Evolution of the dragonfly head-arresting system

The arrester or fixation system of the head in adult Odonata is unique among arthropods. This system involves the organs of two body segments: the head and the neck. It consists of a skeleton–muscle apparatus that sets the arrester parts in motion. The parts comprise formations covered with complicated microstructures: fields of microtrichia on the rear surface of the head and post-cervical sclerites of the neck. The arrester immobilizes the head during feeding or when the dragonfly is in tandem flight. Thus, it may serve as an adaptation to save the head from violent mechanical disturbance and to stabilize gaze in a variety of behavioural situations. This study shows the evolutionary trend of the arrester in the order Odonata by using scanning electron microscopy and measurements of arrester structures in 227 species from 26 odonate families. The arrester design occurring in the Epiophlebiidae, Gomphidae, Neopetaliidae, Petaluridae and Chlorogomphinae is suggested to be the basic one. Two convergent pathways of head-arrester evolution among Zygoptera and Anisoptera are proposed. The possible functional significance of the arrester system is discussed.     

Phylogenetic analysis of higher-level relationships of Odonata

Abstract. This is the most comprehensive analysis of higher‐level relationships in Odonata conducted thus far. The analysis was based on a detailed study of the skeletal morphology and wing venation of adults, complemented with a few larval characters, resulting in 122 phylogenetically informative characters. Eighty‐five genera from forty‐five currently recognized families and subfamilies were examined. In most cases, several species were chosen to serve as exemplars for a given genus. The seven fossil outgroup taxa included were exemplar genera from five successively more distant odonatoid orders and suborders: Tarsophlebiidae (the closest sister group of Odonata, previously placed as a family within ‘Anisozygoptera’), Archizygoptera, Protanisoptera, Protodonata and Geroptera. Parsimony analysis of the data, in which characters were treated both under equal weights and implied weighting, produced cladograms that were highly congruent, and in spite of considerable homoplasy in the odonate data, many groupings in the most parsimonious cladograms were well supported in all analyses, as indicated by Bremer support. The analyses supported the monophyly of both Anisoptera and Zygoptera, contrary to the well known hypothesis of zygopteran paraphyly. Within Zygoptera, two large sister clades were indicated, one comprised of the classical (Selysian) Calopterygoidea, except that Amphipterygidae, which have traditionally been placed as a calopterygoid family, nested within the other large zygopteran clade comprised of Fraser's ‘Lestinoidea’ plus ‘Coenagrionoidea’ (both of which were shown to be paraphyletic as currently defined). Philoganga alone appeared as the sister group to the rest of the Zygoptera in unweighted cladograms, whereas Philoganga + Diphlebia comprised the sister group to the remaining Zygoptera in all weighted cladograms. ‘Anisozygoptera’ was confirmed as a paraphyletic assemblage that forms a ‘grade’ towards the true Anisoptera, with Epiophlebia as the most basal taxon. Within Anisoptera, Petaluridae appeared as the sister group to other dragonflies.     

Molecules, morphology and fossils: a comprehensive approach to odonate phylogeny and the evolution of the odonate wing

We undertook a comprehensive morphological and molecular phylogenetic analysis of dragonfly phylogeny, examining both extant and fossil lineages in simultaneous analyses. The legitimacy of higher‐level family groups and the phylogenetic relationship between families were tested. Thirteen families were supported as monophyletic (Aeshnidae, Calopterygidae, Chlorocyphidae, Euphaeidae, Gomphidae, Isostictidae, Lestidae, Libellulidae, Petaluridae, Platystictidae, Polythoridae, Pseudostigmatidae and Synthemistidae) and eight as non‐monophyletic (Amphipterygidae, Coenagrionidae, Corduliidae, Megapodagrionidae, Protoneuridae and Synlestidae), although Perilestidae and Platycnemididae were recovered as monophyletic under Bayesian analyses. Nine families were represented by one species, thus monophyly was not tested (Epiophlebiidae, Austropetaliidae, Chlorogomphidae, Cordulegastridae, Macromiidae, Chorismagrionidae, Diphlebiidae, Lestoideidae and Pseudolestidae). Epiprocta and Zygoptera were recovered as monophyletic. Ditaxinerua is supported as the sister lineage to Odonata, Epiophlebiidae and the lestid‐like damselflies are sister to the Epiprocta and Zygoptera, respectively. Austropetaliidae + Aeshnidae is the sister lineage to the remaining Anisoptera. Tarsophlebia's placement as sister to Epiprocta or as sister to Epiprocta + Zygoptera was not resolved. Refinements are made to the current classification. Fossil taxa did not seem to provide signals crucial to recovering a robust phylogeny, but were critical to understanding the evolution of key morphological features associated with flight. Characters associated with wing structure were optimized revealing two wing character complexes: the pterostigma–nodal brace complex and the costal wing base & costal–ScP junction complex. In turn, these two complexes appear to be associated; the pterostigma–nodal brace complex allowing for further modification of the wing characters comprised within the costal wing base & costal–ScP junction complex leading the modern odonate wing. © The Willi Hennig Society 2008.     

Coding characters from different life stages for phylogenetic reconstruction: a case study on dragonfly adults and larvae,including a description of the larval head anatomy of Epiophlebia superstes (Odonata: Epiophlebiidae)

The exclusive use of characters coding for specific life stages may bias tree reconstruction. If characters from several life stages are coded, the type of coding becomes important. Here, we simulate the influence on tree reconstruction of morphological characters of Odonata larvae incorporated into a data matrix based on the adult body under different coding schemes. For testing purposes, our analysis is focused on a well‐supported hypothesis: the relationships of the suborders Zygoptera, ‘Anisozygoptera’, and Anisoptera. We studied the cephalic morphology of Epiophlebia, a key taxon among Odonata, and compared it with representatives of Zygoptera and Anisoptera in order to complement the data matrix. Odonate larvae are characterized by a peculiar morphology, such as the specific head form, mouthpart configuration, ridge configuration, cephalic musculature, and leg and gill morphology. Four coding strategies were used to incorporate the larval data: artificial coding (AC), treating larvae as independent terminal taxa; non‐multistate coding (NMC), preferring the adult life stage; multistate coding (MC); and coding larval and adult characters separately (SC) within the same taxon. As expected, larvae are ‘monophyletic’ in the AC strategy, but with anisopteran and zygopteran larvae as sister groups. Excluding larvae in the NMC approach leads to strong support for both monophyletic Odonata and Epiprocta, whereas MC erodes phylogenetic signal completely. This is an obvious result of the larval morphology leading to many multistate characters. SC results in the strongest support for Odonata, and Epiprocta receives the same support as with NMC. Our results show the deleterious effects of larval morphology on tree reconstruction when multistate coding is applied. Coding larval characters separately is still the best approach in a phylogenetic framework. © 2015 The Linnean Society of London     

Carotenoids of dragonflies,from the perspective of comparative biochemical and chemical ecological studies

Carotenoids of 20 species of dragonflies (including 14 species of Anisoptera and six species of Zygoptera) were investigated from the viewpoints of comparative biochemistry and chemical ecology. In larvae, β-carotene, β-cryptoxanthin, lutein, and fucoxanthin were found to be major carotenoids in both Anisoptera and Zygoptera. These carotenoids were assumed to have originated from aquatic insects, water fleas, tadpoles, and small fish, which dragonfly larvae feed on. Furthermore, β-caroten-2-ol and echinenone were also found in all species of larvae investigated. In adult dragonflies, β-carotene was found to be a major carotenoid along with lutein, zeaxanthin, β-caroten-2-ol, and echinenone in both Anisoptera and Zygoptera. On the other hand, unique carotenoids, β-zeacarotene, β,ψ-carotene (γ-carotene), torulene, β,γ-carotene, and γ,γ-carotene, were present in both Anisoptera and Zygoptera dragonflies. These carotenoids were not found in larvae. Food chain studies of dragonflies suggested that these carotenoids originated from aphids, and/or possibly from aphidophagous ladybird beetles and spiders, which dragonflies feed on. Lutein and zeaxanthin in adult dragonflies were also assumed to have originated from flying insects they feed on, such as flies, mosquitoes, butterflies, moths, and planthoppers, as well as spiders. β-Caroten-2-ol and echinenone were found in both dragonfly adults and larvae. They were assumed to be metabolites of β-carotene in dragonflies themselves. Carotenoids of dragonflies well reflect the food chain during their lifecycle.     

A molecular phylogeny of the Odonata (Insecta)

Abstract. We estimated the phylogeny of the order Odonata, based on sequences of the nuclear ribosomal genes 5.8 S, 18S, and ITS1 and 2. An 18S‐only analysis resolved deep relationships well: the order Odonata, as well as suborders Zygoptera and Epiprocta (Anisoptera + Epiophlebia), emerged as monophyletic. Some other deep clades resolved well, but support for more recently diverged clades was generally weak. A second, simultaneous, analysis of the 5.8S and 18S genes with the intergenic spacers ITS1 and 2 resolved some recent branches better, but appeared less reliable for deep clades with, for example, suborder Anisoptera emerging as paraphyletic and Epiophlebia superstes recovered as an Anisopteran, embedded within aeshnoid‐like anisopterans and sister to the cordulegastrids. Most existing family levels in the Anisoptera were confirmed as monophyletic clades in both analyses. However, within the corduliids that form a major monophyletic clade with the Libellulidae, several subclades were recovered, of which at least Macromiidae and Oxygastridae are accepted at the family level. In the Zygoptera, the situation is complex. The lestid‐like family groups (here called Lestomorpha) emerged as sister taxon to all other zygopterans, with Hemiphlebia sister to all other lestomorphs. Platystictidae formed a second monophylum, subordinated to lestomorphs. At the next level, some traditional clades were confirmed, but the tropical families Megapodagrionidae and Amphipterygidae were recovered as strongly polyphyletic, and tended to nest within the clade Caloptera, rendering it polyphyletic. Platycnemididae were also non‐monophyletic, with several representatives of uncertain placement. Coenagrionids were diphyletic. True Platycnemididae and non‐American Protoneurids are closely related, but their relationship to the other zygopterans remains obscure and needs more study. New World protoneurids appeared relatively unrelated to old world + Australian protoneurids. Several recent taxonomic changes at the genus level, based on morphology, were confirmed, but other morphology‐based taxonomies have misclassified taxa considered currently as Megapodagrionidae, Platycnemididae and Amphipterygidae and have underestimated the number of family‐level clades.     

Molecular phylogeny of the higher taxa of Odonata (Insecta) inferred from COI, 16S rRNA, 28S rRNA,and EF1‐α sequences

In this study, we sequenced both two mitochondrial genes (COI and 16S rRNA) and nuclear genes (28S rRNA and elongation factor‐1α) from 71 species of Odonata that represent 7 superfamilies in 3 suborders. Phylogenetic testing for each two concatenated gene sequences based on function (ribosomal vs protein‐coding genes) and origin (mitochondrial vs nuclear genes) proved limited resolution. Thus, four concatenated sequences were utilized to test the previous phylogenetic hypotheses of higher taxa of Odonata via Bayesian inference (BI) and maximum likelihood (ML) algorithms, along with the data partition by the BI method. As a result, three slightly different topologies were obtained, but the BI tree without partition was slightly better supported by the topological test. This topology supported the suborders Anisoptera and Zygoptera each being a monophyly, and the close relationship of Anisozygoptera to Anisoptera. All the families represented by multiple taxa in both Anisoptera and Zygoptera were consistently revealed to each be a monophyly with the highest nodal support. Unlike consistent and robust familial relationships in Zygoptera those of Anisoptera were partially unresolved, presenting the following relationships: ((((Libellulidae + Corduliidae) + Macromiidae) + Gomphidae + Aeshnidae) + Anisozygoptera) + (((Coenagrionidae + Platycnemdidae) + Calopterygidae) + Lestidae). The subfamily Sympetrinae, represented by three genera in the anisopteran family Libellulidae, was not monophyletic, dividing Crocothemis and Deielia in one group together with other subfamilies and Sympetrum in another independent group.     

蜻蜓目高级阶元系统发育关系研究进展

综述了对蜻蜓目系统发育的研究。对一些规模较大较全面的研究结果做了描述,并对其进行比较;对最近的基于分子的和基于形态的研究的共同点做了总结,其共同支持的观点包括:束翅亚目处于分支的基部为一并系类群,差翅亚目与间翅亚目关系较近。     

Temperature-body size responses in insects: a case study of British Odonata

1. Body size is highly correlated with physiological traits, fitness, and trophic interactions. These traits are subject to change if there are widespread reductions of body size with warming temperatures, which is suggested as one of the ‘universal’ ecological responses to climate change. However, general patterns of body size response to temperature in insects have not yet emerged. 2. To address this knowledge gap, we paired the wing length (as a proxy for body size) of 5331 museum specimens of 14 species of British Odonata with historical temperature data. Three sets of analyses were performed: (i) a regression analysis to test for a relationship between wing length and mean seasonal temperature within species and subsequent comparisons across species and suborders; (ii) an investigation of whether the body size of species has an effect on sensitivity to warming temperature; and (iii) a linear-mixed effects model to investigate factors that potentially affect temperature–size response. 3. The regression analysis indicated that wing length is negatively correlated with mean seasonal temperatures for Zygoptera, whereas Anisoptera showed no significant correlation with temperature. 4. There is a significant decline in wing length of all Zygoptera (but not Anisoptera) with collection date, suggesting that individuals emerging later in the season are smaller. 5. Life-cycle type was not important for predicting wing length–temperature responses, whereas sex, species, and suborder were indicated as important factors affecting the magnitude of temperature–size responses in Odonata. 6. Overall, wing lengths of Zygoptera are more sensitive to temperature and collection date than Anisoptera.     

Comparative morphology of the thorax musculature of adult Anisoptera (Insecta: Odonata): Functional aspects of the flight apparatus

Due to their unique flight mechanism including a direct flight musculature, Odonata show impressive flight skills. Several publications addressed the details of this flight apparatus like: sclerites, wings, musculature, and flight aerodynamics. However, 3D-analysis of the thorax musculature of adult dragonflies was not studied before and this paper allows for a detailed insight. We, therefore, focused on the thorax musculature of adult Anisoptera using micro-computed tomography. Herewith, we present a comparative morphological approach to identify differences within Anisoptera: Aeshnidae, Corduliidae, Gomphidae, and Libellulidae. In total, 54 muscles were identified: 16 prothoracic, 19 mesothoracic, and 19 metathoracic. Recorded differences were for example, the reduction of muscle Idlm4 and an additional muscle IIIdlm1 in Aeshna cyanea, previously described as rudimentary or missing. Muscle Iscm1, which was previously reported missing in all Odonata, was found in all investigated species. The attachment of muscle IIpcm2 in Pantala flavescens is interpreted as a probable adaption to its long-distance migration behaviour. Furthermore, we present a review of functions of the odonatan flight muscles, considering previous publications. The data herein set a basis for functional and biomechanical studies of the flight apparatus and will therefore lay the foundation for a better understanding of the odonatan flight.     

Neotropical dragonflies (Insecta: Odonata) as indicators of ecological condition of small streams in the eastern Amazon

Sensitive and cost‐effective indicators of aquatic ecosystem condition in Amazon streams are necessary to assess the effects of anthropogenic disturbances on those systems in a viable and ecologically meaningful manner. We conducted the present study in the municipality of Paragominas, state of Pará, northern Brazil, where we sampled adult dragonflies in 50 100‐m‐long wadeable stream sites in 2011. We collected 1769 specimens represented by 11 families, 41 genera and 97 species. The suborder Zygoptera contributed 961 individuals and Anisoptera 808. Among the 97 recorded species, nine were classified as useful indicators of ecological condition, with four species being associated with more degraded streams (three Anisoptera, one Zygoptera) and five with more preserved streams (all were Zygoptera). Anisoptera (dragonflies) tend to provide more useful indicators of more degraded environments because they have more efficient homeostatic mechanisms and are more mobile, enabling them to tolerate a wider range of environmental conditions. By contrast, Zygoptera (damselflies) tend to provide a more useful role as indicators of more preserved environments and high levels of environmental heterogeneity because of their smaller body sizes and home ranges and greater ecophysiological restrictions. We conclude from our assessment of this low‐order Amazonian stream system that (i) the occurrence of specific odonate species is strongly associated with the configuration of riparian vegetation, (ii) agricultural activities appear to be the main factor determining changes in the composition of odonate assemblages and (iii) these insects can act as useful indicators of the ecological consequences of riparian habitat loss and disturbance. Because generalist species invade moderately degraded areas, those areas may have high species richness but host few species of Zygoptera. Therefore, preserving dense riparian vegetation is necessary to maintain aquatic ecological condition, and that condition can be rehabilitated by planting new trees. Both require enforcing existing environmental regulations, various types of incentives and educating local communities.     

Wing length allometry in Odonata: differences between families in relation to migratory behaviour

In insects, wing shape and body size are correlated with several aspects of behaviour, and the optimal morphology of wings is a trade-off between a number of functional demands in relation to behaviour (e.g. foraging, migration and sexual display). Dragonflies are spectacularly skilful flyers and present a range of different wing shapes, but to date, no detailed studies have been conducted in this group on wing length allometry in relation to body size. In this paper, we use published data on body length and wing length in all European and North American dragonflies to investigate differences in wing length allometries among Odonata taxa (suborders and families) and to relate these to behavioural patterns. We found different wing allometries between Zygoptera and Anisoptera, which are probably related to the flight mode and wing form of the two suborders. Among the Anisoptera, the Libellulidae showed a distinct wing length allometry from all other anisopteran families and migrants differed from non-migrant species. The first dichotomy is likely to reflect the adaptation of wing morphology of Libellulidae to sit-and-wait behaviour and to brief foraging flights (most species of this family are perchers) with respect to all other families, members of which are typically flyers. The second dichotomy reflects the trend of migrating species to have relatively longer wings than non-migrating members of the same family. Finally, wing length allometry differed among all the zygopteran families analysed, and this pattern suggested that each family evolved a particular wing morphology in response to peculiarities in behaviour, habitat and flight mode.     

Do different dispersal capacities influence the biogeography of the western Palearctic dragonflies (Odonata)?

The biogeography of the western Palearctic has been intensively studied for more than a century. Recent advances in genetics have allowed the testing of old theories based on distribution patterns, although these analyses are obviously restricted to a reduced number of specific genetic data sets. On the other hand, an increased knowledge on the distributions of species and advances in computer capacities have allowed more detailed biogeographical analyses based on species presence/absence. In the present study, we selected the Odonata as the study group. For all 162 species native to the western Palearctic, we compiled their respective presence or absence in 97 predefined biogeographical regions. Using cluster analyses and principal component analyses, both based on Jaccard similarity coefficients, we analysed the differentiation among these regions and species. In subsequent analyses, the data set was reduced to the Zygoptera, Anisoptera, and the western Palearctic endemics. All analyses consistently showed different faunal regions and faunal elements. In particular, the (1) western and (2) eastern Mediterranean; (3) Central and (4) Northern Europe; and (5) the British Isles were invariably found in all cases. Although the two major Mediterranean regions were characterized by several endemic faunal elements, Northern Europe and the British Isles lacked such elements, but were characterized by faunal compositions strongly deviating from the rest of the western Palearctic region. Moderate differences between Zygoptera and Anisoptera existed, with the latter more clearly redrawing the Mediterranean refuge areas, whereas the former reflected to a greater extent the postglacial expansion patterns from these regions. In general, our findings underline the old biogeographical theories, but refine especially our understanding of the Atlanto‐ and Ponto‐Mediterranean region. Central Europe, comprising the area with the highest species numbers of our whole study region, unravels as a crossroad of postglacial immigrations, but might also represent a region of in situ glacial survival. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 177–195.     

Interactions among coexisting larval Odonata: an in situ experiment using small enclosures

Field experiments using small replicated enclosures focused on interactions between larval populations of Epitheca cynosura and Ladona deplanata (Odonata: Anisoptera) — two species that emerge in early spring. The presence of Epitheca reduced the total biomass of Ladona, but Ladona had no significant effect on Epitheca. These early-emerging species reduced the biomass of small instars of late-emerging Anisoptera which colonized enclosures during the experiments; and the late-emerging Anisoptera seem to have inhibited colonization by Zygoptera larvae. Results are consistent with the importance of predatory (cannibalism or mutual predation) interactions in this community.     

Patterns of sexual dimorphism in flight agility in territorial and non-territorial Odonata

Journal of Ethology - Adult dragonflies (Anisoptera) and damselflies (Zygoptera) are amongst the most accomplished flying insects on the planet. The main functions of spatial displacement by flight...     

THE ORGANIZATION OF FLIGHT MUSCLE FIBERS IN THE ODONATA

The cytological organization of flight muscle fibers of Odonata has been investigated. These fibers, in representatives of the Zygoptera and Anisoptera, have been compared and found to be similar, except that, in the former, pairs of lamellar fibrils, rather than single fibrils, alternate with the mitochondria. In each instance, in these synchronous muscles, the actin filaments of the myofibrils are found to lie opposite to and midway between pairs of myosin filaments—a configuration previously reported in asynchronous flight muscle fibers. The disposition of the T system and sarcoplasmic reticulum membranes in glutaraldehyde-fixed anisopteran muscle is described in detail: the T system tubules are shown to be radially continuous across the fiber, and are derived as openmouthed invaginations from the surface cell-membrane. The detailed organization of the dyad junctions between these tubules and the adjoining cisternae of the sarcoplasmic reticulum is described. The accessibility of the T system interior to diffusion exchange with the general extracellular milieu has been investigated by studies on the penetration of ferritin into the fiber: molecules of this marker have been found to diffuse solely along the T system tubules, and their presence in the tubule extremities adjoining the centrally placed nuclei confirms the morphological evidence suggesting that these tubules provide open diffusion channels extending across the radius of the fiber. The possible physiological role of these membrane components and their distribution in synchronous muscles of insects and vertebrates and in asynchronous insect flight muscle are discussed.     

Odonata larvae of Kaliningrad oblast: Species composition and distributional features

In this study, a total of 27 species of dragonfly and damselfly larvae were identified, including 14 species from suborder Anisoptera and 13 from Zygoptera. The dominant species are Libellula quadrimaculata, Enallagma cyathigerum, and Erythromma najas. Frequently occurring species are Coenagrion hastulatum, C. lunulatum, C. pulchellum, Ischnura elegans, Platycnemis pennipes, Lestes virens, and Libellula depressa. Rare species are Lestes sponsa, Libellula fulva, Gomphus flavipes, Gomphus vulgatissimus, Sympetrum flaveolum, Aeshna viridis, Ae. cyanea, Ae. grandis, Ae. mixta, Calopteryx splendens, C. virgo, and Ophiogomphus cecilia.     

The sense of smell in Odonata: An electrophysiological screening

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.     

广西大明山蜻蜓调查

2011年8月对广西大明山的蜻蜓物种资源进行了调查。结果表明广西大明山蜻蜓种类隶属2亚目11科46种,差翅亚目Anisoptera4科27种,束翅亚目Zygoptera7科19种。其中广西新纪录种8种(亚种)。     

Insect predation on pike fry

Laboratory tests evaluated the predatory impact of the macroinvertebrates Erythromma najas larvae (Odonata, Zygoptera: Coenagrionidae), Notonecta glauca (Heteroptera: Notonectidae), Ilyocoris cimicoides (Heteroptera: Naucoridae), Libellula depressa larvae (Odonata, Anisoptera: Libellulidae), Dytiscus marginalis larvae (Coleoptera: Dytiscidae) and Anax imperator larvae (Odonata, Anisoptera: Aeshnidae) on 3-, 12-, 21- and 30-day-old pike fry Esox lucius . All these insect predators captured and ate pike fry during the test, although the numbers killed varied among species. Dytiscus marginalis, Anax imperator and Notonecta glauca were the most Voracious predators.