Influences on variation in territorial tenures of male white-faced dragonflies (Leucorrhinia intacta) (Odonata: Libellulidae)

Some individuals in species with extended periods of territorial occupancy may change territory locations within a single bout of territorial activity. Length of occupancy of mating territories among males in a local population of white-faced dragonflies (Leucorrhinia intacta) varied from more than 6 h to 15 min or less. Males with short tenures often established territories in several locations on the pond during a day. Several hypotheses have been proposed to explain shifting territorial sites rather than remaining in a single site during one bout of territoriality. We attempted to test the hypothesis that males shift to leave low-quality sites. Site quality may be affected by costs of defense in relation to intruder rate and the mating benefits of holding the territory. To test whether variation in these possible effects of benefits and costs of territoriality influenced tenure, we manipulated local quality of oviposition substrate and perch density. The quality of oviposition substrate, but not perch density, influenced both potential benefits and costs of territoriality. Female density was higher in areas with good substrate, but so were rates of males intruding into the territories, rates of chasing by territorial males, and local density of territorial males. More matings occurred in areas with good substrate, but among males with tenures of 15 min or more, mating success per male and tenure lengths did not differ statistically among treatments. Defense costs were low for all treatments and perhaps were not an important influence on tenure duration. Territorial males in this population probably adjusted local density to expected mating success by initial choice of site rather than by varying tenure length. Variation in tenure length at a site resulted, in part, from stochastic external factors, such as predation attempts.     

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).     

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.     

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.     

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.     

Dragonflies (Odonata) of thermal springs in Central Asia

Sixteen species of dragonflies were recorded in 12 thermal springs of Central Asia. Of these, Ischnura forcipata, I. pumilio, Orthetrum anceps, O. brunneum, and Sympetrum haritonovi were widely distributed in hydrothermal waters. In the high mountains of Pamir, dragonflies can develop only in hot springs. The maximum altitude recorded for the habitat of O. brunneum was 3950 m a.s.l.     

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.     

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.     

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.     

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.     

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.     

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.     

The structure and function of the genitalia in the Libellulidae (Odonata)

An outline of the morphology of the secondary genitalia of libellulid dragonflies is given together with a more detailed treatment of the components of the fourth (distal) segment of the penis. The variations of penis structure in 70 species, representing ten of the 11 subfamilies, are surveyed and catalogued. There is remarkable diversity in the presence and form of penis structures, particularly those associated with the medial process, namely the flagelium, cornua and inner lobes. A comparable survey of females shows that they can be provisionally divided into ten types according to the number and size of their sperm-storage organs. Comparisons of penis structures with the sperm-storage organs of females suggests that in species whose females store large amounts of sperm, usually doing so in one or a pair of organs, males possess highly extensible, lobate and bristly penes, commonly equipped with cornua. By contrast, in those species in which the females store small volumes of sperm, usually doing so in three organs, the penes bear either flagella equipped with proximally directed spines or with large terminal barbs, or they have narrow, spiny inner-lobes which resemble flagella. Some functional interpretations are proposed in the light of sperm competition. The variability of genital structure can be interpreted adaptively and does not closely reflect phylogenetic relationships. Functionally convergent structures have evolved a number of times from different origins.     

Choice of Lookout Posts by Territorial Amberwing Dragonflies, Perithemis tenera (Anisoptera: Libellulidae)

Many territorial insects use specific perches, or lookout posts, from which they perceive and react to conspecific males and females. We investigated the lookout post choice of territorial male amberwing dragonflies (Perithemis tenera). An observational study indicated that males rarely perched directly at their oviposition site; rather, they perched farther out from shore than, and within 2 m from, their oviposition site. In an experimental study, we provided an array of perches at different distances from the shore and oviposition site to eliminate perch limitation as a factor in perch choice. The results of the experimental study confirmed the patterns evident in the observational study; males perched farther from shore than their oviposition site was located. Interestingly, in both the observational and the experimental study, when neighbors were close, a male perched away from his closest neighbor, which usually resulted in his oviposition site being closer to his neighbor than he was. Thus, male amberwings apparently alter their perch choice within their territories in response to the location of other males. These lookout post locations may provide the best opportunity for the territorial male to perceive passing females and intruding males, while minimizing conflict with their neighbors.     

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.     

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 antennal sensilla of the adult of Libellula depressa (Odonata: Libellulidae)

An ultrastructural investigation (SEM, TEM) on the antennal flagellum of the adult of the dragonfly Libellula depressa (Odonata:Libellulidae) revealed sensilla located in pits on the lateral-ventral side of the antenna. These sensilla are represented by sensilla coeloconica and by deeply sunken sensilla. The sensilla coeloconica are innervated by three unbranched dendrites, which enter the peg and show a dendrite sheath ending at the base of the peg. The peg has no socket and its cuticle is irregular with wide pore-like structures at the base of which actual pores are visible. The structure of these coeloconic sensilla is in agreement with that reported for single-walled insect chemoreceptors. The deeply sunken sensilla are represented by two kinds of sensilla styloconica, named type-1 and type-2, located at the bottom of deep cavities appearing as simple openings on the antennal surface. These sensilla are no-pore sensilla with inflexible socket and unbranched dendrites and, notwithstanding their structural differences, share common features typical of thermo/hygroreceptors. The presence of chemoreceptors in adult dragonflies sheds light on evolutionary trends in insect perception; the previously unknown occurrence of thermo/hygroreceptors in dragonflies is very important in view of the reported ability of Odonata to thermoregulate heliothermically.     

Energy expenditure and food intake of territorial male Pachydiplax longipennis (Odonata: Libellulidae)

ABSTRACT.

• 1 Field observations of males of the dragonfly, Pachydiplax longipennis (Burmeister), were used to determine how individuals of this species allocate energy to different activities during territory occupancy.
• 2 The effects of biological and physical factors on the species’daily activity pattern were examined. The proportion of time spent in flight was independent of temperature but increased asymptotically with increasing population density.
• 3 Measurements of assimilation efficiency and the quantity of faeces produced per day were used to calculate daily intake of food. An independent estimate of food consumption was derived from data on gut contents and clearance rate.
• 4 Food intake appears to exceed only slightly the energy required to maintain a territory, with little available for other activities. The activity pattern may be determined in part by the amount of energy available to individuals.