Mechanisms of sperm removal and sperm transfer in Orthetrum coerulescens (Fabricius) (Odonata: Libellulidae)

ABSTRACT An account of the structure of the secondary genitalia of male Orthetrum coerulescens (Fabricius) is given together with an outline of the internal genitalia of the female. Observations on the experimental inflation of the penis are described. No sperm was released during fast inflations and deflations of the penis, but sustained inflation of the penis led to the release of a clear fluid followed by the slow discharge of sperm. Examination of natural copulations in the field has shown that rapid rhythmic movements at up to 5 Hz take place in the male's third abdominal segment throughout most of copulation, but towards the end they cease though inflation is maintained. It is suggested that the rapid movements coincide with the removal of rival sperm from the female and that sperm is transferred to the female only during the maintained inflation. Possible mechanisms of sperm translocation, sperm removal and sperm transfer are discussed with reference to male and female genital structures and the action of relevant muscles.     

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.     

Fecundity and oviposition in three skimmers,Orthetrum japonicum,O. albistylum andO. triangulare (Odonata: Libellulidae)

Females of three skimmers,Orthetrum japonicum, O. albistylum andO. triangulare, were captured in forests and paddy fields. They were dissected and examined for the number of submature and mature eggs.Orthetrum japonicum andO. albistylum continuously developed eggs throughout their reproductive period. The volume of mature eggs inO. albistylum was the smallest among the three species. The number of mature eggs in ovaries decreased from morning to afternoon, suggesting that all of them oviposited around noon. The number of mature eggs laid byO. japonicum, O. albistylum andO. triangulare was estimated at about 400, 800 and >2000 a day, respectively.     

Flexible Mate Guarding Tactics in the Dragonfly Sympelrum Internum (Odonata: Libellulidae)

Mate guarding–a behaviour prevalent in odonates–is a post copulatory association during which males prevent females from re-mating. Some species use two forms of guarding: contact mate guarding, which is energetically costly but highly effective and non-contact mate guarding, which is less costly but less effective. This study aimed to determine if male Sympetrum internum (Odonata:Libellulidae) adjust the duration of contact mate guarding according to environmental, temporal and physiological factors. There was a significant interaction between male density and season on duration of contact mate guarding. Early in the season males increased the duration of contact guarding as the density of rivals increased. Later in the season males guarded mates longer irrespective of male density. Wind and temperature did not detectabiy alter the duration of contact mate guarding, suggesting that the trade-off between current and future reproductive success was more important than were physiological costs.     

On the restoration of the last relict population of a dragonfly <Emphasis Type="Italic">Urothemis edwardsii</Emphasis> Selys (Libellulidae: Odonata) in the Mediterranean

The restoration of endangered relict populations is challenging in conservation biology because they require specific environmental conditions within an inhospitable regional climate. Urothemis edwardsii Selys is the most endangered dragonfly in the Mediterranean with only one known relict small population (Lac Bleu) left in Northeast Algeria. With the absence of successful (re-)colonization over the last two decades, the restoration of the species became a top priority. To improve the status of the species in Northeast Algeria, we carried out a reintroduction and translocation scheme during 2011–2015 and assessed the changes in distribution and population size. Our restoration plan led to the emergence of three populations of which one was restored (Lac Noir), one resulted from successful translocation (Lac Tonga Northeast), and one established after successful colonization (Lac Tonga Southwest). In three localities (Lac Noir, Lac Tonga Northeast, and Lac Tonga Southwest), signs of population growth were observed, whereas no significant trend in the source population (Lac Bleu) was detected. A new population (El Graeate) was also recorded in 2015, but its origin is uncertain. Capture-mark-recapture on adults conducted in 2015 in two sites (Lac Bleu and Lac Noir) showed low recapture rates and no sign of dispersal between the two sites. Dispersal capacity of the species and conservation implications of adult distribution are discussed. This study highlights the importance of using biological indicators in selecting host habitats for the restoration of critically threatened populations.     

Aspects of life history of the Afrotropical endangered Acisoma inflatum (Selys, 1889) (Odonata: Libellulidae) in Northeast Algeria

Summary

Acisoma inflatum

is a threatened dragonfly in the Mediterranean, where only a few relict populations remain. It is listed as endangered in North Africa, where no data on its biology, behaviour or ecology are available. These latter are important to understand how this species is adapted to local environmental conditions and for establishing effective conservation plans. In this study, the life history of A. inflatum is investigated in a north-east Algerian population. By combining field and laboratory investigations on embryonic development and regular collection of exuviae during the emergence season, it is revealed that the species has direct and synchronous embryonic development, with 75% of all eggs hatching after 10 days of oviposition, and a hatching period ranging from 9 to 15 days. Emergences were asynchronous with half of the larval population (EM₅₀) emerging within the first 25 days of a 64-day emergence season, which matches a typical ‘summer species’ emergence pattern. Sex ratio at emergence was not significantly different from unity. Body size of exuviae did not show a significant pattern across the season. Exuviae males and females did not differ in their vertical stratification. Height of exuviae fixation during mergence was not random, but was rather positively correlated with support height where the relationship was logarithmic. Apparent lifespan of adults was not significantly different between sexes with a mean of 4.6 ± 2.9 days (± SD). Males and females reached their sexual maturity after two days of emergence. The information provided in this study has not been reported before and thus will be helpful in future conservation efforts.     

Foraging behavior ofPachydiplax longipennis (Odonata: Libellulidae)

Food intake, prey availability, and prey capture behavior at feeding areas were quantified in the dragonflyPachydiplax longipennis by observing focal individuals on artificial perches, where they exhibited marked short-term site fidelity. Prey capture success was high and relatively constant, but the frequency and duration of feeding flights depended on the time of day and season (at least in part because of associated variation in temperature and other physical factors), study site, sex, density of other dragonflies, and prey density. Individuals rapidly responded temporally and spatially to changes in prey availability, particularly to localized prey concentrations.     

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.     

Wind Drift Compensation in Migrating Dragonflies Pantala (Odonata: Libellulidae)

Tailwind drift compensation serves to maximize a migrant's flight distance on a given amount of energy, and crosswind drift compensation serves to hold a course true and minimize the distance flown. With full or part compensation, airspeeds are predicted to increase with greater crosswind drift. To test whether migrating dragonflies compensated for wind drift, I measured the velocity and heading of Pantala hymenaea and P. flavescens in natural flight over a lake and the ambient wind speed and direction. P. hymenaea flew north-easterly (58°), whereas P. flavescens flew significantly more east–north easterly (74°) throughout the day. Pantala spp. demonstrated part compensation for changes in crosswind drift within individuals (mean compensation = 54%, P = 0.0000), evidence for use of a ground reference to correct for drift when flying over water. Among individuals, P. flavescens compensated for crosswind drift. P. hymenaea overcompensated and then drifted downwind on one morning and compensated for crosswind drift on the next. As predicted from optimal migration theory, airspeed (5.0 m/s for both species with no tailwind) decreased with tailwind velocity both among individuals (data for both species pooled [n = 19], P < 0.0001) and within each individual as it crossed the lake (P = 0.0016).     

Hygroreceptors in the larva of Libellula depressa (Odonata: Libellulidae)

Ultrastructural and electrophysiological (single-cell recordings) investigations were carried out on the coeloconic sensilla borne by the apical antenna of the larvae of Libellula depressa (Odonata: Libellulidae). These sensilla appear as pegs located in pits. One of them is a compound sensillum constituted of two fused pegs in a common pit and the other two are single pegs located in separated pits close to each other. Coeloconic sensilla show position and ultrastructural details very similar to those described in insect hygroreceptors. The electrophysiological recordings on the apical antennae of the last larval instar of L. depressa clearly show the presence of moist and dry cells responding antagonistically to humidity changes. This study gives the first evidence of hygroreceptors in dragonfly larvae and represents the first electrophysiological approach to larval sensilla of aquatic insects. The presence of hygroreceptors in L. depressa larvae is in agreement with the hygropositive response shown by these insects in laboratory and field behavioural experiments.     

Shade Alone Reduces Adult Dragonfly (Odonata: Libellulidae) Abundance

We demonstrate that physical habitat conditions influence adult dragonfly (Odonata: Anisoptera) riparian site selection. In naturally treeless riparian areas of South Africa, invasive trees create shade and reduce native vegetation. We hypothesized that most breeding odonates select riparian areas (1) without shade, and (2) with high density and variety of understory perch structures. In two experiments at reservoir shorelines, we varied shade and perch structures. Dragonfly abundances (predominantly Trithemis species) were lower at sites with high (75%) or moderate (55%) shade cover than at sites with no shade, and lower at bare sand sites than sites containing stick perches. Perch density and variety (variety of heights and diameters) did not affect dragonfly abundance. These results indicate that shade alone directly reduces dragonfly habitat selection, isolating one aspect of habitat change that can alter insect behaviors.     

Phylogeny of the dragonfly genus Sympetrum (Odonata: Libellulidae)

The libellulid dragonfly genus Sympetrum has been recognized since 1833, but lacks any morphological synapomorphies to unite the taxon. Previous researchers have disagreed over which species belong in Sympetrum, bringing the monophyly of the genus into question. We use DNA sequence data from 6 genetic loci (16S, tRNA-valine, 12S, elongation factor 1 alpha, cytochrome oxidase subunit I, and the second internal transcribed spacer region) and 25 morphological characters (mainly genitalic) to test the monophyly of Sympetrum with Bayesian inference and maximum likelihood analyses. Under Bayesian inference, all Sympetrum species included in this study form a clade, which also contains the Hawaiian monotypic genus Nesogonia, often considered a close relative of Sympetrum. Phylogenetic analyses also reveal at least six strongly supported clades (treated as species groups) within Sympetrum, but relationships between these species groups remain unresolved or unsupported. Although the relationships between Sympetrum species groups remain unresolved, several species groups include taxa from multiple biogeographic regions/continents, and the species group sister to the rest of Sympetrum contains migratory species from the New World and Africa. This pattern suggests a complex biogeographic history in Sympetrum shaped by vicariance and dispersal. Preliminary estimates of the divergence dates of Sympetrum species groups outline a rapid radiation of the groups approximately 32-38 million years ago, possibly influenced by cooling and drying climates of the late Eocene and early Oligocene.     

The histological and histochemical studies on the ovary in relation to vitellogenesis in the dragonfly, Orthetrum chrysis Selys (Libellulidae: Odonata).

The ovaries consist of large number of panoistic ovarioles in the last instar nymph and the adult dragonfly Orthetrum chrysis (Selys). In the nymph the vitellaria are compactly filled with the primary oocytes and the vitellogenesis takes place only in the adult stage. During vitellogenesis oocytes change widely in their shape, size and cytological organisation and their developmental stages can be divided into pre-vitellogenic, early-vitellogenic, vitellogenic, late-vitellogenic and maturation age. PAS-positive material appears first around the germinal vesicle in the early-vitellogenic stage and lateron it migrates towards the periphery. Glycogen appears in the late-vitellogenic stage. DNA is abundantly present in the nuclei of the oocytes during the pre-vitellogenic and completely absent in early-vitellogenic, vitellogenic, late-vitellogenic and maturation stages. It is observed in the nuclei of follicular epithelial cells of all the stages. RNA is abundantly present in cytoplasm of the pre-vitellogenic oocytes but lateron is gradually decreases. During the early-vitellogenic and vitellogenic stages high concentration of RNA in the follicular epithelial cells has been observed. The protein bodies appear first in the interfollicular spaces and towards the periphery of the oocytes just near the enveloping follicular epithelial cells, during the early-vitellogenic stage suggesting the formation of yolk proteins from the haemolymph. In Orthetrum chrysis the sudanophilic bodies appear first in the follicular cells and then lie in the peripheral region of the oocytes suggesting the incorporation of yolk lipid either from the follicular epithelium or from the haemolymph through the follicular epithelium. The phospholipids are synthesised in pre-vitellogenic to the late-vitellogenic stages. In the late-vitellogenic stages the phospholipid granules are present abundantly in the follicular epithelium while in the maturation stage they disappear suggesting their utilisation in the formation of membranes like vitelline and chorion. The neutral fats are present in the form of large number of droplets in the oocytes during the maturation stage.     

Aggregative oviposition behaviour of Helicoverpa spp. (Lepidoptera: Noctuidae) in contaminated chickpea crops

Abstract Aggregative egg distribution by Helicoverpa spp. within experimental field plots of chickpea containing various contaminant plant species was documented at different stages of plant growth. More eggs were laid on contaminant faba and wheat plants than on the surrounding chickpea. The level of egg aggregation on faba and wheat plants increased with general crop growth, becoming evident before a significant disparity in growth rates between these species and the surrounding chickpea became apparent. In chickpea plots contaminated with canola, aggregative oviposition activity on the contaminant plants was observed in the early seedling stage whilst a clear height differential between the two species was evident. Total Helicoverpa egg density on contaminant plants was not correlated with height above chickpea for any of the contaminant species. It is proposed that the aggregative oviposition behaviour of Helicoverpa spp. in contaminated chickpea results from differential apparency of the contaminant plants to ovipositing moths. Differential apparency may result from several factors including growth rate and morphological differences between chickpea and the contaminants. Physiological changes in chickpea, including growth stage-related changes in acid production by the foliage, may also contribute to the greater apparency of contaminant plants in chickpea plots. The implications of differential host plant apparency for insect pest management are discussed.     

Egg envelope synthesis and chorion modification after oviposition in the dragonfly Libellula depressa (Odonata, Libellulidae).

Libellula depressa (Odonata, Libellulidae) is an exophytic dragonfly ovidepositing eggs in clutches on the surface of floating plants and algae. The present work investigates, at ultrastructural level, the gradual differentiation of the egg envelopes and the chorionic changes after egg deposition in water. The ovary of the mature female of L. depressa is composed of numerous strings of panoistic ovarioles, where the eggshell formation takes place gradually throughout the activity of the follicle cells. The present data show that the egg envelopes are constituted of a very thick electrondense vitelline envelope, a thin endochorion and an extremely thick exochorion composed of a fibrillar matrix resting on a thin electrondense layer. After deposition in water, L. depressa eggs, initially white and almost transparent, gradually become brown spots in a semitransparent jelly coat, rich of incorporated debris. The jelly coat enveloping the eggs of L. depressa derives exclusively from the exochorion, constituted of a fibrillar matrix, which swell at contact with water. The jelly-like coat performs an adhesive function and presumably a protective role during egg segmentation and ensuing larval hatching.     

Apical antennal sensilla in nymphs of Libellula depressa (Odonata: Libellulidae)

Abstract. In an ultrastructural study of the apical antenna of the last nymphal stages of Libellula depressa (Odonata: Libellulidae), we found long sensilla trichodea, 2 sensory pegs, and a coeloconic sensillum on the last article of the flagellum (the distal part of the antenna). The long sensilla trichodea are mechanoreceptors, almost identical to the long filiform hairs of some terrestrial insects and the first sensilla of this kind to be described in aquatic insects. Particular attention was given to the complex coeloconic sensillum, a compound sensillum innervated by 2 groups of 3 neurons wrapped in a dendritic sheath. A cuticular sleeve envelops the distal portion of the outer dendritic segment. The cuticle of the coeloconic sensillum shows wide channels and is contiguous to the underlying granular and fibrillar layer. Similar structures on the antennae of the adults of other dragonflies were identified as chemoreceptors in previous studies. We hypothesize that this larval coeloconic sensillum might likewise have a chemosensory function, responding to molecules that diffuse through the cuticle and the underlying granular and fibrillar layer, as no clear pore or pore-tubule system is visible. Alternative functions are also explored on the basis of morphological details.     

On the respiratory mechanism during underwater oviposition in a damselfly Calopteryx cornelia Selys

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.     

Two gynandromorphs of Sympetrum striolatum (Charpentier, 1840) (Odonata: Libellulidae)

Two gynandromorphs of the dragonfly Sympetrum striolatum , with different features, are described here, with special emphasis on the genitalic structures.     

Egg clutch patterning in Lestes virens (Odonata,Lestidae) with evolutionary emphasis on endophytic oviposition in lestid dragonflies

Egg deposition within plants is one of the most widely distributed and ancient behaviors in Odonata. The resulting clutch consists of eggs placed in peculiar pattern that can be a characteristic for certain groups of Odonata. Despite their importance for paleontological and evolutionary research, data on egg‐clutch positioning are missing or insufficient for most species. Here, patterning of egg clutches in Lestes virens was measured and described in detail for the first time. The female usually produces a linear row of single eggs directed at an angle rightward or leftward to the longitudinal axis of plant substrate. Less often eggs are arranged in egg‐sets consisting of up to 4 eggs. Apparently, the female insect follows the rigid behavior stereotypes during oviposition and is unable to easily switch to the alternate stereotypical behavior of single egg deposition or production of multiegg sets. Based on a literature review and original data, egg clutch patterning of European Lestidae is overlaid on preexisting phylogenies. The resulting evolutionary scenario of egg‐clutch patterning can be considered in the framework of egg‐laying behavior in Lestidae.