Test for latitudinal variation of life history,behavior and mortality in the strictly univoltine damselfly Sympecma fusca (Zygoptera: Lestidae)

A wealth of evidence shows that combinations of ecological stressors interact in shaping life history traits, but little is known about how ecological stressors combine with different seasonal time constraints to shape life history, behavior and mortality across populations. We studied life history, behavior and mortality rate in two latitudinally distant populations of the strictly univoltine, adult‐overwintering damselfly Sympecma fusca. Results from laboratory common‐garden and outdoor experiments indicated countergradient variation of larval development time and growth rate: the more time‐constrained larvae showed faster development and a higher growth rate. This finding led to larger size at emergence in the more time‐constrained individuals. Under conditions of intraspecific interaction (outdoor experiment), northern individuals showed lower survival than southern ones, presumably due to cannibalism. In the absence of intraspecific interactions (laboratory experiment), northern and southern larvae did not differ in survival. Finally, laboratory‐grown northern and southern larvae did not differ in activity level. This is the first time that compensation for seasonal time constraints has been shown in a temperate odonate species that overwinters in the adult stage.     

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.     

First Baltic amber megapodagrionid damselfly (Odonata: Zygoptera)

Electropodagrion szwedoi n. gen., n. sp., first Baltic amber megapodagrionid damselfly, is described. The European and North American fossils document a very high diversity and a much wider distribution of this group of damselflies during the Cenozoic than today. A checklist of described fossil species of damselflies of the family Megapodagrionidae is given.     

Mechanical properties of the endophytic ovipositor in damselflies (Zygoptera, Odonata) and their oviposition substrates

Damselfly females use their ovipositor valves to saw aquatic plants in order to insert their eggs into the plant tissues. Stiffness of the plant substrata is therefore an important parameter for oviposition substrate choice by females. Using a force transducer combined with a motorised micromanipulator, the bending stiffness of the ovipositor at the axial compressional load was studied in seven European damselfly species and compared to the local stiffness of seven preferred plant substrates. The puncture force of tested plant samples ranged from 105 to 1500 mN, and their local stiffness ranged from 208 to 1776 N/m. The bending stiffness of the ovipositor was estimated as 173-409 N/m depending on the damselfly species. Using original and literature data, a significant positive correlation between mechanical properties of the ovipositor and preferred oviposition substrates was demonstrated. Possible behavioural adaptations to overcome high stiffness of plant tissues during oviposition are discussed.     

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.     

The fine structure of sense organs in the ovipositor of the parasitic wasp, Orgilus lepidus Muesebeck

Three types of sensilla were observed in the ovipositor, including a multicellular sensillum presumed to respond to both chemical and mechanical stimuli, plus two types of campaniform sensilla. Four or five bipolar chemosensory cells innervate each multicellular sensillum, witln the dendrites terminating at an 800 ,Å dia. pore in the cuticular wall. The dendrite of an associated mechanosensory neuron is inserted upon a slender shaft of cuticle which extends inward from the wall of the ovipositor. This mechanosensory neuron may he activated by stretching when the ovipositor is bent. The dendrite of each campaniform sensillum ends in a cavity in the wall of the ovipositor, and are probably activated by stresses and vibrations as the wasp probes for a host. Sensilla of each type are present in the medial and lateral stylets of the ovipositor. Earlier behavioral studies indicated that the parasite probably uses these sense organs to locate hosts and distinguish healthy from already parasitized hosts.     

Redefining the damselfly families: a comprehensive molecular phylogeny of Zygoptera (Odonata)

An extensive molecular phylogenetic reconstruction of the suborder Zygoptera of the Odonata is presented, based on mitochondrial (16S, COI) and nuclear (28S) data of 59% of the 310 genera recognized and all (suspected) families except the monotypic Hemiphlebiidae. A partial reclassification is proposed, incorporating morphological characters. Many traditional families are recovered as monophyletic, but reorganization of the superfamily Coenagrionoidea into three families is proposed: Isostictidae, Platycnemididae and Coenagrionidae. Archboldargia Lieftinck, Hylaeargia Lieftinck, Palaiargia Förster, Papuargia Lieftinck and Onychargia Selys are transferred from Coenagrionidae to Platycnemididae, and Leptocnemis Selys, Oreocnemis Pinhey and Thaumatagrion Lieftinck from Platycnemididae to Coenagrionidae. Each geographically well‐defined clade of Platycnemididae is recognized as a subfamily, and thus Disparoneurinae (i.e. Old World ‘Protoneuridae’) is incorporated, Calicnemiinae is restricted, and Allocnemidinae (type genus: Allocnemis Selys) subfam.n ., Idiocnemidinae (type genus: Idiocnemis Selys) subfam.n . and Onychargiinae (type genus: Onychargia Selys) subfam.n . and Coperini trib.n . (type genus: Copera Kirby) are described. Half of Coenagrionidae belongs to a well‐supported clade incorporating Coenagrion Kirby and the potential subfamilies Agriocnemidinae, Ischnurinae and Pseudagrioninae. The remainder is less well defined, but includes the Pseudostigmatidae and New World Protoneuridae that, with Argiinae and Teinobasinae, may prove valid subfamilies with further evidence. Ninety‐two per cent of the genera formerly included in the polyphyletic Amphipterygidae and Megapodagrionidae were studied. Pentaphlebiidae, Rimanellidae and Devadattidae fam.n . (type genus: Devadatta Kirby) are separated from Amphipterygidae, and Argiolestidae, Heteragrionidae, Hypolestidae, Philogeniidae, Philosinidae and Thaumatoneuridae from Megapodagrionidae. Eight further groups formerly placed in the latter are identified, but are retained as incertae sedis; the validity of Lestoideidae, Philogangidae and Pseudolestidae is confirmed. For some families (e.g. Calopterygidae, Chlorocyphidae) a further subdivision is possible; Protostictinae subfam.n . (type genus: Protosticta Selys) is introduced in Platystictidae. Numerous new combinations are proposed in the Supporting Information. Many long‐established families lack strong morphological apomorphies. In particular, venation is incongruent with molecular results, stressing the need to review fossil Odonata taxonomy: once defined by the reduction of the anal vein, Protoneuridae dissolves completely into six clades from five families.     

Spatial distribution of mate-searching males in the damselfly,Cercion c. calamorum (Odonata: Zygoptera)

Both sexes of the coenagrionid damselflyCercion c. calamorum meet at, or near, ponds for oviposition. Spatial distribution of mate-searching males at a study pond was established with the following 2 simple and independent rules: (1) males don't go far from the roosting sites of the previous night; (2) once males reach one of the oviposition sites, the length of their stay is related to site quality. The first rule may correspond to the decrease in the number of receptive females at greater distances from the shore, and the second rule may relate to the higher predictability of female arrival at better oviposition sites. Contribution from the Laboratory of Animal Ecology, Kyoto University, No. 543.     

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.     

Central projections from contact chemoreceptors of the locust ovipositor and adjacent cuticle

Contact chemoreceptors (basiconic sensilla) located on the ovipositor and genital segments of the locust serve to control the chemical features of the substrate before and during oviposition. They occur dispersed and also crowded in fields between mechanosensory exteroceptors sensitive to touch or wind (trichoid and filiform sensilla). The central nervous projections of the four chemosensory and one mechanosensory neurons from single basiconic sensilla were stained selectively, focusing on receptors on the ovipositor valves, which usually contact the substrate during the pre-oviposition probing movements. All axons and neurites from one contact chemoreceptor usually stay close together in most of their projections. Segregation occurs mainly when single axons terminate in one neuromere while the others proceed to a different neuromere or ganglion. For projections from one chemoreceptor, there is evidence neither for functional segregation of mechanosensory from chemosensory afferent terminals nor for specific segregation between different chemosensory afferents. The projections from sensilla of dorsal cuticle tend to project rather uniformly along the midline of the terminal ganglion. Comparative staining of touch- and wind-sensitive hair receptor neurons shows mostly central projections, similar to those of neighbouring contact chemoreceptors. From the typical intersegmental projections of most primary afferents and from the lack of segregation into glomerular structures, we conclude that integration of chemosensory information from the genital segments is distributed in the terminal and the 7th abdominal ganglion.     

Asymmetric intraspecific competition among larvae of the damselfly Ischnura elegans (Zygoptera: Coenagrionidae)

Abstract. 1. A laboratory competition experiment is described in which the growth and development rates of larvae of the damselfly Ischnura elegans (Lind.) were measured over an entire instar.
2. Two larval instars which commonly occur together in the field were used in the experiment; they were maintained with a superabundance of prey and either larvae from the same or the larger/smaller instar.
3. Small larvae suffered increased development times and decreased size increases at the moult in the presence of large larvae but similar interference effects were not evident when these smaller larvae were in the presence of other small larvae.
4. Development time and size increases of large larvae were not significantly affected by the presence of small larvae.
5. Irrespective of the instar combinations investigated, interference effects were reduced when there were more perches available, although in only a few cases was this reduction significant.
6. The consequences of the asymmetric competition reported in the experiment for the study of lifetime reproductive success in damselflies are discussed. Late emerging adults may incur reduced reproductive success.     

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.     

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 ecological role of caudal lamellae loss in the larval damselfly,Ischnura posita (Hagen) (Odonata: Zygoptera)

Summary Damselfly larvae may autotomize and regenerate any of their 3 caudal lamellae. At least one missing or regenerating lamella was evident in 50.1% of field collected Ischnura posita larvae. Lamellae loss during molting is very infrequent (1 out of 117 recorded molts). Laboratory trials indicate that conspecifics remove lamellae and that this process is density dependent. The percentage of larvae losing lamellae during 24 h trials ranged from 73.5 at the highest density tested to 17.3 at the lowest density. I. posita larvae are cannibalistic. The presence of lamellae reduces an individual's chance of being cannibalized. More than twice as many final instar lamellae-less larvae were cannibalized during 24 h trials than analogous individuals having 3 lamellae at experimental initiation. Costs are also associated with lamellae autotomy. 1) Although individuals without lamellae can swim they are more reluctant to release from a wooden stalk and swim when threatened (9% release) than are larvae with lamellae (29% release). Since swimming is part of their repertoire of anti-predator behaviors this behavioral shift should be detrimental. 2) Caudal lamellae function in O₂ uptake. Trials were conducted with larvae having and not having lamellae in an experimental horizontal oxygen gradient system. Relative to larvae without lamellae, those with lamellae preferred deeper depths at PO₂ values greater than 70 torr. Many lamellae-less larvae distributed themselves at the water surface throughout the range of PO₂ values tested. Differential depth distribution between larvae with and without lamellae is highly significant (P < 0.01).     

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.