Seasonal polyphenism in wing coloration affects species recognition in rubyspot damselflies (Hetaerina spp.)

Understanding how phenotypic plasticity evolves and in turn affects the course of evolution is a major challenge in modern biology. By definition, biological species are reproductively isolated, but many animals fail to distinguish between conspecifics and closely related heterospecifics. In some cases, phenotypic plasticity may interfere with species recognition. Here, we document a seasonal polyphenism in the degree of dark wing pigmentation in smoky rubyspot damselflies (Hetaerina titia) – a shift so pronounced that it led early researchers to classify different forms of H. titia as separate species. We further show how the seasonal colour shift impacts species recognition with the sympatric congener Hetaerina occisa. Interspecific aggression (territorial fights) and reproductive interference (mating attempts) are much more frequent early in the year, when H. titia more closely resembles H. occisa, compared to later in the year when the dark phase of H. titia predominates. Using wing colour manipulations of tethered damselflies, we show that the seasonal changes in interspecific interactions are caused not only by the seasonal colour shift but also by shifts in discriminatory behaviour in both species. We also experimentally tested and rejected the hypothesis that learning underlies the behavioural shifts in H. occisa. An alternative hypothesis, which remains to be tested, is that the seasonal polyphenism in H. titia wing coloration has resulted in the evolution of a corresponding seasonal polyphenism in species recognition in H. occisa. This study illustrates one of the many possible ways that plasticity in species recognition cues may influence the evolution of interspecific interactions.     

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.     

Field estimates of reproductive success in a model insect: behavioural surrogates are poor predictors of fitness

Understanding, and therefore measuring, factors that determine fitness is a central problem in evolutionary biology. We studied a natural population of Coenagrion puella (Odonata: Zygoptera) over two entire breeding seasons, with over a thousand individuals uniquely marked and genotyped, and all mating events at the rendezvous site recorded. Using a parentage analysis, fitness of individuals in the first generation was quantified as the numbers of offspring that survived to maturity. Although mating behaviour can be predicted by environmental and demographical variables, the numbers of mature offspring produced (fitness) cannot, and crucially, are poorly correlated with behavioural observations of mating. While fitness of both sexes was positively related to mating behaviour and to female's ectoparasite burden, these behavioural observations explained little more variance in offspring production than environmental and demographical variables. Thus, we demonstrate that behavioural measures of reproductive success are not necessarily reliable estimates of fitness in natural populations.     

Resource limitation,predation risk and compensatory growth in a damselfly

Periods of poor nutrition during early development may have negative fitness consequences in subsequent periods of ontogeny. In insects, suppression of growth and developmental rate during the larval stage are likely to affect size and timing of maturity, which in turn may lead to reduced reproductive success or survivorship. In light of these costs, individuals may achieve compensatory growth via behavioural or physiological mechanisms following food limitation. In this study, we examined the effects of a temporary period of food restriction on subsequent growth and age and size at maturity in the larval damselfly Ischnura verticalis (Odonata: Coenagrionidae). We also asked whether this temporary period of reduced nutrition affected subsequent foraging behaviour under predation risk. I. verticalis larvae exposed to a temporary food shortage suffered from a reduced growth rate during this period relative to a control group that was fed ad libitum. However, increased growth rates later in development ensured that adult body size measurements (head and pronotum widths) did not differ between the treatments upon emergence. In contrast, adult dry mass did not catch up to that of the controls, indicating that the increased growth rates for size dimensions occur at the cost of similar gains in mass. Predators reduced foraging effort of larvae, but this reduction did not differ between control larvae and those previously exposed to poor nutrition.     

Significance of temporal changes when designing a reservoir for conservation of dragonfly diversity

While there has been much focus in biodiversity conservation that transcends place, few studies transcend time. Yet an appreciation of vegetational and hydrological succession is essential for maintaining a habitat that has been created with the aim of conserving a particular group of organisms. This is a study of changes in a dragonfly assemblage over a period of 13 years at a biodiversity-rich, southern hemisphere reservoir. A total of 30 dragonfly species were recorded in this study, compared to 12 species before the reservoir was constructed in 1988, and 26 species in 1993, with 25 species resident in both 1993 and 2001. Two of these are local endemics. One other endemic was lost to succession in 1993 but reappeared in 2001. Three other species never reappeared after succession in 1993, yet six other species appeared after this date. Multivariate analyses identified structural and compositional vegetation, especially marginal forest, percentage vegetation cover, percentage shade, as the most important environmental variables determining dragonfly species composition. Other important environmental variables were grasses of tall, medium and short height categories, submerged vegetation, water flow and amount of open water. Not surprisingly, successional changes in vegetation physiognomy and in water conditions significantly increased Odonata species richness and diversity over the years. More importantly, the study shows that to maintain both high species richness, including endemics, it is essential to maintain a variety of biotopes using selective management of the marginal vegetation without allowing succession to proceed to a point where overgrowth of the bank and silting of the bottom begin to impoverish the fauna.     

An integrative approach to species discovery in odonates: from character‐based DNA barcoding to ecology

Modern taxonomy requires an analytical approach incorporating all lines of evidence into decision‐making. Such an approach can enhance both species identification and species discovery. The character‐based DNA barcode method provides a molecular data set that can be incorporated into classical taxonomic data such that the discovery of new species can be made in an analytical framework that includes multiple sources of data. We here illustrate such a corroborative framework in a dragonfly model system that permits the discovery of two new, but visually cryptic species. In the African dragonfly genus Trithemis three distinct genetic clusters can be detected which could not be identified by using classical taxonomic characters. In order to test the hypothesis of two new species, DNA‐barcodes from different sequence markers (ND1 and COI) were combined with morphological, ecological and biogeographic data sets. Phylogenetic analyses and incorporation of all data sets into a scheme called taxonomic circle highly supports the hypothesis of two new species. Our case study suggests an analytical approach to modern taxonomy that integrates data sets from different disciplines, thereby increasing the ease and reliability of both species discovery and species assignment.     

First fossil Calopterygoidea (Odonata: Zygoptera) from Southeastern Asia: A new genus and species from the Paleogene of China

Sinocalopteryx shangyongensis nov. gen., nov. sp., the first fossil calopterygoid from eastern Asia, is described from the earliest Eocene of Southwest China. Although the new genus has the principle synapomorphies of Calopterygoidea, it possesses a unique structure (possible reversal) in the pattern of vein RP1/2.     

Flexible Mate Guarding Tactics in the Dragonfly Sympelrum Internum (Odonata: Libellulidae)

Mate guarding–a behaviour prevalent in odonates–is a post copulatory association during which males prevent females from re-mating. Some species use two forms of guarding: contact mate guarding, which is energetically costly but highly effective and non-contact mate guarding, which is less costly but less effective. This study aimed to determine if male Sympetrum internum (Odonata:Libellulidae) adjust the duration of contact mate guarding according to environmental, temporal and physiological factors. There was a significant interaction between male density and season on duration of contact mate guarding. Early in the season males increased the duration of contact guarding as the density of rivals increased. Later in the season males guarded mates longer irrespective of male density. Wind and temperature did not detectabiy alter the duration of contact mate guarding, suggesting that the trade-off between current and future reproductive success was more important than were physiological costs.