The copulatory complex of Hemianax ephippiger (Burmeister) (Aeshnidae: Anisoptera)

Like other Anisoptera Hemianax ephippiger (Burmeister) bears two sets of copulatory apparatuses. The first set which is primary in nature is located on the ventral surface of the ninth abdominal segment and consists of the vestigial penis, the gonocoxites, the supra-anal appendages, the infra-anal appendage. The supra-anal appendages are well developed, lanceolate with serrate margin and spinous apices. They are dorso-ventrally flattened and furnished with long hairs and black teeth-like tubercles (T). The secondary copulatory complex is lodged inside the membranous fossa located on the ventral surface of second and third abdominal segments. It consists of anterior lamina, the posterior lamina, the supporting framework, the hamules, the penis sheath, the penis vesicle and the penis. The anterior lamina is deeply cleft at the posterior margin to accommodate the ovipositor of the female during copulation. Both the portions of the supporting framework are well developed. The anterior portion is somewhat rectangular while the posterior portion is nearly U-shaped. The anterior hamules are well developed with truncated and hooked portions, while the posterior hamules are very much reduced. The penis sheath is forcepate with well developed rectangular flap which hangs over the penis. The penis is stout, clearly demarcated into three segments, and bears a prominent groove on the mid-ventral line.     

Counting the spots: a molecular and morphological phylogeny of the spotted darner Boyeria (Odonata: Anisoptera: Aeshnidae) with an emphasis on European taxa

Boyeria irene and Boyeria cretensis are species of spotted dragonflies belonging to the ‘darner’ family, Aeshnidae. In 1991, Peters classified Boyeria from Crete as B. cretensis, based on adult morphological characters. In this study, we used molecular evidence to determine if indeed B. irene and B. cretensis are different species. DNA was sequenced from samples of B. irene (from France, Switzerland, Tunisia, Spain and Italy) and B. cretensis (from Crete). These species were recovered as two different clades with strong support. We conclude that B. irene and B. cretensis are different species, with evidence based on molecular and morphological differences. In addition, we present the first phylogenetic hypothesis for Boyeria for which we have sequenced all but three species. Lastly, we discuss different scenarios that may have led to the present‐day distribution and speciation patterns of Mediterranean Boyeria.     

A new Mesozoic Chinese genus of aeshnopteran dragonflies (Odonata: Anisoptera: Progobiaeshnidae)

Mongoliaeshna sinica gen. et sp. n., third record of the Mesozoic aeshnopteran family Progobiaeshnidae is described from the Lower Cretaceous of Yixian Formation in Liutiaogou (Ningcheng County, Inner Mongolia, China).     

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.     

福建头蜓属一新种(蜻蜓目:蜓科)

记述采自福建省邵武市的头蜓属1新种,邵武头蜓Cephalaeschnashaowuensis,sp.nov.。模式标本存放在漳州教育学院。     

Molecular phylogeny of the Western Palaearctic Cordulegaster taxa (Odonata: Anisoptera: Cordulegastridae)

Although Odonata are a key component of many freshwater ecosystems, their taxonomy and evolutionary history is still far from being well resolved. In the present study, we report the first molecular phylogeny for the Western Palaearctic Cordulegaster genus (Odonata: Anisoptera: Cordulegastridae). We sequenced fragments of both mitochondrial and nuclear genes [cytochrome c oxidase I (COI) and Internal Transcribed Spacer‐1 (ITS‐1)] from eight species and 13 subspecies, from western, southern and central Europe, Turkey, and Morocco. Our data support the existence of two major groups corresponding to the traditional boltonii‐ and bidentata‐groups. Both groups are monophyletic based on COI sequences and the distinctiveness of Cordulegaster princeps, Cordulegaster trinacriae, Cordulegaster picta and Cordulegaster heros relative to Cordulegaster boltonii, and Cordulegaster helladica and Cordulegaster insignis relative to Cordulegaster bidentata, is confirmed. All species are also monophyletic for ITS‐1, with the exception of Cordulegaster helladica buchholzi, which shares the haplotype with C. insignis. Although moderate levels of genetic diversity were found within C. boltonii, there was no clear separation among the four subspecies, with the exception of the populations of Cordulegaster boltonii algirica from North Africa. Similarly, no genetic differentiation was found between the two subspecies of C. bidentata, Cordulegaster bidentata bidentata and Cordulegaster bidenta sicilica. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111, 49–57.     

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.     

A nonoccluded virus in nymphs of the dragonfly Leucorrhinia dubia (Odonata,Anisoptera)

This work establishes the sequence of events as the water mold, Coelomomyces psorophorae, develops and enters mosquito larvae through the cuticles of these insects. The fungal cyst shows a number of adaptations for a parasitic function. A bulb-shaped appressorium is produced at germination which is initiated and/or maintained by a microtubular skeleton. The tip of the appressorium secretes a dense amorphous substance, then produces a narrow penetration tube which grows through the larval cuticle. The penetration tube maintains its narrow diameter as it grows inside the host epidermal cell. The parasite protoplasm is injected into a host epidermal cell. The protoplast is squeezed through the narrow tube by the (probably rapid) expansion of a vacuole in the cyst body at the end distal to the penetration point. There is a correlation of cuticular texture with adhesion patterns of cysts. A cuticular collar is always seen in successful penetrations. The sequence of development after attachment of the fungal zygotes is: germination between 1 and 2 hr; penetration between 3 and 4 hr; and injection between 6.5 and 8 hr.     

A remarkable new genus and species of dragonfly (Odonata: Anisoptera: Libellulidae) from Brazil and notes on its bionomics and phylogenetic affinities

Orionothemis felixorioni n. gen., n. sp. from Bahia state, Brazil, is described and illustrated from larvae, reared adults and an immature adult male taken in association with its possible larval shuck. This taxon exhibits remarkable features among the Odonata, such as enormous dorsal and lateral spines perpendicular to the body axis, totally fused last abdominal segments in the larva, strongly differentiated and sexually dimorphic posterior legs, and the incompletely chitinized eighth abdominal tergite of the adult. Orionothemis is closely related to Elasmothemis (Neotropical) and Onychothemis (South-East Asia). The larvae were collected in the abundant immersed vegetation in a clear and cool stream in the Brazilian ‘planalto’ (central plateau) in an area that is endangered by deforestation and irrigation. Better knowledge of the libellulid or more generally odonata larvae could help taxonomy and phylogeny of the group.     

Predatory versatility in a larval dragonfly, Hemianax papuensis (Odonata: Aeshnidae)

Larvae of the large dragonfly Hemianax papuensis used four disparate, prey-specific predatory behaviours. Arthropod prey moving on a substratum were stalked and then attacked from a distance. Arthropod prey moving in the water column or at the water surface were approached using 'jet' propulsion and then attacked from a distance. Snails, an unusual prey for an arthropod, were stalked; then the larva manoeuvred about them until a specific orientation was achieved before an attack was made from close range. Dead snails were scavenged, using tactics very similar to those used with live snails, but non-snail carrion was rarely taken. There was no evidence that the possession, by H. papuensis , of specialized behaviours for an atypical prey-snails-lowered its success when attacking other types of 'typical' prey.     

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.     

Diversity of Anisoptera (Odonata): infering speciation processes from patterns of morphological diversity

With roughly 2500 described species Anisoptera are among the species-poor suborders within insects. However, morphological and ecological variability are truly impressive. Anisoptera are classified into about 15 families of variable species richness. In this analysis phylogenetic research is integrated with comparative approaches to investigate possible explanations of differential speciation rates within this suborder. A short review of phylogenetic work based on morphological characters is compared to published molecular phylogenies. Sistergroup comparisons are used to elucidate whether a) sexual selection, b) duration of life cycles, or c) differentiation in body size, have had a detectable effect on speciation rate. In all three analyses effects of distributional range and latitudinal distribution were controlled. These analyses suggest sexual selection promotes speciation and an increase in body size is positively correlated with speciation rate. The evolutionary significance of these results is discussed and experimental approaches that should advance our understanding of anisopteran diversity are suggested.     

Ventilatory movements of the abdomen and branchial apparatus in dragonfly larvae (Odonata: Anisoptera)

Use of cine-photography in the analysis of ventilatory movements has necessitated a reappraisal of the ventilatory patterns previously described in dragonfly larvae. Movements of the abdominal exoskeleton, monitored by cine-photography, phototransistor and strain gauge, are discussed in relation to known muscular activity. Internal movements of the branchial apparatus, which consists of two chambers and three valves, are also illustrated in the intact animal. The mechanism of "gulping" ventilation ( V _a) in dragonfly larvae is discussed.     

Functional anatomy of the secondary copulatory apparatus of the male dragonfly Tramea virginia (Odonata: Anisoptera)

In the dragonfly Tramea virginia, the secondary copularoty apparatus (SCA) on the venter of the second and third abdominal segments consists of the anterior and posterior laminae, a pair of gential lobes and hamules, the ligula, genital fossa, supporting frame work, and the four segmented, highly specialized penis. The penis head represnts the distal or fourth segment of the penis and bears paired lateral and apical lobes and a medial process. The medial process includes a sperm reservior, a short sperm tube, and paired cornua and inner lobes. The sperm reservoir opens into the sperm tube which bears a terminal orifice between the inner lobes. Gross morphology of the SCA of T. virginia is similar to that of other libellulids except that the gential lobes and hamules are comparatively large in size, the ligula is triangular in shape with an apical flat rectangular lobe and stiff basal setae, the penis bears a pair of cuticular denticles below the spur of the second segment, the apical lobes are inflatable without spines and bristles, and the cornua with smooth outer and spiny inner surface collectively signifying the species-specific anatomical features and the functional specialization for sperm transfer and removal. © 1993 Wiley-Liss, Inc.     

The mechanoreceptors on the endophytic ovipositor of the dragonfly Aeshna cyanea (Odonata,Aeshnidae)

This study investigates the mechanoreceptors located on the cutting valvulae of the ovipositor of the dragonfly Aeshna cyanea (Aeshnidae), using both SEM and TEM, with the aim of providing an overview of the sensory equipment of an odonatan endophytic ovipositor. Four kinds of sensilla have been described. Notwithstanding their different external and internal morphology, they show features typical of mechanoreceptors. Three of them are evident along the external surface of the two cutting valvulae in the form of sub-spherical pegs, pit organs type 1 (holes) and pit organs type 2 (depressions), these last are similar to amphinematic scolopidia, while the fourth type is represented by subintegumental mononematic scolopidia having no direct relationship with the cuticle. In spite of their structural differences, the morphology of the described mechanoreceptors is consistent with performing a main role in allowing the perception of compression/stretching of the thick cuticle of the valvulae and their bending due to the pressure acting on the distal end of the ovipositor during substrate penetration. Such an organization is coherent with the need of endophytic Odonata to be able to evaluate the stiffness of the plant where to lay eggs.     

Larval development of Aeshna cyanea (Müller, 1764) (Odonata: Aeshnidae) in Central Italy

A three-year investigation was carried out on the life cycle of Aeshna cyanea (Müller, 1764) (Odonata, Aeshnidae) in temporary freshwater pools in Central Italy. The instars were discriminated by size and scatter plot, based on measurements of labium length, head width, metafemur length, forewing-pad length and total larval body length. The prolarvae instar was derived by Dyar's law. The mean increase value index between following and previous instar was between 1.26 and 1.33 for isometric variables, and around 1.96 for the wing-pad allometric variable. A. cyanea entered diapause mainly at the F-2 instar, placing it almost intermediate between the Southern Spain populations, which usually overwintered in the F-3 instar, and those of England and Central Europe, who spent their last winter in F-1. A. cyanea appeared to be a `summer species', as defined by Corbet (1962), and the population we studied had a semivoltine life cycle.