Changes in reproductive behavior in adult damselfly Calopteryx splendens (Odonata: Calopterygidae) in response to flood

The reproductive behavior of adult Calopteryx splendens males and females inhabiting the Nida River, south Poland, was studied and compared during a pre‐flood and a post‐flood year. The flood disturbance in 2010 caused a decrease in aquatic macrophytes, thus reducing availability of potential territories and consequently, significantly influencing male behavior towards a frequent non‐territorial strategy. Many males in the post‐flood population had damaged wings due to extremely aggressive contests. Male–male tandems were commonly observed; this is an uncommon behavior in C. splendens. Although the sex ratio was male‐biased throughout the whole study, we observed more males in the post‐flood year. We also observed less‐frequent copulations and ovipositions during the post‐flood year. The only unchanged characteristic was population density, which did not differ before and after the flood disturbance. Floods have significant impact on damselfly reproductive sites and this, due to changes in behavior and sex ratio, may result in further consequences on population dynamics.     

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.     

Responses of rectal pumping to oxygen lack by larval Calopteryx splendens (Zygoptera: Odonata)

Abstract. Larvae of most stadia of Calopteryx splendens (Harris) have been shown to pump water in and out of the rectum for much of the time. In each cycle of pumping there are from three to seven inhalant stokes which progressively fill the ileum with water. A short pause intervenes before a strong compression of segments 5–8 expels the water in a jet. The pattern resembles the gulping ventilation of aeshnid and libellulid dragonfly larvae. Larvae made hypoxic or otherwise stressed show a rise in the frequency and amplitude of pumping. The increase in frequency is brought about by a reduction of the pause duration, while the gain of amplitude results from a rise in both the number of inhalations/cycle and the magnitude of the stroke volume of each inhalation. Some evidence concerning the role of stretch receptors in the execution of the cycle is presented. It is proposed that rectal pumping promotes the exchange of ions and dissolved gases between the rectal water and the hindgut epithelia, supplying mainly the chloride-absorbing cells in the rectal pads. Pumping also stirs the water in the vicinity of the caudal lamellae which are probably the insect's major sites of gaseous exchange.     

Habitat choice as the defensive behavior of Calopteryx splendens Harris larvae (Odonata, Calopterygidae): Impact of the predator species and of the population density of damselflies

Laboratory experiments have demonstrated that the change in the height of dispersal of Calopteryx splendens Harris larvae on perches is a protective strategy in the presence of fish. Specificity of protective behavior of damselflies in the presence of pelagic or bottom-dwelling fishes, or of a complex of both types of fishes is described. The effectiveness of the habitat choice is reduced in the presence of the complex of fishes. The use of an optimal protective strategy by all the larvae was observed only in the case when their population density was the lowest. The increase in the population density resulted in the increased importance of intraspecies competition (cannibalism) and decreased effectiveness of protective behavior against heterospecific predators.     

Incidence of Anagrus obscurus (Hymenoptera: Mymaridae) egg parasitism on Calopteryx haemorrhoidalis and Platycnemis pennipes (Odonata: Calopterygidae: Platycnemididae) in Italy

Very little is known about the incidence of egg parasitoids in odonates, perhaps because Odonata eggs are well protected by stems or leaves, sometimes below water. In Central Italy (Pontecorvo, Frosinone Province) two damselflies, Calopteryx haemorrhoidalis and Platycnemis pennipes, occur in high densities. In August 2007 we collected 30 stems of the aquatic plant Potamogeton sp. used as substrate for oviposition and incubated eggs in the laboratory. Most stems (24 for C. haemorrhoidalis and 23 for P. pennipes) contained Odonata eggs. Parasitoids emerged from 12 stems, with a mean parasitism of 2% for C. haemorrhoidalis and 6% for P. pennipes, and a maximum of 14% and 50%, respectively. Furthermore, we observed egg‐laying of 19 females of C. haemorrhoidalis and 11 of P. pennipes, and marked the stems where oviposition was observed. Clutches remained in the river for five days and were then collected and incubated. Parasitoids emerged from 11 of 30 stems, with an average parasitism of 8% for C. haemorrhoidalis and 3% for P. pennipes (maximums of 50% and 29%, respectively). All parasitoids belonged to the family Mymaridae, and were identified as Anagrus (Anagrus) obscurus Förster, 1861, sensu Soyka, 1955 . This is the first time that this species is described as an egg parasitoid of odonates, and that the egg parasitoid of C. haemorrhoidalis and P. pennipes is identified. Our data suggest that egg parasitism might be a significant selective factor for both odonates in the studied locality, affecting female oviposition behavior.     

Resource defense polygyny by Hetaerina rosea Selys (Odonata: Calopterygidae): influence of age and wing pigmentation

Current evidence suggests that in Hetaerina damselflies males exhibit lek mating system. In this study, in order to answer if the same occurs in Hetaerina rosea Selys, we manipulated vegetation substrates used as territories and quantified the number of visiting females, males defending territories and fight intensity. We also examined whether body size and wing pigmentation are selectable traits in male-male competition, and if age affects male territorial behavior. Our results showed that males with larger pigmented areas won more contests, independently of body size. Old males changed from territoriality to sneaking strategy. Contrary to other Hetaerina species, males of H. rosea do not display lek behavior, but defend resources according to the resource defense polygyny strategy.     

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.     

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.     

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.     

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.     

Body temperature regulation is associated with climatic and geographical variables but not wing pigmentation in two rubyspot damselflies (Odonata: Calopterygidae)

It has been proposed that wing pigmented spots function in temperature control in male calopterygids. Using two rubyspot species Hetaerina americana Fabricius and Hetaerina vulnerata Hagen in Selys, the present study investigated whether (i) wing spot size and colour‐modified aspect can predict temperature gain after a cooling event; (ii) wing spot size is related to the temperature needed to fly and how long it takes to initiate flight; and (iii) wing spot size is related to seasonality and altitude. The results obtained do not support any of these relationships. The results also indicate that H. vulnerata can achieve flight at 8 °C less than H. americana. The present study further investigates whether the species differ in their latitudinal and geographical distribution, and respond differently to bioclimatic variables. The results obtained provide support for this particular hypothesis, showing that H. vulnerata inhabits higher altitudes, and is able to tolerate colder environments compared with H. americana. Wing spots in the two Hetaerina species do not help in thermoregulation, although both species show different temperature control abilities. This difference in thermoregulatory ability may enable the species to colonize different environments and reduce interspecific competition.     

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.     

The Evolution of Wing Shape in Ornamented-Winged Damselflies (Calopterygidae, Odonata)

Flight has conferred an extraordinary advantage to some groups of animals. Wing shape is directly related to flight performance and evolves in response to multiple selective pressures. In some species, wings have ornaments such as pigmented patches that are sexually selected. Since organisms with pigmented wings need to display the ornament while flying in an optimal way, we might expect a correlative evolution between the wing ornament and wing shape. We examined males from 36 taxa of calopterygid damselflies that differ in wing pigmentation, which is used in sexual displays. We used geometric morphometrics and phylogenetic comparative approaches to analyse whether wing shape and wing pigmentation show correlated evolution. We found that wing pigmentation is associated with certain wing shapes that probably increase the quality of the signal: wings being broader where the pigmentation is located. Our results also showed correlated evolution between wing pigmentation and wing shape in hind wings, but not in front wings, probably because hind wings are more involved in signalling than front wings. The results imply that the evolution of diversity in wing pigmentations and behavioural sexual displays might be an important driver of speciation due to important pre-copulatory selective pressures.     

Sex matters: Otolith shape and genomic variation in deacon rockfish (Sebastes diaconus)

Little is known about intraspecific variation within the deacon rockfish (Sebastes diaconus), a recently described species found in the northeast Pacific Ocean. We investigated population structure among fish sampled from two nearshore reefs (Siletz Reef and Seal Rock) and one offshore site (Stonewall Bank) within a <50‐km² area off the Oregon coast. Fish from the three sample sites exhibited small but statistically significant differences based on genetic variation at >15,000 neutral loci, whether analyzed independently or classified into nearshore and offshore groups. Male and females were readily distinguished using genetic data and 92 outlier loci were associated with sex, potentially indicating differential selection between males and females. Morphometric results indicated that there was significant secondary sexual dimorphism in otolith shape, but further sampling is required to disentangle potential confounding influence of age. This study is the first step toward understanding intraspecific variation within the deacon rockfish and the potential management implications. Since differentiation among the three sample sites was small, we consider the results to be suggestive of a single stock. However, future studies should evaluate how the stock is affected by differences in sex, age, and gene flow between the nearshore and offshore environments.     

Post-copulatory mate guarding by males of the demselfly Hetaerina vulnerata Selys (Odonata: Calopterygidae)

Males of the calopterygid damselfly Hetaerina vulnerata remain with their mates after copulating with them. The species exhibits two unusual features of post-copulatory mate guarding. First, a male will often leave his territory to accompany a female in tandem on a search for oviposition sites elsewhere. Second, a male will perch near his ovipositing female even though she completely submerges when egg-laying and cannot be captured and mated by another male while she is underwater. These activities carry two potential costs: (1) a male may miss other receptive females while guarding one mate and (2) he may lose his territory to an interloper while he is absent. These costs were low, however, because territorial males secured only one mating per 3.6 days on average. Moreover, 23 times out of 26, territorial males reclaimed their plots quickly after being away for 30–60 min. The gain from postcopulatory guarding came from being present to recapture a female should she fly up from the water after rejecting an oviposition site. There was a 40% chance that a female would leave one site to search for another during an oviposition bout. If the male were not present, his mate would be captured and mated by another individual (no female ever selected an oviposition site without being carried to it by a male). Her new partner would fertilize the remaining eggs in the female's clutch (if sperm precedence occurs in this species). The total number of eggs fertilized by a male will be affected by how well he prevents any one mate from copulating again before she lays her entire clutch and the total number of receptive females he captures. The variation in the degree of mate guarding by male odonates seems to be the evolutionary outcome of differences in fitness gains derived from these two competing activities in different ecological settings.