive asymmetry among distantly related species are major drivers of community assembling. We investigated the phylogenetic structure of odonate assemblages in central Brazil in a watershed characterized by variations in stream width, vegetation cover, aquatic vegetation, and luminosity. We observed general clustering in communities according to two indices of phylogenetic structure. Phylogenetic beta diversity coupled with Mantel tests and RLQ analysis evidenced a correlation between the riverine gradient and phylogenetic structure. Larger rivers with aquatic vegetation were characterized by anisopterans, while most zygopterans stayed in small and shaded streams. These results indicate niche conservatism in Odonata habitat occupancy, and that the environment is a major influence on the phylogenetic structure of these communities. We suggest that this is due to clade-specific ecophysiological requirements, and because closely related species may also have competitive advantages and dominate certain preferred habitats

Sanchez-Guillen, R.A., Martinez-Zamilpa, S.M., Jimenez-Cortes, J.G., Forbes, M.R., Cordoba-Aguilar, A., 2013. Maintenance of polymorphic females: do parasites play a role? Oecologia. 171, 105-113.
Abstract: The role of parasites in explaining maintenance of polymorphism is an unexplored research avenue. In odonates, female-limited color polymorphism (one female morph mimicking the conspecific male and one or more gynochromatic morphs) is widespread. Here we investigated whether parasitism contributes to color polymorphism maintenance by studying six species of female dimorphic damselflies using large databases of field-collected animals. We predicted that androchrome females (male mimics) would be more intensively parasitized than gynochrome females which is, according to previous studies, counterbalanced by the advantages of the former when evading male harassment compared to gynochrome females. Here we show that in Ischnura denticollis and Enallagma novahispaniae, androchrome females suffer from a higher degree of parasitism than gynochromatic females, and contrary to prediction, than males. Thus, our study has detected a correlation between color polymorphism and parasitic burden in odonates. This leads us to hypothesize that natural selection, via parasite pressure, can explain in part how androchrome and gynochrome female color morphs can be maintained. Both morphs may cope with parasites in a different way: given that androchrome females are more heavily parasitized, they may pay a higher fecundity costs, in comparison to gynochrome females

Sanmartin-Villar, I., Cordero-Rivera, A., 2016. The inheritance of female colour polymorphism in Ischnura genei (Zygoptera: Coenagrionidae), with observations on melanism under laboratory conditions. PeerJ. 4, e2380.
Abstract: Current research on female colour polymorphism in Ischnura damselflies suggests that a balanced fitness trade-off between morphotypes contributes to the maintenance of polymorphism inside populations. The genetic inheritance system constitutes a key factor to understand morph fluctuation and fitness. Ischnura genei, an endemic species of some Mediterranean islands, has three female colour morphs, including one androchrome (male-coloured) and two gynochromes. In this study, we reared two generations of I. genei under laboratory conditions and tested male behavioural responses to female colour morphs in the field. We recorded ontogenetic colour changes and studied morph frequency in three populations from Sardinia (Italy). Morph frequencies of laboratory crosses can be explained by a model based on an autosomal locus with three alleles and sex-restricted expression, except for one crossing of 42 families with unexpected offspring. The allelic dominance relationship was androchrome > infuscans > aurantiaca. Old individuals reared in the laboratory exhibited different levels of melanism in variable extent depending on sex and morph. Results of model presentations indicate a male preference for gynochrome females and the lack of recognition of androchromes as potential mates. Aurantiaca females were the most frequent morph in the field (63-87%). Further studies in other populations and islands are needed to understand the maintenance of this polymorphism

Sherratt, T.N., Laird, R.A., Hassall, C., Lowe, C.D., Harvey, I.F., Watts, P.C., Cordero-Rivera, A., Thompson, D.J., 2010. Empirical evidence of senescence in adult damselflies (Odonata: Zygoptera). J. Anim Ecol. 79, 1034-1044.
Abstract: 1. Age-dependent increases in mortality have been documented in a variety of species of insect under laboratory conditions. However, while strong statistical evidence has been presented for senescence in vertebrate populations in the wild, we know little about the rate and shape of senescence in wild populations of insects. 2. Odonates (damselflies and dragonflies) provide excellent candidate species for evaluating demographic senescence as they are large enough to be marked individually and they are easily re-sighted without recapture. The prevailing opinion - based entirely on qualitative examination of the declines in log numbers alive with time since marking - is that odonates exhibit age-independent daily survivorship. 3. Here, we examine mark-recapture data on the Azure Damselfly Coenagrion puella over two consecutive seasons. For the first time, we evaluate and compare the fit of quantitative models that not only account for weather-dependent daily variation in daily re-sighting rates, but also age-dependent variation in daily survivorship. 4. Models with age-dependent declines in daily survivorship provide a more parsimonious explanation for the data than similar models without these age-dependent effects. In general, models in which mortality increases in an exponential (Gompertz) fashion explain the mark-recapture sequences more efficiently than a range of alternative models, including those in which mortality increases as a power function (Weibull) or reaches a plateau (logistic). These results are indicative of a general senescent decline in physiological functioning, which is particularly marked after 15 days as a mature adult. 5. Weather (temperature, sun and precipitation) and initial mite load influenced the probability of daily re-sighting. Weather and mite load also influenced daily survivorship, but their effects differed between seasons. 6. Overall, fitting models with age as an explicit covariate demonstrates that odonates do indeed senesce. This contradicts previously held assumptions that Odonata do not exhibit age-dependent survivorship in the wild

Sherratt, T.N., Hassall, C., Laird, R.A., Thompson, D.J., Cordero-Rivera, A., 2011. A comparative analysis of senescence in adult damselflies and dragonflies (Odonata). J. Evol. Biol. 24, 810-822.
Abstract: Any population whose members are subject to extrinsic mortality should exhibit an increase in mortality with age. Nevertheless, the prevailing opinion is that populations of adult damselflies and dragonflies do not exhibit such senescence. Here, we challenge this contention by fitting a range of demographic models to the data on which these earlier conclusions were based. We show that a model with an exponential increase in age-related mortality (Gompertz) generally provides a more parsimonious fit than alternative models including age-independent mortality, indicating that many odonates do indeed senesce. Controlling for phylogeny, a comparison of the daily mortality of 35 odonate species indicates that although male and female mortalities are positively correlated, mortality tends to be higher in males of those species that exhibit territoriality. Hence, we show for the first time that territoriality may impose a survivorship cost on males, once the underlying phylogenetic relationships are accounted for

Simon, E., Kis, O., Jakab, T., Kolozsvari, I., Malnas, K., Harangi, S., Baranyai, E., Miskolczi, M., Tothmeresz, B., Devai, G., 2017. Assessment of contamination based on trace element concentrations in Gomphus flavipes (Odonata: Insect) larvae of the Upper Tisza Region. Ecotoxicol. Environ. Saf 136, 55-61.
Abstract: Odonata larvae are frequently used to assess the contamination of aquatic systems, because they tolerate a wide range of chemical and biological conditions in freshwater systems. In early 2000, the sediments of the Hungarian section of the River Tisza and the River Szamos were strongly enriched with heavy metals by an accidental mining spill. Earlier studies demonstrated higher contamination levels in the Szamos than in the Tisza, based on sediment analysis. The aim of our study was to assess the contamination in the Upper Tisza Region, along the upper reach of the Tisza, and the lower reach of the Szamos, based on the trace element concentrations of the Gomphus flavipes larvae. We collected 269 dragonfly specimens for the analyses. The Al, Ba, Cr, Cu, Fe, Mn, Pb, Sr and Zn element contents were analysed in the dragonfly larvae by microwave plasma atomic emission spectrometry (MP-AES). Significantly higher Ba and Cu concentrations were found in the dragonfly larvae of the Tisza than the Szamos. In spite of this, the Cr, Mn, Pb, Sr and Zn concentration was significantly lower in the dragonfly larvae of the Tisza than the Szamos. For all trace elements significant differences were found along the Tisza. Significant differences were also found in all trace element concentrations of dragonfly larvae among studied localities in the Szamos, except in the cases of Al and Ba. Our results demonstrated that the Szamos was more contaminated with Cr, Mn, Pb, Sr and Zn than the Tisza, but that the Tisza was more contaminated with Ba and Cu than the Szamos, based on the trace element concentrations in Gomphus flavipes larvae, which was likely to have been caused by the tributaries of the Tisza. In summary, our results indicated a continuous pollution of the Tisza and the Szamos and their tributaries

Siepielski, A.M., Beaulieu, J.M., 2017. Adaptive evolution to novel predators facilitates the evolution of damselfly species range shifts. Evolution. 71, 974-984.
Abstract: Most species have evolved adaptations to reduce the chances of predation. In many cases, adaptations to coexist with one predator generate tradeoffs in the ability to live with other predators. Consequently, the ability to live with one predator may limit the geographic distributions of species, such that adaptive evolution to coexist with novel predators may facilitate range shifts. In a case study with Enallagma damselflies, we used a comparative phylogenetic approach to test the hypothesis that adaptive evolution to live with a novel predator facilitates range size shifts. Our results suggest that the evolution of Enallagma shifting from living in ancestral lakes with fish as top predators, to living in lakes with dragonflies as predators, may have facilitated an increase in their range sizes. This increased range size likely arose because lakes with dragonflies were widespread, but unavailable as a habitat throughout much of the evolutionary history of Enallagma because they were historically maladapted to coexist with dragonfly predators. Additionally, the traits that have evolved as defenses against dragonflies also likely enhanced damselfly dispersal abilities. While many factors underlie the evolutionary history of species ranges, these results suggest a role for the evolution of predator-prey interactions.

Sniegula, S., Golab, M.J., Drobniak, S.M., Johansson, F., 2016. Seasonal time constraints reduce genetic variation in life-history traits along a latitudinal gradient. J. Anim Ecol. 85, 187-198.
Abstract: Time constraints cause strong selection on life-history traits, because populations need to complete their life cycles within a shorter time. We therefore expect lower genetic variation in these traits in high- than in low-latitude populations, since the former are more time-constrained. The aim was to estimate life-history traits and their genetic variation in an obligately univoltine damselfly along a latitudinal gradient of 2730 km. Populations were grown in the laboratory at temperatures and photoperiods simulating those at their place of origin. In a complementary experiment, individuals from the same families were grown in constant temperature and photoperiod that mimicked average conditions across the latitude. Development time and size was faster and smaller, respectively, and growth rate was higher at northern latitudes. Additive genetic variance was very low for life-history traits, and estimates for egg development time and larval growth rate showed significant decreases towards northern latitudes. The expression of genetic effects in life-history traits differed considerably when individuals were grown in constant rather than simulated and naturally variable conditions. Our results support strong selection by time constraints. They also highlight the importance of growing organisms in their native environment for correct estimates of genetic variance at their place of origin. Our results also suggest that the evolutionary potential of life-history traits is very low at northern compared to southern latitudes, but that changes in climate could alter this pattern

Sniegula, S., Janssens, L., Stoks, R., 2017. Integrating multiple stressors across life stages and latitudes: Combined and delayed effects of an egg heat wave and larval pesticide exposure in a damselfly. Aquat. Toxicol. 186, 113-122.
Abstract: To understand the effects of pollutants in a changing world we need multistressor studies that combine pollutants with other stressors associated with global change such as heat waves. We tested for the delayed and combined impact of a heat wave during the egg stage and subsequent sublethal exposure to the pesticide esfenvalerate during the larval stage on life history and physiology in the larval and adult stage of the damselfly Lestes sponsa. We studied this in a common garden experiment with replicated central- and high latitude populations to explore potential effects of local thermal adaptation and differences in life history shaping the multistressor responses. Exposure of eggs to the heat wave had no effect on larval traits, yet had delayed costs (lower fat and flight muscle mass) in the adult stage thereby crossing two life history transitions. These delayed costs were only present in central-latitude populations potentially indicating their lower heat tolerance. Exposure of larvae to the pesticide reduced larval growth rate and prolonged development time, and across metamorphosis reduced the adult fat content and the flight muscle mass, yet did not affect the adult heat tolerance. The pesticide-induced delayed emergence was only present in the slower growing central-latitude larvae, possibly reflecting stronger selection to keep development fast in the more time-constrained high-latitude populations. We observed no synergistic interactions between the egg heat wave and the larval pesticide exposure. Instead the pesticide-induced reduction in fat content was only present in animals that were not exposed to the egg heat wave. Our results based on laboratory conditions highlight that multistressor studies should integrate across life stages to fully capture cumulative effects of pollutants with other stressors related to global change.

Start, D., Kirk, D., Shea, D., Gilbert, B., 2017. Cannibalism by damselflies increases with rising temperature. Biol. Lett. 13.
Abstract: Trophic interactions are likely to change under climate warming. These interactions can be altered directly by changing consumption rates, or indirectly by altering growth rates and size asymmetries among individuals that in turn affect feeding. Understanding these processes is particularly important for intraspecific interactions, as direct and indirect changes may exacerbate antagonistic interactions. We examined the effect of temperature on activity rate, growth and intraspecific size asymmetries, and how these temperature dependencies affected cannibalism in Lestes congener, a damselfly with marked intraspecific variation in size. Temperature increased activity rates and exacerbated differences in body size by increasing growth rates. Increased activity and changes in body size interacted to increase cannibalism at higher temperatures. We argue that our results are likely to be general to species with life-history stages that vary in their temperature dependencies, and that the effects of climate change on communities may depend on the temperature dependencies of intraspecific interactions.

Stoks, R., Cordoba-Aguilar, A., 2012. Evolutionary ecology of Odonata: a complex life cycle perspective. Annu. Rev. Entomol. 57, 249-265.
Abstract: 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

Suarez-Tovar, C.M., Sarmiento, C.E., 2016. Beyond the wing planform: morphological differentiation between migratory and non-migratory dragonfly species. J. Evol. Biol. 29, 690-703.
Abstract: Migration is a significant trait of the animal kingdom that can impose a strong selective pressure on several structures to overcome the amount of energy that the organism invests in this particular behavior. Wing linear dimensions and planform have been a traditional focus in the study of flying migratory species; however, other traits could also influence aerodynamic performance. We studied the differences in several flight-related traits of migratory and non-migratory Libellulid species in a phylogenetic context to assess their response to migratory behavior. Wings were compared by linear measurements, shape, surface corrugations, and microtrichia number. Thorax size and pilosity were also compared. Migratory species have larger and smoother wings, a larger anal lobe that is reached through an expansion of the discoidal region, and longer and denser thoracic pilosity. These differences might favor gliding as an energy-saving displacement strategy. Most of the changes were identified in the hind wings. No differences were observed for the thorax linear dimensions, wetted aspect ratio, some wing corrugations, or the wing microtrichiae number. Similar changes in the hind wing are present in clades where migration evolved. Our results emphasize that adaptations to migration through flight may extend to characteristics beyond the wing planform and that some wing characteristics in libellulids converge in response to migratory habits whereas other closely related structures remain virtually unchanged. Additionally, we concluded that despite a close functional association and similar selective pressures on a structure, significant differences in the magnitude of the response may be present in its components. This article is protected by copyright. All rights reserved

Suhonen, J., Honkavaara, J., Rantala, M.J., 2010. Activation of the immune system promotes insect dispersal in the wild. Oecologia. 162, 541-547.
Abstract: Dispersal has important ecological and evolutionary consequences but is a poorly understood behaviour. We experimentally tested whether activation of the immune system affects dispersal in male damselflies, Calopteryx virgo, from three natural populations. We show that males that contained an experimentally inserted artificial pathogen, a nylon monofilament implant, had higher dispersal rates and flew further than control males, but not further than sham manipulated males. Our data suggest that dispersal may reduce the risk of further infections if immune system activation indicates high parasite infection risk in the present habitat. We, thus, suggest that parasites may play an important role in the evolution of host dispersal

Suhonen, J., Korkeamaki, E., Salmela, J., Kuitunen, M., 2014. Risk of local extinction of Odonata freshwater habitat generalists and specialists. Conserv. Biol. 28, 783-789.
Abstract: Understanding the risk of a local extinction in a single population relative to the habitat requirements of a species is important in both theoretical and applied ecology. Local extinction risk depends on several factors, such as habitat requirements, range size of species, and habitat quality. We studied the local extinctions among 31 dragonfly and damselfly species from 1930 to 1975 and from 1995 to 2003 in Central Finland. We tested whether habitat specialists had a higher local extinction rate than generalist species. Approximately 30% of the local dragonfly and damselfly populations were extirpated during the 2 study periods. The size of the geographical range of the species was negatively related to extinction rate of the local populations. In contrast to our prediction, the specialist species had lower local extinction rates than the generalist species, probably because generalist species occurred in both low- and high-quality habitat. Our results are consistent with source-sink theory

Suvorov, A., Jensen, N.O., Sharkey, C.R., Fujimoto, M.S., Bodily, P., Wightman, H.M., Ogden, T.H., Clement, M.J., Bybee, S.M., 2016. Opsins have evolved under the permanent heterozygote model: insights from phylotranscriptomics of Odonata. Mol. Ecol., in press.
Abstract: Gene duplication plays a central role in adaptation to novel environments by providing new genetic material for functional divergence and evolution of biological complexity. Several evolutionary models have been proposed for gene duplication to explain how new gene copies are preserved by natural selection but these models have rarely been tested using empirical data. Opsin proteins, when combined with a chromophore, form a photopigment that is responsible for the absorption of light, the first step in the phototransduction cascade. Adaptive gene duplications have occurred many times within the animal opsins gene family, leading to novel wavelength sensitivities. Consequently, opsins are an attractive choice for the study of gene duplication evolutionary models. Odonata (dragonflies and damselflies) have the largest opsin repertoire of any insect currently known. Additionally, there is tremendous variation in opsin copy number between species, particularly in the long wavelength sensitive (LWS) class. Using comprehensive phylotranscriptomic and statistical approaches we tested various evolutionary models of gene duplication. Our results suggest that both the blue sensitive (BS) and LWS opsin classes were subjected to strong positive selection that greatly weakens after multiple duplication events, a pattern that is consistent with the permanent heterozygote model. Due to the immense interspecific variation and duplicability potential of opsin genes among odonates, they represent a unique model system to test hypotheses regarding opsin gene duplication and diversification at the molecular level.

Svensson, E.I., Waller, J.T., 2013. Ecology and sexual selection: evolution of wing pigmentation in calopterygid damselflies in relation to latitude, sexual dimorphism, and speciation. Am. Nat. 182, E174-E195.
Abstract: Our knowledge about how the environment influences sexual selection regimes and how ecology and sexual selection interact is still limited. We performed an integrative study of wing pigmentation in calopterygid damselflies, combining phylogenetic comparative analyses, field observations and experiments. We investigated the evolutionary consequences of wing pigmentation for sexual dimorphism, speciation, and extinction and addressed the possible thermoregulatory benefits of pigmentation. First, we reconstructed ancestral states of male and female phenotypes and traced the evolutionary change of wing pigmentation. Clear wings are the ancestral state and that pigmentation dimorphism is derived, suggesting that sexual selection results in sexual dimorphism. We further demonstrate that pigmentation elevates speciation and extinction rates. We also document a significant biogeographic association with pigmented species primarily occupying northern temperate regions with cooler climates. Field observations and experiments on two temperate sympatric species suggest a link between pigmentation, thermoregulation, and sexual selection, although body temperature is also affected by other phenotypic traits such as body mass, microhabitat selection, and thermoregulatory behaviors. Taken together, our results suggest an important role for wing pigmentation in sexual selection in males and in speciation. Wing pigmentation might not increase ecological adaptation and species longevity, and its primary function is in sexual signaling and species recognition

Swaegers, J., Janssens, S.B., Ferreira, S., Watts, P.C., Mergeay, J., McPeek, M.A., Stoks, R., 2014. Ecological and evolutionary drivers of range size in Coenagrion damselflies. J. Evol. Biol. 27, 2386-2395.
Abstract: Geographic range size is a key ecological and evolutionary characteristic of a species, yet the causal basis of variation in range size among species remains largely unresolved. One major reason for this is that several ecological and evolutionary traits may jointly shape species' differences in range size. We here present an integrated study of the contribution of ecological (dispersal capacity, body size and latitudinal position) and macroevolutionary (species' age) traits in shaping variation in species' range size in Coenagrion damselflies. We reconstructed the phylogenetic tree of this genus to account for evolutionary history when assessing the contribution of the ecological traits and to evaluate the role of the macroevolutionary trait (species' age). The genus invaded the Nearctic twice independently from the Palearctic, yet this was not associated with the evolution of larger range sizes or dispersal capacity. Body size and species' age did not explain variation in range size. There is higher flight ability (as measured by wing aspect ratio) at higher latitudes. Species with a larger wing aspect ratio had a larger range size, also after correcting for phylogeny, suggesting a role for dispersal capacity in shaping the species' ranges. More northern species had a larger species' range, consistent with Rapoport's rule, possibly related to niche width. Our results underscore the importance of integrating macroecology and macroevolution when explaining range size variation among species

Takahashi, Y., Suyama, Y., Matsuki, Y., Funayama, R., Nakayama, K., Kawata, M., 2016. Lack of genetic variation prevents adaptation at the geographic range margin in a damselfly. Mol. Ecol. 25, 4450-4460.
Abstract: What limits a species' distribution in the absence of physical barriers? Genetic load due to asymmetric gene flow and the absence of genetic variation due to lack of gene flow are hypothesized to constrain adaptation to novel environments in marginal populations, preventing range expansion. Here, we examined the genetic structure and geographic variation in morphological traits in two damselflies (Ischnura asiatica and I. senegalensis) along a latitudinal gradient in Japan, which is the distribution centre of I. asiatica and the northern limit of I. senegalensis. Genomewide genetic analyses found a loss of genetic diversity at the edge of distribution in I. senegalensis but consistently high diversity in I. asiatica. Gene flow was asymmetric in a south-north direction in both species. Although body size and wing loading showed decreasing latitudinal clines (smaller in north) in I. asiatica in Japan, increasing latitudinal clines (larger in north) in these phenotypic markers were observed in I. senegalensis, particularly near the northern boundary, which coincided well with the location where genetic diversity began a sharp decline. In ectothermic animals, increasing latitudinal cline in these traits was suggested to be established when they failed to adapt to thermal gradient. Therefore, our findings support the possibility that a lack of genetic variation rather than geneflow swamping is responsible for the constraint of adaptation at the margin of geographic distribution

Takamura, K., 1996. Life cycle of the damselfly Calopteryx atrata in relation to pesticide contamination. Ecotoxicology. 5, 1-8.
Abstract: The life cycle of the damselfly Calopteryx atrata was investigated in relation to pesticide contamination occurring in its aquatic habitat. Calopteryx atrata emerged from the River Onogawa around May and stayed as immature adults in forests away from the stream. From late June to mid-August, mature adults were engaged in reproduction at the stream. On the other hand, pesticide contamination occurred from April to August with its peak in May and June, following transplantation of young rice plants. Mature nymphs of C. atrata experienced pesticide contamination, but may have tolerated it. Hatched nymphs had high susceptibility to two of the commonly used insecticides, fenitrothion (mortality occurred at >4.0 mug l(-1) in 24 h and at >2.0 mug l(-1) in 48 h) and fenthion (>2.0 mug l(-1) in 24 h and >1.0 mug l(-1) in 48 h). Hatching was estimated to occur mainly in August, when pesticide contamination was not as high as the susceptibility level. However, the level of pesticide contamination in August is variable due to its origin from aerial spraying, so hatched nymphs may experience a hazardous amount of pesticides depending on the year or place. The population of C. atrata does not escape the risk of pesticide contamination completely and may be affected by it

Tennessen, K.J., 2016. Psaironeura angeloi, a new species of damselfly (Zygoptera: Coenagrionidae) from Central and South America. Zootaxa. 4078, 28-37.
Abstract: Psaironeura angeloi sp. nov. (Holotype male deposited in FSCA: ECUADOR, Esmeraldas Province, small stream 5.6 km NW of Lita, 00.893 degrees N 78.510 degrees W, 4.II.1997, KJT leg.) is described and illustrated based on specimens from Ecuador, Panama, Costa Rica, and Nicaragua, bringing the total number of species in the genus to five. The new species is closely related to P. remissa (Calvert), a Mexican/northern Central American species with broad, foliate male cerci, but is distinct in that the long flagella of the genital ligula lack a small sharp spine unique to P. remissa, labrum and clypeus are orange-red, and the back of the head is mostly pale in both males and females. In life, the eyes of the new species are bright red in males versus green and black in P. remissa

Tiitsaar, A., Kaasik, A., Teder, T., 2013. The effects of seasonally variable dragonfly predation on butterfly assemblages. Ecology. 94, 200-207.

Abstract: Where predation is seasonally variable, the potential impact of a predator on individual prey species will critically depend on phenological synchrony of the predator with the prey. Here we explored the effects of seasonally variable predation in multispecies assemblages of short-lived prey. The study was conducted in a landscape in which we had previously demonstrated generally high, but spatially and seasonally variable dragonfly-induced mortality in adult butterflies. In this system, we show that patterns of patch occupancy in butterfly species flying during periods of peak dragonfly abundance are more strongly associated with spatial variation in dragonfly abundance than patch occupancy of species flying when dragonfly density was low. We provide evidence indicating that this differential sensitivity of different butterfly species to between-habitat differences in dragonfly abundance is causally tied to seasonal variation in the intensity of dragonfly predation. The effect of dragonfly predation could also be measured at the level of whole local butterfly assemblages. With dragonfly density increasing, butterfly species richness decreased, and butterfly species composition tended to show a shift toward a greater proportion of species flying during periods of off-peak dragonfly abundance

Troast, D., Suhling, F., Jinguji, H., Sahlen, G., Ware, J., 2016. A Global Population Genetic Study of Pantala flavescens. PLoS. One. 11, e0148949.
Abstract: Among terrestrial arthropods, the dragonfly species Pantala flavescens is remarkable due to their nearly global distribution and extensive migratory ranges; the largest of any known insect. Capable of migrating across oceans, the potential for high rates of gene flow among geographically distant populations is significant. It has been hypothesized that P. flavescens may be a global panmictic population but no sufficient genetic evidence has been collected thus far. Through a population genetic analysis of P. flavescens samples from North America, South America, and Asia, the current study aimed to examine the extent at which gene flow is occurring on a global scale and discusses the implications of the genetic patterns we uncovered on population structure and genetic diversity of the species. This was accomplished using PCR-amplified cytochrome oxidase one (CO1) mitochondrial DNA data to reconstruct phylogenetic trees, a haplotype network, and perform molecular variance analyses. Our results suggested high rates of gene flow are occurring among all included geographic regions; providing the first significant evidence that Pantala flavescens should be considered a global panmictic population.

Tsuchiya, K., Hayashi, F., 2014. Left-handed sperm removal by male Calopteryx damselflies (Odonata). Springerplus. 3, 144.
Abstract: Male genitalia in several insect species are asymmetry in right and left shape. However, the function of such asymmetric male genitalia is still unclear. We found that the male genitalia of the damselfly Calopteryx cornelia (Odonata: Calopterygidae) are morphologically symmetric just after emergence but asymmetric after reproductive maturation. Males remove rival sperm stored in the female bursa copulatrix (single spherical sac) and the following spermatheca (Y-shaped tubular sac) prior to their own ejaculation to prevent sperm competition. Males possess the aedeagus with a recurved head to remove bursal sperm and a pair of spiny lateral processes to remove spermathecal sperm. The right lateral process is less developed than the left, and sperm stored in the right spermathecal tube are rarely removed. Experiments involving surgical cutting of each lateral process demonstrated that only the left process functions in spermathecal sperm removal. Thus, males of C. cornelia are left-handed in their sperm removal behaviour at copulation

Tsubaki, Y., Kato, C., Shintani, S., 2006. On the respiratory mechanism during underwater oviposition in a damselfly Calopteryx cornelia Selys. J. Insect Physiol 52, 499-505.
Abstract: Calopteryx cornelia females oviposit almost exclusively underwater in forest streams. Field observation showed that the duration of uninterrupted submerged oviposition ranged between 20 and 120 min and the number of eggs laid was linearly related to the time spent underwater. By holding a damselfly under water in a small jar, we measured the maximum 'submergence potential', which was defined as the time elapsed between placing the insect underwater and asphyxiation. A series of experiments showed that there was no gender difference in the submergence potential. This was about 120 min if a damselfly was allowed to change its position while under water. The submergence potential was shorter if the damselflies were kept motionless, if air bubbles trapped on the wing surfaces were removed by coating with Vaseline or if the water was hypoxic. By contrast, submergence potential was longer if a part of the wings were kept above the water surface, or if the water was agitated using a magnetic stirrer. These results suggest that ovipositing C. cornelia females depend for oxygen on the physical-gill action of the thin air layer trapped on the body and wing surfaces. Respiration capacity under water is not likely to be a limiting factor for ovipositing females during the production of a single clutch

Tuzun N, Op de BL, Stoks R. 2017. Sexual selection reinforces a higher flight endurance in urban damselflies. Evol Appl 10:694-703.
Abstract: Urbanization is among the most important and globally rapidly increasing anthropogenic processes and is known to drive rapid evolution. Habitats in urbanized areas typically consist of small, fragmented and isolated patches, which are expected to select for a better locomotor performance, along with its underlying morphological traits. This, in turn, is expected to cause differentiation in selection regimes, as populations with different frequency distributions for a given trait will span different parts of the species' fitness function. Yet, very few studies considered differentiation in phenotypic traits associated with patterns in habitat fragmentation and isolation along urbanization gradients, and none considered differentiation in sexual selection regimes. We investigated differentiation in flight performance and flight-related traits and sexual selection on these traits across replicated urban and rural populations of the scrambling damselfly Coenagrion puella. To disentangle direct and indirect paths going from phenotypic traits over performance to mating success, we applied a path analysis approach. We report for the first time direct evidence for the expected better locomotor performance in urban compared to rural populations. This matches a scenario of spatial sorting, whereby only the individuals with the best locomotor abilities colonize the isolated urban populations. The covariation patterns and causal relationships among the phenotypic traits, performance and mating success strongly depended on the urbanization level. Notably, we detected sexual selection for a higher flight endurance only in urban populations, indicating that the higher flight performance of urban males was reinforced by sexual selection. Taken together, our results provide a unique proof of the interplay between sexual selection and adaptation to human-altered environments

Van, G.H., Stoks, R., Matthysen, E., Valck, F., De, B.L., 1999. Male choice for female colour morphs in Ischnura elegans (Odonata, Coenagrionidae): testing the hypotheses. Anim Behav. 57, 1229-1232.
Abstract: The occurrence of different conspecific female colour morphs, with one of the morphs resembling the male, is supposed to have consequences for mate choice. There are two hypotheses linking mate choice and female colour polymorphism. First, males may mate predominantly with female morphs that differ from the male because they do not recognize androchrome females as females (male mimic hypothesis). Second, males may be more attracted to the most common morph in the population (habituation hypothesis). We tested these hypotheses in five populations of the same species, Ischnura elegans, with a range of androchrome frequencies. In each population we performed binary choice experiments in small cages. Males did not consistently prefer gynochrome females but mated predominantly with the most common morph in the population. Moreover, a reanalysis of the available damselfly data in the literature also supported the habituation hypothesis. Copyright 1999 The Association for the Study of Animal Behaviour

Van, G.H., De, B.L., Stoks, R., 2005. Reversible switches between male-male and male-female mating behaviour by male damselflies. Biol. Lett. 1, 268-270.
Abstract: For many animal groups, both sexes have been reported to attempt to mate with members of their own sex. Such behaviour challenges theories of sexual selection, which predict optimization of reproductive success. We tested male mate choice between opposite- and same-sex members in the damselfly Ischnura elegans. Binary choice experiments were conducted following exposure periods in insectaries with only males or with both sexes present. We show that switches in choice between the opposite sex and the same sex can be induced and reversed again by changing the social context. We argue that the observed reversibility in male-male- and male-female-directed mating behaviour is maladaptive and a consequence of strong selection on a male's ability to alter choice between different female colour morphs

Van, P.N., De, B.L., De, J.M., Vanhaecke, L., Stoks, R., Bervoets, L., 2014. Can damselfly larvae (Ischnura elegans) be used as bioindicators of sublethal effects of environmental contamination? Aquat. Toxicol. 154 , 270-277.
Abstract: The present study measured various pesticides and trace metals, together with sublethal effect biomarkers (lipid, protein and glycogen levels, acetylcholinesterase (AChE) and glutathione-S-transferase (GST) activities) in damselfly larvae (Ischnura elegans) at sixteen sampling sites in Flanders (Belgium). Four pesticides (chloridazon, dichlorvos, terbutylazine, metolachlor), some of them hardly measurable in surface water, and all trace metals were above the limit of quantification in damselfly tissue. A principal component analysis (PCA) on the accumulated pollutant concentrations returned five pollutant axes explaining 85.8% of the total variation. Based on these PCA-axes a hierarchical cluster analysis revealed that the 16 sampled ponds could be classified in 7 groups. Increasing dichlorvos levels in the animals resulted in a lower body mass. Body mass was negatively correlated with GST and AChE activities, lipid and glycogen levels. The present findings provide evidence of toxicity-induced sublethal stress of dichlorvos accumulation in natural populations of I. elegans

Verzijden, M.N., Scobell, S.K., Svensson, E.I., 2014. The effects of experience on the development of sexual behaviour of males and females of the banded demoiselle (Calopteryx splendens). Behav. Processes 109 Pt B , 180-189.
Abstract: Mate preferences can vary in the direction of the preference, as well as the strength of the preference, and both direction and strength of preference are known to be plastic in many species. Preferences might have a learned component, and current and past social context may influence an individual's choosiness. In the damselfly species Calopteryx splendens, females increase the strength of their mate preferences with sexual experience. Here we show that sexually naive females selectively respond to conspecific courtship as soon as physical contact has been established, suggesting a role for tactile cues perceived through interspecific morphological differences in secondary reproductive traits. In addition our data also shows that males and females selectively respond to the intensity of the courtship of the potential, conspecific mate, while ignoring such information in heterospecific potential mates. These results underscore that mate choice is the result of dynamic interactions between the sexes, where both current and past information are integrated. This article is part of a Special Issue entitled: Cognition in the Wild

Villalobos-Jimenez, G., Hassall, C., 2017. Effects of the urban heat island on the phenology of Odonata in London, UK. Int. J. Biometeorol., in press.
Abstract: Urbanisation is one of the major drivers of ecosystem change and includes increased temperatures in cities leading to an urban heat island (UHI). This study quantified the phenological response of odonates across London, UK, from 1990 to 2012, using a database of 1,031,277 historical sightings. The ordinal flight dates of each species were used to calculate the leading edge, middle and trailing edge of the flight period (P5, P50 and P95, respectively). The results suggest that the phenology of odonates is affected by the UHI only at a community level: no significant changes in the P5 or P50 of the flight period were found, although the P95 shows a mean advance of 4.13 days compared to rural areas, thus suggesting a contraction of the flight period in urban areas. However, only one individual species (Sympetrum striolatum) exhibited an advance in the P95 of the flight period in urban areas compared to rural areas. On the other hand, climate change (minimum temperature) had a much stronger impact on the phenology of odonates at the community level with a significant advance of 6.9 days degrees C-1 in the P5 of the flight period, 3.1 days degrees C-1 in the P50 and 3.3 days degrees C-1 in the P95 flight date. Similarly, a significant advance in P5 was found in 7 of the 15 species tested in response to minimum temperature, and 2 species showed a significant advance in P50 in response to minimum temperature, but no species showed a shift in the P95 flight date due to minimum temperature. As shown in previous studies, life history influences the phenological response of odonates, with spring species and those species lacking an egg diapause being the most responsive to increased temperatures, although summer species and species with obligate egg diapause also respond to the UHI by advancing the P95 by 3.8 and 4.5 days, respectively, compared to rural areas, thus contracting the flight period. The present study shows that the UHI has negligible impacts on emergence patterns of odonates compared to climate change, which may result from the capacity of aquatic habitats to buffer the microclimatic conditions of the surrounding terrestrial habitats. We conclude by highlighting the importance of climate change on freshwater habitats over the impacts of the UHI

Watts, P.C., Rouquette, J.R., Saccheri, I.J., Kemp, S.J., Thompson, D.J., 2004. Molecular and ecological evidence for small-scale isolation by distance in an endangered damselfly, Coenagrion mercuriale. Mol. Ecol. 13, 2931-2945.
Abstract: Coenagrion mercuriale (Charpentier) (Odonata: Zygoptera) is one of Europe's most threatened damselflies and is listed in the European Habitats directive. We combined an intensive mark-release-recapture (MRR) study with a microsatellite-based genetic analysis for C. mercuriale from the Itchen Valley, UK, as part of an effort to understand the dispersal characteristics of this protected species. MRR data indicate that adult damselflies are highly sedentary, with only a low frequency of interpatch movement that is predominantly to neighbouring sites. This restricted dispersal leads to significant genetic differentiation throughout most of the Itchen Valley, except between areas of continuous habitat, and isolation by distance (IBD), even though the core populations are separated by less than 10 km. An urban area separating some sites had a strong effect on the spatial genetic structure. Average pairwise relatedness between individual damselflies is positive at short distances, reflecting fine-scale genetic clustering and IBD both within- and between-habitat patches. Damselflies from a fragmented habitat have higher average kinship than those from a large continuous population, probably because of poorer dispersal and localized breeding in the former. Although indirect estimates of gene flow must be interpreted with caution, it is encouraging that our results indicate that the spatial pattern of genetic variation matches closely with that expected from direct observations of movement. These data are further discussed with respect to possible barriers to dispersal within the study site and the ecology and conservation of C. mercuriale. To our knowledge, this is the first report of fine-scale genetic structuring in any zygopteran species

Wellenreuther, M., Sanchez-Guillen, R.A., 2016. Nonadaptive radiation in damselflies. Evol. Appl. 9, 103-118.
Abstract: Adaptive radiations have long served as living libraries to study the build-up of species richness; however, they do not provide good models for radiations that exhibit negligible adaptive disparity. Here, we review work on damselflies to argue that nonadaptive mechanisms were predominant in the radiation of this group and have driven species divergence through sexual selection arising from male-female mating interactions. Three damselfly genera (Calopteryx,Enallagma and Ischnura) are highlighted and the extent of (i) adaptive ecological divergence in niche use and (ii) nonadaptive differentiation in characters associated with reproduction (e.g. sexual morphology and behaviours) was evaluated. We demonstrate that species diversification in the genus Calopteryx is caused by nonadaptive divergence in coloration and behaviour affecting premating isolation, and structural differentiation in reproductive morphology affecting postmating isolation. Similarly, the vast majority of diversification events in the sister genera Enallagma and Ischnura are entirely driven by differentiation in genital structures used in species recognition. The finding that closely related species can show negligible ecological differences yet are completely reproductively isolated suggests that the evolution of reproductive isolation can be uncoupled from niche-based divergent natural selection, challenging traditional niche models of species coexistence

Wiederman, S.D., O'Carroll, D.C., 2013. Selective attention in an insect visual neuron. Curr. Biol. 23, 156-161.
Abstract: Animals need attention to focus on one target amid alternative distracters. Dragonflies, for example, capture flies in swarms comprising prey and conspecifics, a feat that requires neurons to select one moving target from competing alternatives. Diverse evidence, from functional imaging and physiology to psychophysics, highlights the importance of such "competitive selection" in attention for vertebrates. Analogous mechanisms have been proposed in artificial intelligence and even in invertebrates, yet direct neural correlates of attention are scarce from all animal groups. Here, we demonstrate responses from an identified dragonfly visual neuron that perfectly match a model for competitive selection within limits of neuronal variability (r(2) = 0.83). Responses to individual targets moving at different locations within the receptive field differ in both magnitude and time course. However, responses to two simultaneous targets exclusively track those for one target alone rather than any combination of the pair. Irrespective of target size, contrast, or separation, this neuron selects one target from the pair and perfectly preserves the response, regardless of whether the "winner" is the stronger stimulus if presented alone. This neuron is amenable to electrophysiological recordings, providing neuroscientists with a new model system for studying selective attention

Wikelski, M., Moskowitz, D., Adelman, J.S., Cochran, J., Wilcove, D.S., May, M.L., 2006. Simple rules guide dragonfly migration. Biol. Lett. 2, 325-329.
Abstract: Every year billions of butterflies, dragonflies, moths and other insects migrate across continents, and considerable progress has been made in understanding population-level migratory phenomena. However, little is known about destinations and strategies of individual insects. We attached miniaturized radio transmitters (ca 300 mg) to the thoraxes of 14 individual dragonflies (common green darners, Anax junius) and followed them during their autumn migration for up to 12 days, using receiver-equipped Cessna airplanes and ground teams. Green darners exhibited distinct stopover and migration days. On average, they migrated every 2.9+/-0.3 days, and their average net advance was 58+/-11 km in 6.1+/-0.9 days (11.9+/-2.8 km d-1) in a generally southward direction (186+/-52 degrees). They migrated exclusively during the daytime, when wind speeds were less than 25 km h-1, regardless of wind direction, but only after two nights of successively lower temperatures (decrease of 2.1+/-0.6 degrees C in minimum temperature). The migratory patterns and apparent decision rules of green darners are strikingly similar to those proposed for songbirds, and may represent a general migration strategy for long-distance migration of organisms with high self-propelled flight speeds

Willkommen, J., Michels, J., Gorb, S.N., 2015. Functional morphology of the male caudal appendages of the damselfly Ischnura elegans (Zygoptera: Coenagrionidae). Arthropod. Struct. Dev. 44, 289-300.
Abstract: Odonata are usually regarded as one of the most ancient extant lineages of winged insects. Their copulatory apparatus and mating behavior are unique among insects. Male damselflies use their caudal appendages to clasp the female's prothorax during both copulation and egg-laying and have a secondary copulatory apparatus for sperm transfer. Knowledge of the functional morphology of the male caudal appendages is the basis for understanding the evolution of these structures in Odonata and respective organs in other insects. However, it is still not exactly known how the zygopteran claspers work. In this study, we applied micro-computed tomography and a variety of microscopy techniques to examine the morphology, surface microstructure, cuticle material composition and muscle topography of the male caudal appendages of Ischnura elegans. The results indicate that the closing of the paraproctal claspers is mainly passive. This indirect closing mechanism is very likely supported by high proportions of the elastic protein resilin present in the cuticle of the paraproctal bases. In addition, the prothoracic morphology of the female plays an important role in the indirect closing of the male claspers. Our data indicate that both structures - the male claspers and the female prothoracic hump - function together like a snap-fastener

Worthen, W.B., Horacek, H.J., 2015. The distribution of dragonfly larvae in a South Carolina stream: relationships with sediment type, body size, and the presence of other larvae. J. Insect Sci. 15.
Abstract: Dragonfly larvae were sampled in Little Creek, Greenville, SC. The distributions of five common species were described relative to sediment type, body size, and the presence of other larvae. In total, 337 quadrats (1 m by 0.5 m) were sampled by kick seine. For each quadrat, the substrate was classified as sand, sand-cobble mix, cobble, coarse, or rock, and water depth and distance from bank were measured. Larvae were identified to species, and the lengths of the body, head, and metafemur were measured. Species were distributed differently across sediment types: sanddragons, Progomphus obscurus (Rambur) (Odonata: Gomphidae), were common in sand; twin-spotted spiketails, Cordulegaster maculata Selys (Odonata: Cordulegastridae), preferred a sand-cobble mix; Maine snaketails, Ophiogomphus mainensis Packard (Odonata: Gomphidae), preferred cobble and coarse sediments; fawn darners, Boyeria vinosa (Say) (Odonata: Aeshnidae), preferred coarse sediments; and Eastern least clubtails, Stylogomphus albistylus (Hagen) (Odonata: Gomphidae), preferred coarse and rock sediments. P. obscurus and C. maculata co-occurred more frequently than expected by chance, as did O. mainensis, B. vinosa, and S. albistylus. Mean size varied among species, and species preferences contributed to differences in mean size across sediment types. There were significant negative associations among larval size classes: small larvae (<12 mm) occurred less frequently with large larvae (>15 mm) than expected by chance, and large larvae were alone in quadrats more frequently than other size classes. Species may select habitats at a large scale based on sediment type and their functional morphology, but small scale distributions are consistent with competitive displacement or intraguild predation.

Zeuss, D., Brandl, R., Brandle, M., Rahbek, C., Brunzel, S., 2014. Global warming favours light-coloured insects in Europe. Nat. Commun. 5, 3874.
Abstract: Associations between biological traits of animals and climate are well documented by physiological and local-scale studies. However, whether an ecophysiological phenomenon can affect large-scale biogeographical patterns of insects is largely unknown. Insects absorb energy from the sun to become mobile, and their colouration varies depending on the prevailing climate where they live. Here we show, using data of 473 European butterfly and dragonfly species, that dark-coloured insect species are favoured in cooler climates and light-coloured species in warmer climates. By comparing distribution maps of dragonflies from 1988 and 2006, we provide support for a mechanistic link between climate, functional traits and species that affects geographical distributions even at continental scales. Our results constitute a foundation for better forecasting the effect of climate change on many insect groups

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