Origin and pathway of the epidermal secretion in the damselfly head-arresting system (Insecta: Odonata)

In damselflies, the arrester system is responsible for an additional attachment of the head to the neck. It consists of a pair of mobile postcervical sclerites (SPC) covered by microtrichia. In their lateral position, SPCs can fixate the head on fields of microtrichia on the back surface of the head. The intact surface of microtrichia of the SPC is usually covered by a lipid-containing secretion. The present study provides ultrastructural data on the secretory epidermis and pore channels adapted to transport the secretion to the cuticle surface. 1) Shock-frozen preparations of the contact area show that microtrichia of co-opted surfaces do not interlock with each other. When co-opted fields are pressed to each other, deformations of flexible microtrichia of the SPC result in an increase of contact area between corresponding structures and consequently in an increase of frictional forces. 2) In the area of the SPC, only electron-lucent vesicles have been found. Histochemical procedures revealed that the material stored in vesicles and liberated on the external surface of the SPC is presumably non-volatile lipid. 3) Only a small number of large pore channels reach the surface of a microtrichium. Most of them terminate with tiny terminal channels, whose diameter is approximately six to twenty times smaller than the diameter of structures of secretory blooms occurring in SEM in shock-frozen and air-dried preparations. The terminal channels do not penetrate the epicuticular layer. It seems that the secretion reaches the epicuticle through terminal channels and diffuses through the epicuticle without any channel structures.     

Female internal genital organs of the damselfly Pseudagrion rubriceps selys (Zygoptera: Odonata)

The morphology, anatomy and histology of the female internal genital organs of the adult damselfly Pseudagrion rubriceps Selys were studied in detail. The paired, long, panoistic ovaries each lead posteriorly into an oviduct. The oviducts unite to form a short common median oviduct leading into the "eighth complex" localized in the 8th abdominal segment. The "eighth complex" comprises the bursa copulatrix, spermatheca and vagina. A pair of prominent accessory reproductive glands, localized in the 9th segment, open into the vagina.     

Alternative mate-finding tactics in a non-territorial damselfly (Odonata: Coenagrionidae)

Males of the non-territorial damselfly Enallagma hageni have two alternative tactics for finding mates: (1) they search the banks of the pond for unmated females (searching tactic), or (2) wait at oviposition sites for females that resurface prematurely from underwater oviposition (waiting tactic). Although the searching tactic yielded more fertilizations than the waiting tactic, for time invested, the waiting tactic became increasingly successful later in the reproductive season due to changes in female oviposition behaviour. The two tactics can be maintained in the population because males can mate by the waiting tactic during the afternoon when few females are available to searchers. Among males visiting the breeding site an equal number of times, males mating by a mixture of tactics were as successful as males mating only by the main tactic. Because marked males were found to use both tactics, these behaviours are interpreted as evidence of behavioural plasticity within individuals, representing one conditional evolutionary strategy.     

Impact of predation by spiders on a territorial damselfly (Odonata: Calopterygidae)

Summary Predation by orb-weaving spiders and crab spiders on the damselfly Calopteryx haemorrhoidalis was studied at a small stream in Southern France. One species of orb-weaving spider, Larinioides folium, caught 76% of the damselflies which fell prey to spiders. Displacement experiments on spiders on sections of bank and the positioning of webs in male territories show that the density and distribution of damselflies is not influenced by orbweb density or by the position of webs. Predation rates corresponded to orb-web density, but neither for sex nor for stage was there a relationship with damselfly density. Mean daily predation rates ranged between 0.9% for females and 4.1% for adult males. Predation risk to adult damselflies by orb-weaving spiders was male biased, whereas among tenerals there was no bias. Males were captured more frequently at territories near the water. Captures show a maximum at noon when territorial disputes of adult males were most frequent. After orbwebs were placed within territories predation rate of males was strongly increased. Predation risk to adult females in the direct vicinity of the stream was less than in the bank vegetation where they perch close to orbwebs. The risk of predation by crab spiders, which catch damselflies at their perching sites, was not sex-biased.     

Behavior of males in a reproductive aggregation of the banded demoiselle Calopteryx splendens (Insecta, Odonata)

The view according to which damselfly males practice two alternative reproductive tactics of access to females is critically discussed. It is widely accepted that some males (“territorial” ones) have priority as potential female partners, while others (“sneakers” or “wanderers”) are incapable of retaining an individual territory. They have a chance of mating only by intruding briefly into the area defended by a “territorial” male when a female is present there. Thus, the tactics of a “territorial” male consists in waiting for a female in its territory and copulating with it “by agreement,” whereas non-territorial males resort to forced copulations. By observation of individually marked males (48 out of 118) it was shown that every male could be regarded as “territorial” during a certain period and as a “wanderer” before and after it. Thus, no correlation between the modes of space use by a male (residence/mobility) and the characters of its external morphology and/or signal behavior appears to be possible in principle. According to the data obtained, a more plausible explanation is that the female chooses not the male but the best area for oviposition. In addition, it was ascertained that adherence to forced copulations cannot constitute successful “tactics” since they rarely result in insemination, neither by “territorial” nor “non-territorial” males. In other words, we are dealing not with certain alternative tactics (i.e., specialized adaptive mechanisms that have evolved in the species) but simply with the results of different sets of circumstances at a given moment.     

Differences in Mating Propensity Between Immature Female Color Morphs in the Damselfly Ischnura elegans (Insecta: Odonata)

Female-limited color polymorphisms occur in a variety of animal taxa where excessive male sexual harassment may explain the coexistence of multiple female color morphs. In the color polymorphic damselfly Ischnura elegans, mature and immature female color morphs coexist at the mating site where males are in search for suitable mating partners. Here, we study male preference and female mating propensity for the two immature female morphs. As would be expected, compared to mature morphs, both immature female morphs mate much less. Within immature females, one morph consistently mates more frequently compared to the other morph, a pattern that is similar for the ontogenetically corresponding mature female morphs. Preference experiments with the two differently colored immature female morphs, however, did not indicate male mate preference for either morph. Low mating frequencies of immature females at natural sites in combination with relatively high attractiveness of immature models in terms of male preference indicate that female behavior influences female mating success.     

Dragonfly associations (Insecta: Odonata) in relation to habitat variables: a multivariate approach

In a dragonfly survey, carried out in a lowland wetland area in eastern Austria, a total of 19 resident species was recorded. Multivariate statistical procedures were used to analyse the relationship between dragonfly assemblage patterns and environmental variables. Besides widespread and euryoecious species with unspecific habitat requirements two dragonfly associations were identified: on the one hand species mainly occurring at temporary natural and near-natural ponds characterised by rush and reed vegetation, on the other hand species preferring permanent waters such as the artificial waterbodies in the investigation area characterised by floating macrophytes. Water persistence and the existence of floating macrophytes determined the formation of species assemblages.     

Possible coevolution of male and female genital form and function in a calopterygid damselfly

In this paper some evolutionary changes of genitalia in the damselfly Calopteryx haemorrhoidalis are investigated by determining their current and past function. Calopteryx haemorrhoidalis males stimulate females by aedeagal frictioning on a set of vaginal sensilla. The aedeagus is considerably variable and positively correlates with volumes of ejected sperm from the spermatheca. Interestingly, females show a significantly reduced sensillum number compared with other family members. Here I explore whether there existed directional selection for aedeagal width at its evolutionary onset; and whether the sensillum reduction evolved to make sperm ejection less effective. Using C. haemorrhoidalis aedeagi in females whose species retained the ancestral conditions (no stimulatory ability and large sensillum numbers), Hetaerina cruentata and C. xanthostoma, my results corroborated these assumptions: variation in aedeagal width inversely correlated with sperm ejection rate while sperm ejection was higher in species with high sensillum numbers. A suggested coevolutionary interpretation of these results in C. haemorrhoidalis is that aedeagal width was favoured which was followed by a sensillum reduction.     

Simple pond restoration measures increase dragonfly (Insecta: Odonata) diversity

Ponds are home to a diverse community of specialized plants and animals and are hence of great conservation concern. Through land-use changes, ponds have been disappearing rapidly and remaining ponds are often threatened by contamination and eutrophication, with negative consequences for pond-dependent taxa like amphibians or dragonflies (Odonata: Anisoptera and Zygoptera). Increasingly, restoration measures such as removal of shading terrestrial vegetation or submerged organic matter are implemented to counteract current threats, but how these measures affect the target taxa is rarely assessed. We tested if and how simple pond restoration measures affectionate diversity. We propose that pond restoration influences the light regime, which promotes aquatic and riparian vegetation important for different dragonfly life stages, thus increasing their diversity. Additionally, we assume that this changes dragonfly species composition between restored and unrestored ponds. We surveyed exuviae in the riparian and aquatic vegetation along the shore of 29 (12 restored, 17 unrestored) man-made ponds in southwest Germany and assessed environmental variables known to affect dragonfly diversity. We identified the cover of tall sedges and submerged macrophytes as the driving biotic variables for dragonfly diversity and species composition, with restoration measures affecting submerged macrophyte cover directly but tall sedges indirectly via available sunlight. This study demonstrates that simple restoration measures not only have a positive effect on overall dragonfly diversity, but also increase habitat suitability for several species that would otherwise be absent. We therefore propose dragonflies as a suitable flagship group for pond conservation.     

Population structure of the rare damselfly,Ischnura gemina (Kennedy) (Odonata: Coenagrionidae)

Summary The population structure of the endemic San Francisco Bay Area damselfly, Ischnura gemina, is examined using mark-recapture methods. Average daily movements, sex ratios, population size, maturation times, survivorship, and dispersion patterns, were recorded and calculated from two small (each less than one hectare) sites 150 m apart in Glen Canyon, San Francisco. Of 563 adults marked over 36 days, 412 (73%) were recaptured at least once. Average daily movements for males and females were less than 6 m, suggesting local movements. However, directional movements of 150 m were observed from one site to the other, indicating dispersal potential. One of the populations was a satellite composed entirely, of emigrating individuals from the other site; no larvae or teneral adults were found at the satellite area. Males were more aggregated than females at both sites. Both sexes were highly clumped at one site but were nearly randomly dispersed at the other site. Total population size for both sexes tended to be constant throughout the sampling period, at about 250. Adult population estimates showed more males were present than, females, but larval counts at one site indicated only a slight excess of males. Average life span estimates ranged from 6.5 days (females) to 23.3 days (males). One male lived at least 36 days. Maturation time for males was about 5–7 days, 7–10 days for females. A long life span and long flight season (March to November) are probably adaptations to the foggy San Francisco climate. All populations of I. gemina located to date are small, possibly originating from founders from nearby demes, and may be subject to different selection pressures. The dispersal potential of I. gemina may increase its chance of survival should small urban demes be threatened with destruction.     

Maximum and actual field feeding-rates in larvae of the damselfly Pyrrhosoma nymphula (Sulzer) (Odonata: Zygoptera)

The paper presents information on factors influencing maximum feeding-rates (measured in the laboratory) in larvae of the damselfly Pyrrhosoma nymphula, and compares estimates of feeding rates in the field with maximum feeding-rates for larvae of the same size at the same temperature. Maximum feeding-rates in Pyrrhosoma larvae of different sizes were measured at 4, 5, 10 and 15°C. At low temperatures (4–5°C) maximum feeding-rate was depressed more than predicted either from the effects of temperature on the gut-clearance time of larvae or their respiratory rate. Maximum feeding-rate declined steadily throughout metamorphosis in the final instar and stopped completely prior to emergence of the adult. Feeding-rates in the field were calculated from ‘balanced’ energy equations, i.e. from the sum of larval growth, respiration and the production of faeces and exuviae. Feeding rates in the field were always much less than maximum feeding-rates for the same size of larva at the same temperature, and at the most were never more than 70% of the latter; they were closer to the maximum in summer than in winter, when they fell as low as 20% of the maximum. The relevance of these findings to studies on the efficiency with which carnivores utilize energy in the field, and the importance of Odonata larvae as predators in aquatic ecosystems is discussed.     

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     

An exploration of genome size diversity in dragonflies and damselflies (Insecta: Odonata)

Like most insect orders, the Odonata (dragonflies and damselflies) remain poorly studied from the perspective of genome size. They exhibit several characteristics that make them desirable targets for analysis in this area, for example a large range in body size, differences in developmental rate, and distinct modes of flight – all of which are related to genome size in at least some animal taxa. The present study provides new genome size estimates and morphometric data for 100 species of odonates, covering about 1/5 of described North American diversity. Significant relationships are reported between genome size and body size (positive in dragonflies, negative in damselflies), and there is also indication that developmental rate and flight are related to genome size in these insects. Genome size is also positively correlated with chromosome number across the order. These findings contribute to an improved understanding of genome size evolution in insects, and raise several interesting questions for future research.     

Disturbed population genetics: suspected introgressive hybridization between two Mnais damselfly species (Odonata)

Mnais costalis and M. pruinosa are damselflies (Odonata: Calopterygidae) with low dispersal abilities, both during their aquatic stream-living immature stage and their flying adult stage. A previous nuclear DNA (nDNA) sequencing and morphology study showed that these two species are very closely related, and cohabit widely in western Japan. The two species, however, segregate microhabitats along a stream: M. costalis lives in the lower reaches, and M. pruinosa in the upper reaches. In this study, our analyses were based on mitochondrial DNA (mtDNA), which usually mutates faster and is more variable among individuals than nDNA, and which is inherited maternally. We found that most COI haplotypes were shared between the two species, and that for most study sites interspecific riverine genetic structures were not clarified by mtDNA analysis. Incongruent population genetic structures based on nDNA and mtDNA suggested hybridization and introgression of mtDNA between the two species.     

Odonata (Insecta) as a tool for the biomonitoring of environmental quality

Despite the fundamental dependence of human populations on water resources, a range of anthropogenic impacts, in particular the removal of riparian vegetation, threaten freshwater environments. One of the most effective means of evaluating the effects of anthropogenic disturbance in aquatic ecosystems is the use of bioindicators, and the insects of the order Odonata are among the most efficient indicators, due to their enormous sensitivity to environmental changes. In this context, the present study aimed to verify which parameters of the odonate community (species richness, abundance/biomass, composition, taxonomic diversity and taxonomic/phylogenetic distinctness) are most effective for the evaluation of the loss of environmental integrity. The study focused on 50 streams in the northeast of the Brazilian state of Pará. The streams were sampled during the dry season, between June and August 2011. The physical characteristics of each stream were evaluated using a Habitat Integrity Index (HII). The species composition provided the best parameter for the evaluation of ecological integrity, providing a relatively accurate assessment at a lower mean research cost than other parameters. Taxonomic diversity and distinctness also provided relatively reliable results, contributed additional information on the evolutionary relationships among the odonate taxa, and also provided a low-cost approach. Deconstructing communities is necessary to detect impacts, considering the considerable variation in the environmental requirements of the different species. Overall, the parameter that best responded to gradients of disturbance was species composition, followed by diversity and taxonomic distinctness. Given these findings, odonate-based biomonitoring should focus on these parameters to guarantee the optimal detection and evaluation of habitat alterations.