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.     

Neurosecretory system of the ventral ganglia in the dragonfly,Orthetrum chrysis (selys) (Odonata: Libellulidae)

Summary The neurosecretory cells of the ventral ganglia in the adult dragonfly, Orthetrum chrysis, are classified into A, B, C1 and C2 cells. The neurosecretory material in the ventral ganglia is composed of PAS-positive material with 1-, 2-glycol groups and some proteins. The proteins rich in cystine or cysteine occur predominantly in the A cells, moderately in C cells and negligibly in B cells. Proteins containing arginine occur in A and B cells only, and those containing basic amino acids occur in C2 cells. The neurosecretory pathways and the neurohemal organs are also described.     

Reproductive behavior of the dragonfly,Orthetrum japonicum (Odonata: Libellulidae)

The reproductive behavior of the dragonfly,Orthetrum japonicum, is described. Behavioral processes of turnover of territorial males, simultaneous guarding of 2 females by a male, and copulation by non-territorial males are described. The males with longer hind wings won the territorial conflicts more frequently. The total duration of territorial residence of a given male was correlated with the number of his matings, but not correlated with the length of his abdomen or hind wings. The territorial site with the lower degree of vegetation cover was occupied by males more consistently. Males in more consistently occupied territorial sites did not have longer abdomen and hind wings than males in less consistently occupied sites. The territorial site where the larger number of copulations was observed was not occupied more consistently. Selection episode analysis using the method of Arnold & Wade (1984a, b) showed that direct selection on the hind wing length favored the short wing and that direct selection on the abdomen length favored the long abdomen during mating.     

Selection and reproductive success in males of the dragonfly,Orthetrum japonicum (Odonata: Libellulidae)

Reproductive success, copulation success, and mating success were measured for a population of male dragonflies,Orthetrum japonicum. Copulation success explained the greatest variation in reproductive success. The proportion of copulations followed by oviposition was positively correlated with the number of oviposited eggs per mating. Directional selection on four morphological characters was estimated. The effect of selection on correlated traits was comparable to that of direct selection. Directional selection varied between traits and between episodes in a single trait. The probability that the observed directional selection on the four morphological traits was expected under the condition of the selective neutrality of traits was not smaller than 5%.     

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.     

Establishment and organization of territories in the dragonfly Sympetrum rubicundulum (Odonata: Libellulidae)

Territory formation by males of the dragonfly Sympetrum rubicundulum was studied at a shallow temporary pond in Pennsylvania. Most males arrived at the pond between 1100 and 1200 hours and sought unoccupied sites in which to land, which they immediately began defending against conspecific males. Incoming males were never observed to challenge previous residents for possession of territories. Most conflicts between males occurred in the mid-afternoon and involved occupants of adjacent territories; residents experienced a clear ownership advantage in these conflicts. Escalated contests for possession of territory sites did not occur, perhaps because unoccupied sites were continually becoming available due to territory abandonment. Certain regions of the pond were consistently favoured by both sexes and most mating occurred in them: the spatial distributions of copulating pairs, ovipositing females and unpaired females were in close agreement with the spatial distribution of territorial males. High density sites were also the first sites occupied in the morning by incoming males. Although most of the pond and the surrounding land was thickly vegetated, territorial males and ovipositing females were both significantly clumped along the water's edge. The spatial distribution of males may have resulted from a balance of factors: peripheral sites were better than central ones for encountering incoming females, but areas far from the water's edge were unsuitable for female oviposition.     

The Morphology of the Stomodeal Nervous System in the Adult Dragon Flies Bradinopyga geminata and Orthetrum chrysis (Libellulidae)

The morphology of the stomodeal nervous system of the adult dragon flies Bradinopyga geminata and Orthetrum chrysis is described. No gastric ganglion or ganglion ingluviale has been found. Instead the oesophageal nerve forks near the junction of the proventriculus and the midgut. The two nerves run on either side of the midline as ingluvial nerves and enter the proventricular ganglionic masses. These ganglionic masses are connected by a transverse nerve, which has been called as the nervus transversus proventriculare. Both bipolar and multipolar types of sensory cells have been found over the surface of the crop. These cell bodies appear to be interconnected by connective tissue. Dendrites of these cells terminate on the longitudinal muscle fibres, surrounding the proventriculus and the midgut. The proximal processes of these cells enter the proventricular ganglionic mass. In methylene blue whole mounts they resemble the stretch receptors, hence it is quite probable that they play some role in the peristaltic movement of the gut. The corpora cardiaca lie dorsal to the pharynx and are connected to the brain by two pairs of nerves, the nervi corporis cardiaci (NCC I, NCC II). Unlike in other insects, the nerve connecting the corpora cardiaca with the corpora allata is slender and arises as a branch of the nerve, nervus corporis allati II. The corpora alata are spherical to ovoid in shape and lie ventral to the nerve cord. Anteriorly they are attached to the inner wall of the hypopharynx and posteriorly to the subesophageal ganglion by a pair of nerves, the nervi corporis allati II.     

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.     

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.     

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.     

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.     

Isotope evidence for latitudinal migrations of the dragonfly Sympetrum fonscolombii (Odonata: Libellulidae) in Middle Asia

• Sympetrum fonscolombii dragonflies are believed to migrate seasonally. In the spring and early summer, the already-mature dragonflies arrive in Middle Asia for reproduction. In the late summer and autumn, summer-generation dragonflies migrate to the south. Their wintering places remain unknown.
• Stable hydrogen (δ²H) and oxygen (δ¹⁸O) isotope analyses were conducted to confirm the migration of S. fonscolombii and determine the wintering area. Stable isotope composition of carbon (δ¹³C) and nitrogen (δ¹⁵N) in wings and legs was used to clarify the habitats in which dragonfly development took place.
• Three cohorts of dragonflies collected in different regions of Middle Asia were used for analysis: (i) immigrants that arrived in the spring, (ii) residents that developed in Middle Asia, and (iii) transit dragonflies migrating to the south during autumn.
• The average δ²H values in the wings were significantly higher in immigrants (−96‰) than in residents (−134‰) and transit individuals (−124‰). High δ¹⁸O and δ¹⁵N values in the tissue of immigrants confirmed their southerly origin.
• Based on the species range and the global distribution of annual averages of δ²H and δ¹⁸O values in precipitation, the latitudinal migrations of S. fonscolombii were inferred to cover the area from the proposed natal regions of immigrants in South-West Asia (below ∼36°N) to Southern Ural and the south of Western Siberia in the north (54–55°N) with a maximum migration distance of more than 4000 km.

     

Influence of temperature and photoperiod on embryonic development in the dragonfly Sympetrum striolatum (Odonata: Libellulidae)

Temperature and photoperiod play major roles in insect ecology. Many insect species have fixed degree‐days for embryogenesis, with minimum and maximum temperature thresholds for egg and larval development and hatching. Often, photoperiodic changes trigger the transfer into the next life‐cycle stadium. However, it is not known whether this distinct pattern also exist in a species with a high level of phenotypic plasticity in life‐history traits. In the present study, eggs of the dragonfly Sympetrum striolatum Charpentier (Odonata: Libellulidae) are reared under different constant and fluctuating temperatures and photoperiodic conditions in several laboratory and field experiments. In general, and as expected, higher temperatures cause faster egg development. However, no general temperature or light‐days for eyespot development and hatching are found. The minimum temperature thresholds are distinguished for survival (2 °C), embryogenesis (6 °C) and larval hatching (above 6 °C). Low winter temperatures synchronize hatching. Above 36 °C, no eyespots are visible and no larvae hatch. In laboratory experiments, light is neither necessary for eyespot development, nor for hatching. By contrast to the laboratory experiments, the field experiment show that naturally changing temperature and photoperiod play a significant role in the seasonal regulation of embryonic development. The post‐eyespot development is more variable and influenced by temperature and photoperiod than the pre‐eyespot development. This developmental plasticity at the end of the embryogenesis might be a general pattern in the Libellulidae, helping them to cope with variation in environmental conditions.     

Ionic and osmotic regulation of the haemolymph of the dragonfly, Libellula quadrimaculata (Odonata : Libellulidae)

Larvae of the widespread dragonfly, Libellula quadrimaculata, were adapted to a series of salt solutions, and the osmotic pressure, and sodium, potassium and chloride concentrations in the haemolymph measured. The regulation of potassium is extremely efficient over the range 0–50 m-mole/l. external concentration. Above this, larvae die. Sodium and chloride are regulated to a lesser extent, the larvae being able to withstand considerable changes in the concentration of these ions in the haemolymph. However, at higher external concentrations, the haemolymph concentration of these ions is maintained below that of the external medium. The osmotic pressure is regulated in parallel with sodium concentration over most of the range tested. However, in higher salinities, the osmotic pressure of the haemolymph does not fall below that of the external medium. This is seen as a strategy to limit the amount of drinking in saline media. Overall, the osmoregulatory system of L. quadrimaculata resembles that of brackish-water insects, rather than that of the more strictly freshwater dragonflies that have been studied.     

The influence of temperature on the functional response of the dragonfly Celithemis fasciata (Odonata: Libellulidae)

Summary Four biennial species (Oenothera erythrosepala, Dontostemon dentatus, Erigeron strigosus, and Erigeron sumatrensis) from the dune system at Azigaura, Japan, were sand-cultured in pots with different nutrient levels, and the critical plant size for the development of a flowering stalk was determined. The critical size decreased in the order O. erythrosepala>E. strigosus>D. dentatus=E. sumatrensis, and corresponded to the order of their distribution along the gradient of soil fertility in the dune system. This correspondence is discussed, and it is concluded that in a less fertile environment, biennials producing seeds after attaining a larger size have an advantage over those producing seeds earlier.     

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.     

The adaptive significance of non-contact mate guarding by males of the dragonfly,Nannophya pygmaea Rambur (Odonata: Libellulidae)

InNannophya pygmaea, ovipositing females were frequently disturbed by conspecific males. Disturbed females often copulated with one of these males or flew away from the pool. Females which flew away from the pool due to male disturbance often returned later the same day and mated with different males. A territorial male would guard his ovipositing mate by hovering above her, presumably trying to prevent her from moving out of his territory. A non-territorial male would also guard his mate in a similar way, both at a vacant water area which was not occupied by any territorial males, or within the territory of a resident male. In addition, both territorial and non-territorial males chased intruding males in an attempt to prevent their mates from being stolen. Territorial males defended their mates better than non-territorial males. Both males and females often mated more than once in the course of a single day. Some territorial males copulated with a new female while another mate oviposited in their territories. This observation supported the “multiple mating hypothesis” proposed by Alcock (1979) and Uéda (1979) but other evidence suggested that this is an inadequate explanation for the non-contact guarding ofN. pygmaea.     

A dragonfly in the desert: genetic pathways of the widespread Trithemis arteriosa (Odonata: Libellulidae) suggest male-biased dispersal

Water-dependent species inhabiting desert regions seem to be a contradiction in terms. Nevertheless, many species have evolved survival strategies for arid conditions. In Odonates (dragonflies and damselflies), both larvae and adults require very different and complex water-associated habitat conditions. The present study investigates the genetic diversity, population structure and dispersal patterns of a desert inhabiting odonate species, the Red-veined Dropwing dragonfly, Trithemis arteriosa. Eight populations from the arid Namibia and four population sites in the more tropical Kenya were compared by using nine microsatellite loci, one non-coding nuclear fragment and the mtDNA fragment ND1. Microsatellite analyses as well as the nuclear fragment reveal a high allelic diversity in all populations with almost no genetic sub-structuring. In contrast, ND1 sequence analyses show sub-structuring and??with two exceptions??only private haplotypes. The conflicting patterns of nuclear versus mitochondrial markers suggest a male-biased dispersal in this species. Results indicate that male dispersal is dependent on the environmental stability of the habitat, while females are philopatric. This life history adaptation would allow females to save energy for mating and oviposition in the demanding environment of a desert region. The results give direct insights into the dispersal pathways of a desert-inhabiting, strongly water dependent flying insect.     

Mechanism of the wing colouration in the dragonfly Zenithoptera lanei (Odonata: Libellulidae) and its role in intraspecific communication

Zenithoptera dragonflies are known for their remarkable bluish colouration on their wings and unique male behaviour of folding and unfolding their wings while perching. However, nothing is known about the optical properties of such colouration and its structural and functional background. In this paper, we aimed to study the relationship between the wing membrane ultrastructure, surface microstructure and colour spectra of male wings in Zenithoptera lanei and test the hypothesis that colouration functions as a signal in territorial fights between males. The results show that the specific wing colouration derives from interference in alternating layers of melanized and unmelanized cuticle in the wing membrane, combined with diffuse scattering in two different layers of wax crystals on the dorsal wing surface, one lower layer of long filaments, and one upper layer of leaf-shaped crystals. The results also show that the thicker wax coverage of the dorsal surface of the wings results in increased brightness and reduced chroma. In the field experiments, we have demonstrated that there is a reduction of aggressive reactions of rivals towards individuals with experimentally reduced amount of blue wing colouration.     

Coupling and oviposition behaviour in two macrodiplacinid libellulids–Aethriamantha rezia (Kirby) and Urothemis assignata Selys (Libellulidae: Odonata)

Coupling and oviposition behaviour were studied in two macrodiplacinid libellulids– Aethriamantha rezia (Kirby) and Urothemis assignata Selys in die Ibadan area of Nigeria. Sperm translocation occurs in both libellulids after the male has secured the female. Pre-mating tandem flight occurs prior to mating. Mating is long in A. rezia (384–716 s, mean 480) and shorter in U. assignata (69–122 s, mean 94). After mating, both libellulids rest in tandem, the duration being longer in A. rezia (25–82 s, mean 42) than in U. assignata (0–24 s, mean 11). This is followed by tandem exploration in A. rezia and later by oviposition, the male guarding the female. In U. assignata , tandem exploration is absent, but oviposition is usually in tandem. The duration of oviposition is similar in both libellulids–6 7–148 s, mean 95, in A. rezia and 63–183 m, mean 133, in U. assignata. Aethriamantha rezia oviposits mainly on Pistia stratiotes while plants utilized by U. assignata are more variable, although P. stratiotes is also preferred. The number of eggs laid per oviposition is high, averaging 435 in A. rezia and 398 in U. assignata. High population of males at water prolongs mating and oviposition in the two species. The ecological habitats of both overlap.