Odonata (dragonflies and damselflies) as a bridge between ecology and evolutionary genomics

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.     

A comparative analysis of senescence in adult damselflies and dragonflies (Odonata)

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.     

An exploration of genome size diversity in dragonflies and damselflies (Insecta: Odonata)

Like most insect orders, the Odonata (dragonflies and damselflies) remain poorly studied from the perspective of genome size. They exhibit several characteristics that make them desirable targets for analysis in this area, for example a large range in body size, differences in developmental rate, and distinct modes of flight – all of which are related to genome size in at least some animal taxa. The present study provides new genome size estimates and morphometric data for 100 species of odonates, covering about 1/5 of described North American diversity. Significant relationships are reported between genome size and body size (positive in dragonflies, negative in damselflies), and there is also indication that developmental rate and flight are related to genome size in these insects. Genome size is also positively correlated with chromosome number across the order. These findings contribute to an improved understanding of genome size evolution in insects, and raise several interesting questions for future research.     

Tolerance of low pH exposure by the eggs of Odonata (dragonflies and damselflies)

The development times and hatching success of the eggs of four species of Odonata (Ischnura verticalis, Lestes congener, Libellula lydia, and Sympetrum vicinum) were unaffected by exposure to soft water at pH 5.1 and 3.5. Tolerance of low pH, soft water conditions by Odonata eggs may in part account for the widespread distribution of Odonata in potentially acid-stressed regions.     

Phylogeny and classification of the Stenophlebioptera (Odonata: Epiproctophora)

The Juraheterophlebiidae, new family of the “heterophlebioid” lineage, the Henrotayiidae, new family of the “anisopteroid” lineage, the Prostenophlebiidae and the Liassostenophlebiidae, new families of the Stenophlebioptera, and three new genera and species of the Stenophlebiidae are described from the Mesozoic of Germany, Spain, England, Kazakhstan, and Mongolia. The phylogenetic positions of the families Erichschmidiidae and Gondvanogomphidae are discussed. A tentative phylogenetic analysis of the Anisopteromorpha is proposed. This significantly extends our knowledge on the palaeogeographical distribution of the Stenophlebioptera and the Epiproctophora (“dragondamselflies”).     

Changes in occurrence,richness, and biological traits of dragonflies and damselflies (Odonata) in California and Nevada over the past century

Increases in water demand, urbanization, and severity of drought threaten freshwater ecosystems of the arid western United States. Historical assessments of change in assemblages over time can help determine the effects of these stressors but, to date, are rare. In the present study, we resurveyed 45 sites originally sampled in 1914–1915 for Odonata (dragonflies and damselflies) adults throughout central California and northwestern Nevada, USA. We examined changes in species occurrence rates, taxonomic richness, and biological trait composition in relation to climate changes and human population increases. While species richness at individual sites did not change significantly, we found that odonate assemblages have become more similar across sites. Homogenization is a result of the expansion of highly mobile habitat generalists, and the decline of both habitat specialists and species with an overwintering diapause stage. Using a multi-species mixed-effects model, we found that overall occurrences of Odonata increased with higher minimum temperatures. Habitat specialists and species with a diapause stage, however, occurred less often in warmer regions and more often in areas with higher precipitation. Habitat specialists occurred less often in highly populated sites. Life history traits of Odonata, such as dispersal ability, habitat specialization, and diapause, are useful predictors of species-specific responses to urbanization and climate change in this region.     

Flight of the dragonflies and damselflies

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’.     

Phylogeny of the higher Libelluloidea (Anisoptera: Odonata): an exploration of the most speciose superfamily of dragonflies

Although libelluloid dragonflies are diverse, numerous, and commonly observed and studied, their phylogenetic history is uncertain. Over 150 years of taxonomic study of Libelluloidea Rambur, 1842, beginning with Hagen (1840), [Rambur, M.P., 1842. Neuropteres. Histoire naturelle des Insectes, Paris, pp. 534; Hagen, H., 1840. Synonymia Libellularum Europaearum. Dissertation inaugularis quam consensu et auctoritate gratiosi medicorum ordinis in academia albertina ad summos in medicina et chirurgia honores.] and Selys (1850), [de Selys Longchamps, E., 1850. Revue des Odonates ou Libellules d'Europe [avec la collaboration de H.A. Hagen]. Muquardt, Bruxelles; Leipzig, 1-408.], has failed to produce a consensus about family and subfamily relationships. The present study provides a well-substantiated phylogeny of the Libelluloidea generated from gene fragments of two independent genes, the 16S and 28S ribosomal RNA (rRNA), and using models that take into account non-independence of correlated rRNA sites. Ninety-three ingroup taxa and six outgroup taxa were amplified for the 28S fragment; 78 ingroup taxa and five outgroup taxa were amplified for the 16S fragment. Bayesian, likelihood and parsimony analyses of the combined data produce well-resolved phylogenetic hypotheses and several previously suggested monophyletic groups were supported by each analysis. Macromiinae, Corduliidae s. s., and Libellulidae are each monophyletic. The corduliid (s.l.) subfamilies Synthemistinae, Gomphomacromiinae, and Idionychinae form a monophyletic group, separate from the Corduliinae. Libellulidae comprises three previously accepted subfamilies (Urothemistinae, a very restricted Tetrathemistinae, and a modified Libellulinae) and five additional consistently recovered groups. None of the other previously proposed subfamilies are supported. Bayesian analyses run with an additional 71 sequences obtained from GenBank did not alter our conclusions. The evolution of adult and larval morphological characters is discussed here to suggest areas for future focus. This study shows the inherent problems in using poorly defined and sometimes inaccurately scored characters, basing groups on symplesiomorphies, and failure to recognize the widespread effects of character correlation and convergence, especially in aspects of wing venation.     

Adaptations of dragonflies (Odonata) under desert conditions

Different types of adaptations of dragonflies to climatic conditions of the desert zone are described. The main habitats of the dragonfly nymphs are lotic and semi-lotic artificial reservoirs. The life cycles are synchronous with seasonal climatic changes, reproduction being restricted to periods of the optimal hygrothermal conditions. Some species reveal vertical seasonal migrations. The labile daily activity rhythms allow the dragonflies to avoid the effects of unfavorable conditions. Adults of many species emerge at night.     

A checklist of the dragonflies (Odonata) of Kenya

A checklist of Odonata has been compiled for Kenya. It is based on an inventory of museum material, publications and personal observations made between 1978 and 1997. Changes of scientific names and synonyms are documented. The list contains 194 valid dragonfly species recorded for Kenya.     

Evolution of reproductive strategies in libellulid dragonflies (Odonata: Anisoptera)

In Libellulidae, oocyte production has been assumed to be continuous, with periods of egg-laying interspersed with periods of resting/eating; however, recent work suggests that two types of oocyte production are common: either (a) continuous or (b) step-wise. These are mirrored in the arrangement of the ovarioles in the ovaries. Likewise, two types of mate-guarding behavior have been observed in Libellulidae: (1) non?Ccontact guarding and (2) tandem guarding in which the male either hovers above the female or is physically attached to her during oviposition. Using molecular (mitochondrial and nuclear) data we explored the evolution of female reproductive traits, focusing on ovariole morphology, as well as guarding behavior, in Libellulidae. Continuous egg production appears to have evolved more than once, as have tandem and non-contact guarding. We discuss how the evolution of different ovariole types and guarding behavior may have been influenced by habitat instability, dispersal and crowded oviposition sites; thus, migratory behavior or habitat availability may have been the driving force of ovariole evolution.     

Distribution patterns of dragonflies (Odonata) in Central Asia

The dragonfly fauna of Central Asia reveals distinct vertical differentiation. Three groups of species can be distinguished: mountain (24 species), plain (18), and plain-mountain (34) ones. The ranges observed can be classified into 7 principal types: plain, mountain, continuous boreo-montane, disjunctive Central Asian boreomontane, disjunctive Tien Shan boreo-montane, Central Asian, and Pamir-Alay plain-mountain. The leading factors determining the distribution of dragonflies are the temperature and the presence of streams suitable for preimaginal development; the former factor determines the potential ranges, and the latter, the actual ones. The present-day composition, structure, and vertical distribution of the dragonfly fauna formed during the historical time, after the development of artificial irrigation canals which provided new habitats for dragonflies.     

Crevice organs: Sensory structures on the wings of dragonflies (Insecta,Odonata)

Summary Crevice organs are small, elongate, innervated indentations in the hard cuticle of one wing vein of aeshnid dragonflies. There are four groups on each wing. The structure and orientation of crevice organs suggest that they detect strains in the cuticle during wing movements.I wish to thank Mr. R. Whitty and staff for technical assistance with the scanning electron microscope and Dr. E.E. Ball for helpful discussion     

Life on the fly: phylogenetics and evolution of the helicopter damselflies (Odonata,Pseudostigmatidae)

Ingley, S.J., Bybee, S.M., Tennessen, K.J., Whiting, M.F. & Branham, M.A. (2012). Life on the fly: phylogenetics and evolution of the helicopter damselflies (Odonata, Pseudostigmatidae). —Zoologica Scripta, 41, 637–650. Helicopter damselflies (Odonata: Pseudostigmatidae) form a relatively small, yet highly specialized group of odonates, including the largest extant odonate (wingspan of ～190 mm). Pseudostigmatids are found throughout Central and South America, with the exception of one species that is found exclusively in East Africa. Pseudostigmatids oviposit exclusively in phytotelmata and forage on orb‐weaver spiders, which they pluck from webs. Pseudostigmatids also exhibit unique forms of both broad and narrow wings. Although the ecology of these behaviours and morphological features have been studied, their phylogenetic origins and evolutionary history are unknown. Here, we examine the origins of pseudostigmatid wing forms, oviposition in phytotelmata and spider feeding within a modern phylogenetic context, testing for single origins of each character. Phylogenetic analyses are based on 59 morphological characters and ～5 kb of sequence data. Our findings include a well‐supported monophyletic Pseudostigmatidae and Coryphagrion grandis as sister to the Neotropical genera. The genus Mecistogaster is paraphyletic, with Pseudostigma nested within the clade. The genus Microstigma is supported as monophyletic and forms a sister group relationship to the clade of Megaloprepus and Anomisma. The sister group relationship to Pseudostigmatidae is less clear. On the basis of this phylogenetic analysis, we propose three new tribes (Coryphagrionini, Microstigmatini and Mecistogastrini). As Pseudostigmatidae is monophyletic, the behaviour of gleaning spiders from webs appears to derive from a single origin. There are two origins of broad wings within Pseudostigmatidae. Oviposition in phytotelmata most certainly evolved multiple times within Coenagrionoidea. These findings provide new insights into pseudostigmatid evolution that can be used to generate hypotheses regarding behaviour and morphological adaptation in this unique and threatened group of damselflies.     

Phylogeny and taxonomy of Macrolepiota (Agaricaceae)

The position and composition of Macrolepiota within the Agaricaceae and its phylogenetic relationships with other members of the family were investigated, using both molecular (ITS and LSU rDNA sequences) and morphological characters. The molecular data separate the genus into two clades. The first clade comprises M. procera, M. mastoidea, M. clelandii and allies and is a sister group of Leucoagaricus and Leucocoprinus. The second, more diverse, clade, with M. rachodes and allies, M. globosa, Chlorophyllum molybdites, Leucoagaricus hortensis and Endoptychum agaricoides, is a sister group of Agaricus. The separation of the two clades is supported by morphological characters, such as the structure of the pileus covering, the stipitipellis and the shape of the germ pore and the spore apex. The two clades are regarded as genera for which the names Macrolepiota and Chlorophyllum are proposed. Macrolepiota nympharum does not belong to either clade but is assigned to the genus Leucoagaricus, close to L. leucothites. Endoptychum depressum is transferred to the genus Agaricus as A. inapertus.