The effects of temperature and prey density on the development rates and growth of damselfly larvae (Odonata: Zygoptera)

Abstract. 1. The effects of prey density and temperature on the feeding and development rates of several late instars of the larvae of three common European damselflies ( Lestes sponsa (Hansemann), Coenagrion puella (L.) and Ischnura elegans van der Linden) were investigated in a laboratory experiment.
2. Functional responses were used to estimate maximum feeding rates. Maximum feeding rates were compared between species, instars and temperatures by expressing prey consumption in terms of prey biomass consumed per unit predator biomass. Lestes was capable of feeding at almost twice the rate of either Coenagrion or Ischnura.
3. Higher feeding rates led to faster development rates and there was an interaction between species and temperature. With the exception of those with very low feeding rates, larvae maintained at higher temperatures, but similar feeding rates, developed faster. Under similar conditions of temperature and feeding rate, Lestes larvae developed faster than larvae of either Coenagrion or Ischnura.
4. Faster development rates at similar rates of food intake were achieved at the cost of reduced size-increases between instars.
5. The differences in the responses of the three species are discussed in the light of their respective life histories, and with reference to a recent model of population regulation in damselflies.     

Functional morphology of the male caudal appendages of the damselfly Ischnura elegans (Zygoptera: Coenagrionidae)

Odonata are usually regarded as one of the most ancient extant lineages of winged insects. Their copulatory apparatus and mating behavior are unique among insects. Male damselflies use their caudal appendages to clasp the female's prothorax during both copulation and egg-laying and have a secondary copulatory apparatus for sperm transfer. Knowledge of the functional morphology of the male caudal appendages is the basis for understanding the evolution of these structures in Odonata and respective organs in other insects. However, it is still not exactly known how the zygopteran claspers work. In this study, we applied micro-computed tomography and a variety of microscopy techniques to examine the morphology, surface microstructure, cuticle material composition and muscle topography of the male caudal appendages of Ischnura elegans. The results indicate that the closing of the paraproctal claspers is mainly passive. This indirect closing mechanism is very likely supported by high proportions of the elastic protein resilin present in the cuticle of the paraproctal bases. In addition, the prothoracic morphology of the female plays an important role in the indirect closing of the male claspers. Our data indicate that both structures – the male claspers and the female prothoracic hump – function together like a snap-fastener.     

Emergence of the damselfly Pyrrhosoma nymphula (Sulzer) (Zygoptera: Coenagrionidae) from two adjacent ponds in northern England

Emergence of the damselfly Pyrrhosoma nymphula was followed closely at two similar and adjacent ponds in northern England. Males emerged earlier than females at both ponds (significantly so at one). The sex ratio was significantly male biased at both ponds. Size of emerging adults declined through the emergence period at both ponds. Adults emerging from one pond did so significantly earlier and were significantly larger than those from the other pond. The density of larvae was approximately twice as high in the pond from which larvae emerged early; this pond was also slightly deeper and usually achieved higher maximum daily water temperatures. These findings are discussed in the light of the conventional view of seasonal regulation in a spring-emerging damselfly.     

Variable microsatellite loci isolated from the azure damselfly,Coenagrion puella (L.) (Zygoptera; Coenagrionidae)

We isolated and characterized 10 polymorphic microsatellite loci from the azure damselfly Coenagrion puella (Zygoptera; Coenagrionidae) as part of a study assessing reproductive success and genetic structure in an isolated population of this species. Levels of genetic diversity were assessed in 50 individuals collected from Queen Elizabeth Country Park, Hampshire, UK. The number of alleles per microsatellite loci ranged from three to 22 and the observed and expected heterozygosities varied between 0.26 and 0.84 and between 0.23 and 0.91, respectively. Two loci showed significant (P < 0.05) heterozygote deficits, likely because of null (non‐amplifying) alleles; one pair of loci was in linkage disequilibrium.     

The natural prey of larvae of the damselfly, Ischnura elegans (Odonata: Zygoptera)

SUMMARY. A method for determining the prey of a damselfly larva under natural conditions is described, with its shortcomings.
Within the limits of the method, it would appear that Ischnura elegans consumes different prey items in roughly the same proportions as their occurrence in the field.
Anomalous results are explained in terms of differences in encounter rates between predator and prey and/or ease with which prey can be handled.     

Fluctuating asymmetry and lifetime mating success are correlated in males of the damselfly Coenagrion puella (Odonata: Coenagrionidae)

Abstract.

• 1 The influence of fluctuating asymmetry on lifetime mating success of males of the damselfly Coenagrion puella (Odonata: Coenagrionidae) was investigated. Fluctuating asymmetry was measured as the difference in length of left and right fore and hind wings.
• 2 Males with more symmetrical wings enjoyed higher lifetime mating success.
• 3 Larger males, in contrast to a previous study of this species, also had higher mating success. This may be attributed to differences in the weather conditions prevailing at the time of the studies.

     

Hybridization between Ischnura graellsii (Vander Linder) and 2. elegans (Rambur) (Odonata: Coenagrionidae): are they different species?

Two closely related damselflies, Ischnura graellsii and I. elegans , were analysed for morphological differences and reproductive isolation in the north coast of Galicia (NW Spain). We compared animals from sympatric and allopatric localities, including I. elegans from Belgium and I. graellsii from southern Spain as pure allopatric populations. A set of morphometric characters were studied by means of multivariate discriminant analysis to determine if these two species can be unambiguously distinguished. Discriminant analysis revealed that I. graellsii and I. elegans are well differentiated on the first two axis (86% and 11%, respectively). I. graellsii individuals are distinguished from I. elegans by their smaller size and, specifically, by their narrower and shorter wings and shorter tibiae. In addition, I. elegans has a narrower space between the branches of each cercus, and greater distance between the branches of each paraproct. Sympatric individuals are morphologically intermediate, suggesting hybridization. When the species were put together in the laboratory, they showed partial temporal separation in mating behaviour, but males of I. elegans readily mated with females of I. graellsii , and hybrid individuals were obtained. The opposite heterospecific cross was almost impossible, apparently because of mechanical problems with the tandem linkage. Laboratory-reared hybrids (from male I. elegans × female I. graellsii ) are morphologically intermediate, mainly resembling the maternal phenotype. Although hybridization between both taxa is common, we suggest maintaining the specific status for both phenotypes because they show incipient reproductive isolation, as it is reported in the literature. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 225–235.     

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.     

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.     

The costs of reduced feeding due to predator avoidance: potential effects on growth and fitness in Ischnura elegans larvae (Odonata: Zygoptera)

ABSTRACT.

• 1 This paper investigates the behaviour, in the laboratory, of a forager simultaneously confronted with the conflicting needs to feed and to avoid predators. The foragers were larvae of the damselfly Ischnura elegans Van der Linden, feeding on Daphnia magna Strauss. The predators were adult females of Notonecta glauca L.
• 2 Patch choice by Ischnura larvae was significantly modified by the presence of predators. Larvae moved to feed in patches of high prey density when predators were absent but preferred dense cover, even though virtually no prey were available, when predators were present. This behaviour was not altered by hunger, up to 12 days without food. In other words, Ischnura larvae were risk averse in their foraging behaviour.
• 3 In experiments with abundant prey available, the feeding rates of Ischnura larvae confined to a single patch were also significantly reduced by the presence of hydrodynamically and chemically detectable predators. Predators detectable only by vision had little effect.
• 4 Calculations made from published data show that reduced larval feeding rates can lead to slower growth and development and prolonged instar durations in Ischnura elegans larvae. This may have important consequences for larval survival and adult reproductive fitness.

     

Evidence for widespread sperm displacement ability among Zygoptera (Odonata) and the means for predicting its presence

Males of the coenagrionid damselflies Argia moesta, A. sedula and hchnura ramburii use similar penis morphology to remove and/or reposition sperm of previous males from the storage organs of females prior to inseminating them. Although the species vary in the degree to which sperm is removed from or packed into the spermatheca, in all three species, sperm is removed from the bursa copulatrix. Since sperm in the bursa probably has priority in fertilizing eggs in at least the first oviposition after mating, sperm precedence can be estimated as the percentage of sperm (by volume) in the bursa belonging to the last male to mate. Estimated sperm precedence for these species is approximately 71% for Argia sedula, 82% for I. ramburii and 93% for A. moesta. These results, combined with similar ones for other damselflies clearly indicate that the ability to displace sperm may be widespread among temperate-zone Zygoptera. Species with each of the four major variations in damselfly penis structure have now been shown to displace sperm using this morphology. The systematic distribution of these major variants suggests several origins of sperm displacement ability within the Zygoptera. Whether or not all damselflies are capable of sperm displacement depends on both the presence of micro-structures used in sperm removal or repositioning and on the presence of sperm of previous males in mating females. It is possible, therefore, to predict that sperm displacement occurs in a damselfly if (1) females mate more than once, (2) mating females store sperm in organs accessible to penis morphology, (3) the distal segment of the male penis has structures similar to those known to be involved in sperm removal or repositioning, and (4) oviposition occurs in tandem or with the male non-contact guarding his mate.     

Pre- and post-maturation survival in adults of the damselfly Pyrrhosoma nymphula (Zygoptera: Coenagrionidae)

The technique of mark-release-recapture was used to study survival before and after sexual maturity in adults of the damselfly Pyrrhosoma nymphula (Sulzer). Fewer females were recaptured upon return to water to breed despite no differences in dispersal or daily survival rate between the sexes over the immature period. Because females took longer to mature than males, their poorer recapture rate was attributed to greater overall mortality during their longer maturation phase. Survivorship curves for tenerals marked at emergence suggested that overall survival of immature adults was similar to, if not better than, that of mature adults. The reasons for this are discussed.
Jolly's model was used to estimate daily survival rates for mature adults. The assumptions of the model were tested rigorously. Estimates for females were statistically less reliable than those for males. Mean reproductive spans for males and females were 6.8 and 6.6 days, respectively, giving mean total adult lifespans of 19.4 days and 21.6 days for individuals surviving the maturation period.
Because neither sex visited the breeding site every day, sampling exclusively at water resulted in underestimation of mean reproductive spans for both sexes. Female reproductive spans were underestimated to a greater extent; because females remain away from water longer between visits, there is a greater chance that they will die before being recaptured.
Mean reproductive spans were also underestimated when only a sub-section of the habitat was sampled. Females were significantly more mobile than males and this increased the likelihood that they would move out of the study area, resulting in more severe underestimation. The importance of obtaining accurate estimates of mature lifespan for females is discussed.     

The effect of invertebrate and vertebrate predators on the foraging movements of Ischnura elegans larvae (Odonata: Zygoptera)

SUMMARY.

• 1 The foraging movements of late instar Ischnura elegans larvae were monitored in laboratory experiments to study the effects of predators on larval feeding behaviour.
• 2 Ischnura larvae are sit-atid-wait, or ambush, foragers, moving occasionally between perches in search of profitable feeding sites. Larval foraging movements, monitored at different densities of Daphnia prey, increased significantly when prey were absent.
• 3 In experiments without prey, larval movement was inhibited by the presence of fish predators, as well as by invertebrate predators (Notonecta glauca), but not by closely related, non-predatory invertebrates (Corixa punctata) or physical disturbance of the water (intermittent air bubbles).
• 4 Further experiments varied Ischnura hunger levels (0–8 days without food) and illumination (light or dark) with and without notonectid predators. Hunger had no consistent effect on penultimate instar behaviour but final instar foraging activity was significantly modified: movements increased after 4 days starvation and decreased again after 8 days. This response was suppressed by the presence of predators. Both larval instars moved significantly less often in the light, even when predators were absent.
• 5 These phenotypically flexible predator-avoidance responses are likely to decrease the risk of predation by both visual and tactile predators. However, predators clearly have an important influence on the feeding niche of Ischnura larvae, and may decrease the overall feeding efficiency, growth rate, and survival of larvae by constraining their movement in search of profitable feeding sites.

     

Phylogeny and taxonomy of the damselfly genus Enallagma and related taxa (Odonata: Zygoptera: Coenagrionidae)

Abstract The zygopteran genus Enallagma has been the subject of numerous behavioural and ecological studies, but phylogenetic relationships among species have been examined only within eastern North America, and even the composition and diagnosis of the genus are unclear on a world-wide basis. Most authorities currently recognize about seventy species within Enallagma , comprising two major radiations, in North America and Africa. This study, using morphological data, demonstrates that the North American and a few related Palaearctic species form a monophyletic group that is quite distinct from the African species. The latter are themselves divided into at least three, and probably four, separate clades, one of which may be related to E. parvum of India. Consequently, three of Kennedy's long disused genera, Africallagma , Amphiallagma and Proischnura ( Kennedy, 1920 ) are resurrected and two new genera, Azuragrion gen.n. and Pinheyagrion gen.n. are established for the remaining African taxa. Finally, Enallagma is divided into two subgenera, Enallagma s.s ., the typical 'bluets', including many North American, Holarctic and Palaearctic species, and Chromatallagma subgen.n., comprising a group of species of more variable colour that is confined to North America, the Caribbean and northernmost South America.     

Correlated morphological and colour differences among females of the damselfly Ischnura elegans

Abstract 1. The female‐limited colour polymorphic damselfly Ischnura elegans has proven to be an interesting study organism both as an example of female sexual polymorphism, and in the context of the evolution of colour polymorphism, as a model of speciation processes. 2. Previous research suggests the existence of correlations between colour morph and other phenotypic traits, and the different female morphs in I. elegans may be pursuing alternative phenotypically integrated strategies. However, previous research on morphological differences in southern Swedish individuals of this species was only carried out on laboratory‐raised offspring from a single population, leaving open the question of how widespread such differences are. 3. The present study therefore analysed multi‐generational data from 12 populations, investigating morphological differences between the female morphs in the field, differences in the pattern of phenotypic integration between morphs, and quantified selection on morphological traits. 4. It was found that consistent morphological differences indeed existed between the morphs across populations, confirming that the previously observed differences were not simply a laboratory artefact. It was also found, somewhat surprisingly, that despite the existence of sexual dimorphism in body size and shape, patterns of phenotypic integration differed most between the morphs and not between the sexes. Finally, linear selection gradients showed that female morphology affected fecundity differently between the morphs. 5. We discuss the relevance of these results to the male mimicry hypothesis and to the existence of potential ecological differences between the morphs.     

On the biology of the damselfly Nosoticta kalumburu Watson & Theischinger (Zygoptera: Protoneuridae)

The habitat and the territorial and reproductive behaviours of a recently named Australian protoneurid damselfly, Nososticta kulumburu Watson & Theischinger are described. Nososticta kalumburu breeds in shallow, narrow (<2 m wide) fast-flowing permanent streams in the Kimberley region of Western Australia. The presence of suitable perches (protruding rocks or overhanging vegetation) seems to be important in determining its distribution. It is a sexually dimorphic species; males have prominent dark brown spots on the wings and a blue pruinescence, which the females lack. Males are strongly territorial. Territories (radius around I m) are small in both high and low density populations and are defended vigorously against conspecific males. Male courtship behaviour towards potential mates involves a vigorous hovering flight in front of, and by the side of, the female, during which the prominent dark brown spots on all four wings are dispalyed. Mean length of copulation is 15.4 min. Two stages of copulation were recognized. It is thought that the first stage which occupies most time (94.7%) is concerned with removing sperm from previous matings in the usual zygopteran manner. Oviposition begins in tandem in the stems of aquatic macrophytes or the roots of terrestrial plants that hang into thc water. Males exhibit plasticity in post-copulatory guarding bchaviour: in pairs undisturbed during the early stages of oviposition, the male may release the female before the current bout of egg-laying has been completed and attempt to regain a territory. The penis is unusual in having a heavily armoured shaft and two pairs of cornua; it is probably used in sperm removal as well as intromission.     

A quantitative study of mortality at emergence in the damselfly Pyrrhosoma nymphula (Sulzer) (Zygoptera: Coenagrionidae)

SUMMARY.

• 1 Emergence of the damselfly Pyrrhosoma nymphula was studied over three seasons in two ponds in northern England.
• 2 Numbers emerging were significantly negatively correlated with temperature and atmospheric pressure.
• 3 Overall mortality at emergence was 28%. of which predators (largely birds, ants and spiders) accounted for 22% and climatic factors 6%.
• 4 Daily mortality estimates were significantly positively correlated with precipitation.
• 5 There was no evidence of density dependent mortality at emergence.