Phylogeography of a facultatively migratory dragonfly, Libellula quadrimaculata (Odonata: Anisoptera)

The biogeography of a widely distributed dragonfly (Libellula quadrimaculata) was examined via a phylogenetic analysis of 416 bp of the mitochondrial cytochrome oxidase I subunit (COI). Phylogenetic analyses under parsimony and minimum evolution produced trees with similar topologies, and revealed strong support for three clades corresponding to populations in Asia, Europe and North America. However, resolution was poor within clades, and genetic distances between populations within continents was quite low (1–2%). Several populations of this species are known to engage in periodic mass migrations, and it is possible that introgression from gene flow due to the mobility of this species has obscured phylogenetic patterns within continents. I was unable to test for phylogenetic patterns coincident with historical glacial refugia given the lack of phylogenetic patterns within continents. However, given that some sequence divergence was observed between populations within continents, it is possible that phylogenetic patterns exist, and subsequent studies should make use of larger data sets, and molecular data from faster evolving genes. Despite the propensity for periodic, short distance migrations in L. quadrimaculata, gene flow appears to be limited and does not influence the phylogenetic relationships of populations between continents.     

Potential distribution of the critically endangered dragonfly Libellula angelina (Odonata: Libellulidae) under shared socio-economic pathways

Libellula angelina is an endangered dragonfly species that is native to East Asia. Recently, their population has become severely reduced through habitat loss. To protect L. angelina populations, we need to understand which factors determine their distribution and how their potential habitats will change in the future. In this study, the habitat preference of L. angelina was identified through field surveys, and the potential distribution of L. angelina and the impact of integrated climate–land cover changes were simulated using the MaxEnt model. Furthermore, the wetland loss scenario was applied to areas where the current trend in wetland loss will continue in the future. The field survey identified that L. angelina prefers small inland wetlands: permanent freshwater, ponds; permanent rivers, ponds; irrigated land; and estuarine waters. From the MaxEnt results, altitude was the variable with the greatest contribution and distance from wetlands was the most unique variable. MaxEnt described the geographic pattern of occurrences under the current climate well, with few areas requiring any further survey. In the future projection, the potential habitat area was increased by up to 48.8% and 30.6% in the 2050s and 2080s, respectively. However, potential habitat loss was expected if wetlands continue to decline as they have done in the last 20 years. The wetland loss scenario resulted in potential habitat losses of 1.9%–2.3% and 4.5%–6.1% in the 2050s and 2080s, respectively. Therefore, to protect L. angelina populations we must minimize the loss of current populations, secure wetlands and strengthen the connectivity between wetlands.     

Biased dispersal of Metrioptera bicolor,a wing dimorphic bush‐cricket

In the highly fragmented landscape of central Europe, dispersal is of particular importance as it determines the long‐term survival of animal populations. Dispersal not only secures the recolonization of patches where populations went extinct, it may also rescue small populations and thus prevent local extinction events. As dispersal involves different individual fitness costs, the decision to disperse should not be random but context‐dependent and often will be biased toward a certain group of individuals (e.g., sex‐ and wing morph‐biased dispersal). Although biased dispersal has far‐reaching consequences for animal populations, immediate studies of sex‐ and wing morph‐biased dispersal in orthopterans are very rare. Here, we used a combined approach of morphological and genetic analyses to investigate biased dispersal of Metrioptera bicolor, a wing dimorphic bush‐cricket. Our results clearly show wing morph‐biased dispersal for both sexes of M. bicolor. In addition, we found sex‐biased dispersal for macropterous individuals, but not for micropters. Both, morphological and genetic data, favor macropterous males as dispersal unit of this bush‐cricket species. To get an idea of the flight ability of M. bicolor, we compared our morphological data with that of Locusta migratoria and Schistocerca gregaria, which are very good flyers. Based on our morphological data, we suggest a good flight ability for macropters of M. bicolor, although flying individuals of this species are seldom observed.     

Larval morphology and range of three West Asiatic species of the genus Onychogomphus Selys, 1854. (Insecta: Odonata)

The previously unknown larvae of O. assimilis (Schneider, 1845) and O. lefebvrei (Rambur, 1842) are described, while that of O. flexuosus (Schneider, 1845) is redescribed, all from material from SW. Turkmenistan and Tajikistan. Their distributions extend from the eastern mediterranean to central-west Asia and are mapped. A differential diagnosis and key to the identification of the larvae of four central Asiatic species of the genus Onychogomphus is given.     

The development of dragonfly larvae (Odonata: Anisoptera) from two streams in north-eastern Victoria, Australia

Developmental stages of larvae of eight species of Anisoptera from the Kiewa River and Middle Creek, Victoria, were determined and their phenology investigated. The last six to nine instars of each species were distinguished by size frequency and scatter plots, using labium width, metafemur and wing-pad measurements, and the early instars were estimated from Dyar's Law. This suggested between 11 to 14 instars. Four species appeared to be univoltine and four species semivoltine.     

Micro-morphological adaptations of the wing nodus to flight behaviour in four dragonfly species from the family Libellulidae (Odonata: Anisoptera)

Adult dragonflies can be divided into two major groups, perchers and fliers, exhibiting notably different flight behaviour. Previous studies have yielded conflicting results regarding the link between the wing macro-morphology and flight style in these two groups. In this study, we present the first systematic investigation of the micro-morphological differences of wings of percher and flier dragonflies in four closely related species from the family Libellulidae. Our results suggest that the shape and material composition of wing microstructural components and, in particular, the nodus are adapted to facilitate the specific wing functioning in fliers and perchers. The findings further indicate a decreasing trend in the area proportion of the soft resilin-dominated cuticle in the nodus in the series of species from typical perchers to typical fliers. Such a reduction in the resilin proportion in the nodus of fliers is associated with an increase in the wing aspect ratio. The knot-shaped protrusion at the nodus of perchers, which becomes notably smaller in that of strong fliers, is likely to act as a mechanical stopper, avoiding large wing displacements. This study aims to develop a novel framework for future research on the relationship between wing morphology and flight behaviour in dragonflies.     

Ontogenetic shifts in functional morphology of dragonfly legs (Odonata: Anisoptera)

Anisopteran leg functions change dramatically from the final larval stadium to the adult. Larvae use legs mainly for locomotion, walking, climbing, clinging, or burrowing. Adults use them for foraging and grasping mates, for perching, clinging to the vegetation, and for repelling rivals. In order to estimate the ontogenetic shift in the leg construction from the larva to the adult, this study quantitatively compared lengths of fore, mid, and hind legs and the relationships between three leg segments, femur, tibia, and tarsus, in larval and adult Anisoptera of the families Gomphidae, Aeshnidae, Cordulegastridae, Corduliidae, and Libellulidae, represented by two species each. We found that leg segment length ratio as well as ontogenetic shift in length ratios was different between families, but rather similar within the families. While little ontogenetic shift occurred in Aeshnidae, there were some modifications in Corduliidae and Libellulidae. The severest shift occurred in Gomphidae and Cordulegastridae, both having burrowing larvae. These two families form a cluster, which is in contrast to their taxonomic relationship within the Anisoptera. Cluster analysis implies that the function of larval legs is primarily responsible for grouping, whereas adult behavior or the taxonomic relationships do not explain the grouping. This result supports the previous hypothesis about the convergent functional shift of leg characters in the dragonfly ontogenesis.     

The Larvae of the Mesozoic family Aeschnidiidae and their Phylogenetic Implications (Insecta, Odonata, Anisoptera)

Four giant dragonfly larvae are described from the Lower Cretaceous of China. Owing to the preservation of wing tracheal venation on the larval wing sheaths, they can be identified as the first undoubted larvae of the extinct Mesozoic family Aeschnidiidae. They are ultimate or penultimate male and female specimens, and a younger larva. The female larva has a very long ovipositor sheath. These larvae have an anisopteran anal pyramid and a very particular spoon-shaped labial mask, with a very narrow prementum and large palps with numerous teeth, suggesting possible affinities of the Aeschnidiidae with the Anisoptera Cavilabiata. The positions of other larvae formally attributed to the Aeschnidiidae are discussed, i.e. Nothomacromia sensibilis (Carle and Wighton, 1990), Sona nectes Pritykina, 1986, and the alleged larvae of Hemeroscopus baissicus Pritykina, 1977. They differ greatly from the true Chinese larval Aeschnidiidae, in the labial mask and female ovipositor, even if they show some similarities in the anal pyramid.     

The wing morphology of limnephilid caddisflies in relation to their habitat preferences

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Relationship between exposure to an insecticide and fluctuating asymmetry in a damselfly (Odonata,Coenagriidae)

In this study, we explored the effects of pesticide on fluctuating asymmetry (FA) levels and mortality of Ceriagrion sp. larvae. The results showed that the mortality of larval damselflies treated with pesticide was significantly higher than that treated with tap water which had been aerated for 48 h, but there were no significant differences among mortality of larvae treated with different concentrations of pesticide. Meanwhile, we found that the level of FA of the first tibia length, one of the seven bilaterally symmetrical traits (First femur length, First tibia length, Second femur length, Second tibia length, Third femur length, Third tibia length and Prementum width), differed significantly with different treatments, whereas the others did not show any significant differences. The Bonferroni (Dunn) t Tests revealed that FA of the first tibia length at 15 × 10⁻⁹ mgl⁻¹ was significantly higher than that at 1.5 × 10⁻⁹ mgl⁻¹ and control. There was no significant relationship between trait size and the absolute difference between their right and left sides. There was also no significant relationship between body size and the absolute difference between right and left sides. Trait size was significantly positively correlated with body size. FA was not associated with mortality. Therefore, we concluded that FA of the first tibia length of Ceriagrion sp. larvae may be induced by sublethal doses of pesticides. That is to say, its FA may be regarded as an indicator of reflecting the level of pesticide stress. Handling editor: K. Martens     

Slow, fast and in between: habitat distribution and behaviour of larvae in nine species of libellulid dragonfly

1. Activity and microhabitat use are important factors determining species performance in habitats that differ in permanence and species composition of top predators. This study examined the relationship between the distribution across a gradient of habitat permanence and an associated transition in the composition of top predators and the behaviour of species of larval dragonflies. It also assessed the relationship between larval behaviour, body size and the duration of the larval stage. In laboratory mesocosms the mobility of the different species was measured, as was the extent to which they associated with artificial vegetation. 2. Species mobility was positively related to their natural occurrence in habitats in which invertebrates or small‐bodied fish were the top predators, and negatively related with the frequency with which species co‐existed with large‐bodied fish, the permanence of the habitat and the length of the larval stage. 3. Rather than falling into strict low and high mobility categories, habitat generalists that occurred across the habitat gradient, co‐existing with different top predators, had variable mobility levels. In these generalists, mobility was positively related to how frequently they were found in natural habitats in which invertebrates were the top predators. 4. The extent to which species utilized the artificial vegetation in mesocosms was associated with the length of the larval period but was not associated with mobility or species habitat distribution in the field.     

Postcopulatory Behavior in Libellula pulchella Drury (Odonata: Libellulidae) and Female Tactics for Avoiding Male Interference with Oviposition

Postcopulatory behavior was studied in Libellula pulchella, a North American dragonfly in which ovipositing females face frequent harassment by unpaired males seeking matings. Although males performed noncontact guarding of their mates after copulation, females received minimal protection since their guarders tended to leave on extended chases of other males when harassment was intense. Ovipositions by unguarded females were even more likely to be terminated by harassment and were disrupted sooner. Female tactics to minimize interference included rapid escape flights, repeated return visits to the water within short time periods, perching when severerly harassed, and proceeding with mating when clasped. Female use of multiple oviposition sites is discussed in the context of guarding effectiveness and mate recognition by males.     

A panel of microsatellite markers to study sperm precedence patterns in the emperor dragonfly Anax imperator (Odonata: Anisoptera)

Odonates were the first group of organisms where sperm competition and last male sperm precedence have been identified. With the development of 10 microsatellites for the emperor dragonfly Anax imperator, the function and priority patterns of the multiple sperm storage organs of females can be studied and compared between species in natural populations. In addition, two microsatellite loci developed for the sister species Anax parthenope, are also highly polymorphic in A. imperator. For the presented 12 microsatellite loci, the number of alleles per locus ranged from two to 24. Observed heterozygosity ranged from 0.07 to 0.88.     

Hybridization between Ischnura graellsii (Vander Linder) and 2. elegans (Rambur) (Odonata: Coenagrionidae): are they different species?

Two closely related damselflies, Ischnura graellsii and I. elegans , were analysed for morphological differences and reproductive isolation in the north coast of Galicia (NW Spain). We compared animals from sympatric and allopatric localities, including I. elegans from Belgium and I. graellsii from southern Spain as pure allopatric populations. A set of morphometric characters were studied by means of multivariate discriminant analysis to determine if these two species can be unambiguously distinguished. Discriminant analysis revealed that I. graellsii and I. elegans are well differentiated on the first two axis (86% and 11%, respectively). I. graellsii individuals are distinguished from I. elegans by their smaller size and, specifically, by their narrower and shorter wings and shorter tibiae. In addition, I. elegans has a narrower space between the branches of each cercus, and greater distance between the branches of each paraproct. Sympatric individuals are morphologically intermediate, suggesting hybridization. When the species were put together in the laboratory, they showed partial temporal separation in mating behaviour, but males of I. elegans readily mated with females of I. graellsii , and hybrid individuals were obtained. The opposite heterospecific cross was almost impossible, apparently because of mechanical problems with the tandem linkage. Laboratory-reared hybrids (from male I. elegans × female I. graellsii ) are morphologically intermediate, mainly resembling the maternal phenotype. Although hybridization between both taxa is common, we suggest maintaining the specific status for both phenotypes because they show incipient reproductive isolation, as it is reported in the literature. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 225–235.     

Wing morphology and migration status,but not body size,habitat or Rapoport's rule predict range size in North‐American dragonflies (Odonata: Libellulidae)

Understanding why species range sizes vary is important for predicting the impact of environmental change on biodiversity. Here we use a multi‐variable approach in a phylogenetic comparative context to understand how four morphological, two ecological, and two eco‐geographical variables are associated with range size, latitudinal range and longitudinal range in 81 species of North‐American libellulid dragonflies. Our results show that: 1) migratory species and species with a more expanded basal hindwing lobe have a larger range size; 2) opposite to Rapoport's rule, latitudinal range is negatively correlated with mid‐range latitude; 3) longitudinal range is predicted by wing morphology and migration; 4) body size and larval habitat are not correlated with range size, latitudinal range or longitudinal range. These results suggest that dispersal‐related traits, such as wing shape and migratory status, are important factors in predicting the range size of libellulid dragonflies. In addition, the reverse Rapoport's rule suggests that more northern‐centred species might be more specialized than more southern‐centred species. We suggest that the variables predicting range size are likely imposed by taxon‐specific morphological, ecological, physiological and behavioural traits. Taxon‐specific knowledge is thus necessary to understand the dynamics of range sizes and is important to implement successful restoration and conservation plans of threatened species.     

Adult habitat preferences, larval dispersal, and the comparative phylogeography of three Atlantic surgeonfishes (Teleostei: Acanthuridae)

Although many reef fishes of the tropical Atlantic are widely distributed, there are large discontinuities that may strongly influence phylogeographical patterns. The freshwater outflow of the Amazon basin is recognized as a major barrier that produces a break between Brazilian and Caribbean faunas. The vast oceanic distances between Brazil and the mid-Atlantic ridge islands represent another formidable barrier. To assess the relative importance of these barriers, we compared a fragment of the mitochondrial DNA (mtDNA) cytochrome b gene among populations of three species of Atlantic surgeonfishes: Acanthurus bahianus, A. chirurgus and A. coeruleus. These species have similar life histories but different adult habitat preferences. The mtDNA data show no population structure between Brazil and the mid-Atlantic islands, indicating that this oceanic barrier is readily traversed by the pelagic larval stage of all three surgeonfishes, which spend approximately 45-70 days in the pelagic environment. The Amazon is a strong barrier to dispersal of A. bahianus (d = 0.024, phiST = 0.724), a modest barrier for A. coeruleus (phiST = 0.356), and has no discernible effect as a barrier for A. chirurgus. The later species has been collected on soft bottoms with sponge habitats under the Amazon outflow, indicating that relaxed adult habitat requirements enable it to readily cross that barrier. A limited ability to use soft bottom habitats may also explain the low (but significant) population structure in A. coeruleus. In contrast, A. bahianus has not been collected over deep sponge bottoms, and rarely settles outside shallow reefs. Overall, adult habitat preferences seem to be the factor that differentiates phylogeographical patterns in these reef-associated species.     

福建狭翅蟌属一新种(蜻蜓目:蟌科)

记述采自福建省华安县的狭翅蟌属一新种华安狭翅蟌（Aciagrion huaanensis Xu, sp.nov.）,模式标本存放于福建省漳州教育学院.     

The larva of Libellula foliata (Kirby, 1889) (Odonata: Libellulidae)

The larva of Libellula foliata (Kirby) is described based upon mature larvae from the Biosphere Reserve of ??El Triunfo?? in the state of Chiapas. It belongs to the small group of species without dorsal protuberances, L. composita (Hagen), L. comanche Calvert and L. saturata Uhler. The following combination of characters permits the separation of L. foliata larva from the other aforementioned larvae: tergites 6?C10 uniformly colored, no lateral spines on segments 8?C9, 5?C6 palpal setae and 3 long premental setae. After this finding, only the larvae of Libellula gaigei Gloyd and L. nodisticta Hagen remain undiscovered for the Mexican species of Libellula.     

The antennal lobe of Libellula depressa (Odonata,Libellulidae)

Here we describe the antennal lobe of Libellula depressa (Odonata, Libellulidae), identified on the basis of the projections of the afferent sensory neurons stemming from the antennal flagellum sensilla. Immunohistochemical neuropil staining as well as antennal backfills revealed sensory neuron terminal arborizations covering a large portion of the antennal lobe. No clear glomerular structure was identified, thus suggesting an aglomerular antennal lobe condition as previously reported in Palaeoptera. The terminal arbors of backfilled sensory neurons do, however, form spherical knots, probably representing the connections between the few afferent neurons and the antennal lobe interneurons. The reconstruction revealed that the proximal part of the antennal nerve is divided into two branches that innervate two spatially separated areas of the antennal lobe, an anterioventral lobe and a larger posteriodorsal lobe. Our data are consistent with the hypothesis that one tract of the antennal nerve of L. depressa contains olfactory sensory neurons projecting into one of the sublobes, while the other tract contains thermo-hygroreceptive neurons projecting into the other sublobe.