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.     

Mating tactics and male wing dimorphism in the damselfly,Mnais pruinosa costalis selys (Odonata: Calopterygidae)

Males of the damselfly,Mnais pruinosa costalis, exhibit wing color dimorphism: one form has orange wings, and the other hyaline wings which resemble female wings. The former is usually territorial and the latter uses sneaky mate securing tactics. When orange-winged males failed to establish territory, they became floaters that day. Hyaline-winged males perched around their territories and often, formed in tandem without any apparent courtship behavior when they found females. Their copulation frequency was higher and copulation duration longer than those of territorial males. A few females oviposited without remating. Total oviposition duration of females with which a hyaline-winged male mated was more than 32 min per male on average in a day Females that copulated with hyaline-winged males often remated with orange-winged residents before oviposition. Total duration of oviposition bouts of females after mating with floaters was short (15 min), while that with territorial residents was long (66 min). As a result, total oviposition duration of females with which an orange-winged male mated was about 40 min in a day. The reproductive success of the hyaline-winged males may be similar to that of the orange-winged males.     

Age-Specific Behavior and Habitat Selection of Adult Male Damselflies, Calopteryx maculata (Odonata: Calopterygidae)

We compared the age, movement, and time–activity budgets of male Calopteryx maculata damselflies occupying off-stream tree-fall gaps with those at stream sites within a 10-ha woodland. All males collected at off-stream sites were younger than males collected at stream sites—as indicated by their significantly higher wing transmittance. Thirty-three percent of teneral males marked at off-stream gaps moved to stream sites within 4 days (mean distance = 140 m), while mature males marked at stream sites never left the stream. In contrast to stream site males, off-stream males spent significantly more time capturing prey and never engaged in aggressive interactions with other males. Behavioral differences were not due to variations in the operative body temperature. However, malaise trapping revealed a greater frequency of suitable prey in forest light gaps. Our findings support the idea that teneral male Calopteryx leave their emergence sites along the stream for off-stream light gaps to forage without interference and build the energy reserves necessary to attain and hold streamside territories.     

Survival after experimental manipulation in the territorial damselfly Hetaerina titia (Odonata: Calopterygidae): more ornamented males are not more pathogen resistant

It has been hypothesized that sexual ornaments communicate pathogen resistance ability. We experimentally explored the relationship between the expression of a male ornamental trait (wing pigmentation) of the damselfly Hetaerina titia and survival after a bacterial challenge. We infected males with Serratia marcescens (a Gram-negative bacteria typical of insects) and compared survival against a group infected with dead bacteria and a noninfected group. Wing pigmentation was entered as a predictor of survival in this comparison. Our study indicated that wing pigmentation was not a good predictor of immune ability against bacteria. This result contradicts previous findings in the same and other calopterygid species in which wing pigmentation intensity inversely correlated with gregarine infection levels. It also contradicts the general idea that ornaments are honest indicators of pathogen defense.     

Age-related changes in flight muscle mass, lipid reserves and flight capacity during adult maturation in males of the territorial damselfly Calopteryx atrata (Odonata: Calopterygidae)

In the territorial damselfly Calopteryx atrata Selys, length of the hindwing, the wing areas and the aspect ratio did not differ significantly among age classes during the pre-reproductive period, while the body mass of males increased about 2.5 times. This is due primarily to increase in mass of thorax and abdomen. The flight muscle mass accounted for the great part of the thorax mass, and began to increase from early in the pre-reproductive period and continued increasing until sexual maturation. The average flight muscle mass of sexually matured males was about 2.4 times of that of the youngest immature ones. On the other hand, the abdomen mass and total lipids increased remarkably in the latter half of the pre-reproductive period. The average total lipid content of mature males was about tenfold of that of the youngest immature ones. The maximum lift production per flesh body mass was positively correlated with the flight muscle mass and total lipid content. Such an increase in flight muscle mass and lipid reserves resulted in the increase of maximum lift force, and probably enhanced flight performance.     

Male dimorphism, territoriality and mating success in the tropical damselfly, Paraphlebia zoe Selys (Odonata: Megapodagrionidae)

The tropical damselfly Paraphlebia zoe has two male morphs: a black-winged (BW) male which is associated with territorial defense of oviposition sites; and a hyaline-winged (HW) male similar in appearance to females, and, compared to the black morph, less frequently found defending territories. In a wild population of this species, we first assessed the relationship between phenotypic traits [male morph, size and territorial status (being territorial or non-territorial)], their role on mating success, and the degree to which a particular territory may contribute to male mating success. Second, to relate a physiological basis of being territorial we compared both morphs in terms of muscular fat reserves and thoracic muscle, two key traits related to territory defense ability. Males of both morphs defended territories although the BW males were more commonly found doing this. BW males were larger than HW males and size predicted being territorial but only within HW males (territorial males were larger) but not in BW males. Male mating success was related to territorial status (territorial males achieved a higher mating success), but not to morph or size. Furthermore, territory identity also explained mating success with some territories producing more matings than others. The BW morph stored more fat reserves which may explain why this morph was more likely to secure and defend a place than the HW morph. However, the HW morph showed higher relative muscle mass which we have interpreted as a flexible strategy to enable males to defend a territory. These results are distant to what has been found in another male dimorphic damselfly, Mnais pruinosa, where the advantage of the non-territorial morph relies on its longevity to compensate in mating benefits compared to the territorial morph.     

Remarkable iridescence in the hindwings of the damselfly Neurobasis chinensis chinensis (Linnaeus) (Zygoptera: Calopterygidae)

The bright green dorsal iridescence of the hindwings of Neurobasis chinensis chinensis males, very rare in Odonata, is known to play a significant role in their courtship behaviour. The mechanism responsible for such high contrast and spectrally pure colour has been investigated and found to be optical interference, producing structural colour from distinct laminations in the wing membrane cuticle. The ventral sides of these iridescent wings are dark brown in colour. In a single continuous membrane of wing cuticle, this is an effect that requires a specialized structure. It is accomplished through the presence of high optical absorption (kappa = 0.13) within two thick layers near the ventral surface of the wing, which leads to superior dorsal colour characteristics. By simultaneously fitting five sets of optical reflectivity and transmissivity spectra to theory, we were able to extract very accurate values of the complex refractive index for all three layer types present in the wing. The real parts of these are n = 1.47, 1.68 and 1.74. Although there is often similarly significant dorsal and ventral colour contrast in other structurally coloured natural systems, very few system designs comprise only a single continuous membrane.     

Male courtship success and female discrimination in the mediterranean fruit fly (Diptera: Tephritidae)

The mating success of individually marked male Mediterranean fruit flies was monitored over 6 consecutive days in the laboratory. Mating frequency was nonrandom, as the numbers of both males that failed to mate and males that mated many (more than four) times were much higher than expected by chance alone. Differential mating success resulted in part from intermale variation in activity level. Male copulatory success was positively correlated with the numbers of courtships performed, attempted copulations (mountings), and females courted. Male-male aggression, on the other hand, and a negligible effect on male mating success. Female choice also appeared to influence male mating frequency. Fewer than 10% of courtships resulted in mating, and in most cases females terminated courtship by simply moving away from the male. Females that did mate generally selected males having higher copulatory scores than previously rejected males.     

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.     

Seasonal dynamics, movements and the effects of experimentally increased female densities on a population of imaginal Calopteryx aequabilis (Odonata: Calopterygidae)

Abstract. 1. A population of Calopteryx aequabilis Say was sampled daily on a tributary of Canard River, Kings County, Nova Scotia, for the entire flight season in 1983 (29 May to 13 August), using capture–mark–recapture techniques.
2. 2701 sightings of 678 individuals were obtained along a 635 m segment of the stream. A maximum daily count of 174 imagines was reached on 11 June, after which the population gradually declined.
3. More females than males were marked but sexually mature males outnumbered females at the water on all but four days.
4. Immigration rather than local emergence accounted for a large proportion of the population after 20 June.
5. Females were consistently vagile; males were site–specific but occasionally moved long distances between captures.
6. Males and females first marked as tenerals became reproductively mature after about 5 days.
7. We experimentally increased female density on a partially isolated section of the study stream to see how increased female numbers affected the demographics and movement patterns of the population.
8. Residence times for introduced and resident females were similar. In contrast, during a similar introduction of males a year earlier, most introduced males disappeared quickly.
9. Males decreased the distance they travelled daily between captures, their total distance travelled and their range following the introduction, and females showed a tendency (not statistically significant) toward increased movements and dispersal, as predicted.     

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.     

Diurnal changes and frequency dependence in male mating preference for female morphs in the damselfly Ischnura senegalensis (Rambur) (Odonata: Coenagrionidae)

Ischnura senegalensis females exhibit color dimorphism, consisting of an andromorph and a gynomorph, which might be maintained under a frequency-dependent process of mating harassment by mate-searching males. Males change their mating preference for female morph depending on prior copulation experience. Binary choice experiments between two female morphs were carried out in four local populations in the early morning (07.00–09.00 hours) and the afternoon (12.00–14.00 hours), times which mark the onset and the end of diurnal mating activity, respectively. According to the line census along the water's edge, the proportion of andromorphs in the female population varied from 21 to 67% throughout the survey period for four local populations. Males showed non-biased preference for female morphs in the early morning in each local population, while they chose the common morph in the afternoon. Male mating preference for female morphs was positively correlated to the proportion of female morphs in the population. If the selective mating attacks on the common female morphs inhibit their foraging and/or oviposition behavior, frequency-dependent male mating attacks might provide a selective force for maintaining the female color dimorphism in I. senegalensis .     

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.     

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.     

Sensory trap as the mechanism of sexual selection in a damselfly genitalic trait (Insecta: Calopterygidae)

During copulation, males of some calopterygid damselfly species displace the sperm stored in the spermatheca: the male genital appendages enter into the spermathecal ducts and physically remove sperm. In Calopteryx haemorrhoidalis, the genital appendages are too wide to penetrate the spermathecae, but males use a different mechanism in which the aedeagus stimulates the vaginal sensilla that control spermathecal sperm release. Since these sensilla are used during egg fertilization and oviposition, it was hypothesized that this function evolved before the male stimulatory ability. I investigated this using Hetaerina cruentata, a species whose position in the Calopterygidae phylogeny is more basal than Calopteryx. Given this position and having determined that males of this species are not able to displace sperm of their conspecific females during copulation, it was expected that H. cruentata females would eject sperm when stimulated with the aedeagi of C. haemorrhoidalis but not when stimulated with the aedeagi of their conspecifics. This prediction was confirmed. In order to investigate the widespread nature of this result, some other Calopteryx species-Calopteryx xanthostoma and Calopteryx virgo-were investigated. The results were similar to those of H. cruentata: conspecific males were unable to stimulate their females, but females ejected sperm when stimulated with C. haemorrhoidalis aedeagi. Morphometric analysis suggests that the mechanistic explanation for the stimulatory ability of C. haemorrhoidalis genitalia is that the aedeagal region that makes contact with the vaginal sensilla is wider in C. haemorrhoidalis than in the other species. These results suggest that the sensory "bias" shown and shared by H. cruentata, Calopteryx splendens, C. virgo, and C. haemorrhoidalis females represents an ancestral condition and that the male stimulatory ability is absent in the evolutionary history of the clade. These pieces of evidence as well as another one presented elsewhere, which indicates that C. haemorrhoidalis males vary in their stimulatory ability, constitute the three criteria for a case of sexual selection via exploitation of a female sensory bias. These results also provide support to the sensory trap hypothesis that indicates that the female bias-in this case, egg fertilization and oviposition-evolved in a context different from sexual selection. Considering that the male genital appendages responsible for physically removing spermathecal sperm in other calopterygids are present in C. haemorrhoidalis, I suggest that males were once able to displace spermathecal sperm physically. Such ability may have been later impeded by a reduction in size of the spermathecal ducts. Possibly, one of the latest events in this sequence is the male's stimulatory ability. This hypothetical series of events suggests a coevolutionary scenario in which the central actor is the sperm stored in the spermathecae.     

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.