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.     

Phylogeographic analysis of a recent radiation of Enallagma damselflies (Odonata: Coenagrionidae)

A phylogenetic hypothesis revealed two recent radiations among species of Enallagma damselflies, and extensive ecological work suggests that both adaptive and nonadaptive processes are involved in these radiations. We analysed the geographical pattern of genetic variability at 868 bp of mitochondrial DNA (mtDNA) among 283 individuals of 5 species displaying little ecological differentiation to identify the ancestral lineage, support their independent evolutionary trajectories and identify historical events and the underlying mechanism for one of these radiations. Nested clade analysis results clearly support a past event of range fragmentation in E. hageni. These Atlantic and Continental hageni races experienced distinct dispersal histories and still maintain nearly nonoverlapping ranges All four other species derive from the Continental hageni. Whereas three species endemic to the Atlantic coastal plain show little genetic variation, E. ebrium shared several haplotypes with the Continental hageni. Contrasting levels of genetic differentiation between E. hageni and E. ebrium in geographical areas associated with distinct events of E. hageni's recent history support the recent origin of this species. Altogether, our results are compatible with a process of radiation via divergence in mate recognition systems within the Continental hageni race following secondary contacts between putative refugial races.     

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.     

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.     

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.     

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.     

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 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.     

Phylogenetic relationships of North American damselflies of the genus Ischnura (Odonata: Zygoptera: Coenagrionidae) based on sequences of three mitochondrial genes

Relationships of North American damselflies of the genus Ischnura (Odonata: Zygoptera: Coenagrionidae) were investigated using a total of 1205 bp from portions of three mitochondrial genes: cytochrome b, cytochrome oxidase II, and 12S ribosomal DNA. Parsimony and neighbor joining analyses reveal a monophyletic group consisting of I. damula, I. demorsa, I. perparva, I. posita posita, I. posita atezca, I. verticalis, and probably I. denticollis, likely reflecting a recent radiation in North America. Ischnura kellicotti, I. barberi, I. prognata, I. hastata, I. ramburii, and I. capreola appear to represent much earlier divergences in the group. Many previous hypotheses of relationships among North American species of Ischnura are not supported by the molecular-based analyses. However, there is agreement in many respects between the results of the molecular phylogenetic analyses and the morphologically based conclusions of Kennedy (1919, "The Phylogeny of the Zygoptera," Ph.D. Dissertation, Cornell University, Ithaca). Although results of single-gene phylogenetic analyses often differ, there are very few cases in which there is strong support for conflicting relationships using different partitions of the data. Combined analysis of all three genes yields trees with stronger support overall than the single-gene analyses, and the combined data trees that result from diverse data treatments are congruent with one another in most respects.     

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.

     

Phylogeny of Neotropical Protoneuridae (Odonata: Zygoptera) and a preliminary study of their relationship with related families

Abstract.  A cladistic analysis of Neotropical Protoneuridae was performed on a data matrix of 48 morphological characters and 43 terminal taxa. Representatives of Paleotropical Protoneuridae, Platycnemididae and Isostictidae were included to test their relationships with Neotropical Protoneuridae. Coenagrionidae was chosen as the outgroup, but alternative analyses with Platycnemididae as the outgroup were also performed. Protoneuridae appears as a polyphyletic clade, with its Paleotropical component being more closely related to Platycnemididae and Isostictidae. Neotropical Protoneuridae appear as a monophyletic clade; included genera considered monophyletic or valid monotypic taxa are Epipleoneura Williamson, 1915 ; Idioneura Selys, 1860; Junix Rácenis, 1968; Neoneura Selys, 1860; Peristicta Hagen in Selys, 1860; Roppaneura Santos, 1966; and Lamproneura De Marmels, 2003. A key to the Neotropical Protoneuridae genera is included.     

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.     

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.     

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.