The evolutionary history of colour polymorphism in Ischnura damselflies.

A major challenge in evolutionary biology consists of understanding how genetic and phenotypic variation is created and maintained. In this study, we investigated the origin(s) and evolutionary patterns of the female-limited colour polymorphism in ischnuran damselflies. These consist of the presence of one to three colour morphs: one androchrome morph with a coloration that is similar to the male and two gynochrome morphs (infuscans and aurantiaca) with female-specific coloration. We (i) documented the colour and mating system of 44 of the 75 taxa within the genus Ischnura, (ii) reconstructed the evolutionary history of colour and mating system to identify the ancestral state, (iii) evaluated the stability of the colour morph status over time and (iv) tested for a correlation between colour and mating system. We found that the ancestral female colour of Ischnura was monomorphic and aurantiaca and that colour morph status changed over time, characterized by many gains and losses across the species tree. Our results further showed that colour polymorphism is significantly more frequent among polyandric species, whereas monandric species tend to be monomorphic. Research on some Ischnura species has shown that colour morphs have evolved to reduce male mating harassment, and our finding that the same phenotypic morphs have evolved multiple times (convergent evolution) suggests that several species in this genus might be experiencing similar selective pressures.

A Review of the Density, Biomass, and Secondary Production of Odonates.

Freshwater insects are highly significant as ecosystem service providers, contributing to provisioning services, supporting services, and cultural services. Odonates are dominant predators in many freshwater systems, becoming top predators in fishless ecosystems. One service that odonates provide is the export of matter and energy from aquatic to terrestrial ecosystems. In this study, we provide a review of the literature aiming to estimate the density, biomass, and secondary production of odonates and discuss to what extent this order of insects is relevant for the fertilization of terrestrial ecosystems. We found published data on 109 species belonging to 17 families of odonates from 44 papers. Odonata larvae are abundant in freshwater systems, with a mean density of 240.04 ± 48.01 individuals m-2 (±SE). Lentic habitats show much higher densities (104.40 ± 55.31 individuals m-2, N = 118) than lotic systems (27.12 ± 5.09, N = 70). The biomass estimations for odonates indicate values of 488.56 ± 134.51 mg m-2 y-1, with similar values in lentic and lotic habitats, which correspond to annual secondary productions of 3558.02 ± 2146.80 mg m-2 y-1. The highest biomass is found in dragonflies of the Aeshnidae, Corduliidae, and Gomphidae families. The available evidence suggests a significant potential contribution of Odonata to the exportation of material from water bodies to land. This is further strengthened by the ability of adult odonates to migrate and to colonize different types of water bodies.

A red flag: an experimental analysis of the function of pink colouration in males of Calopteryx haemorrhoidalis asturica (Odonata: Calopterygidae).

Males display phenotypic characteristics that may be associated with their quality, allowing non-random mating and post-copulatory female choice. In the damselfly Calopteryx haemorrhoidalis asturica, males have a conspicuous pink colouration in the underside of abdominal segments 8-10, which they exhibit during pre- and post-copulatory courtship. We hypothesized that this colouration functions to increase male mating success and/or to elicit females to oviposit. We estimated mating and oviposition success of 27 males, and on the following day, treated males had their segments 8-10 painted black and control males the seventh segment. We recorded the number of male-male fights and courtships, whether the courtship ended in copulation, and whether the female remained in the territory and laid eggs. Our results indicate that the mating success of male C. h. asturica was not significantly affected by the removal of the pink colouration of the abdominal tip, but this colouration clearly affected their success in enticing females to oviposit. Courtship frequency, fat content and muscle mass were positively correlated to male mating rate, and the number of aggressive encounters was negatively correlated. Our study yields experimental evidence for the function of pink colouration of male C. h. asturica, in the context of post-copulatory sexual selection.

Sex wars: a female genital spine forces male damselflies to shorten copulation duration.

In some species, males use weapons to harm females, increasing their short-term fitness. Here we show that females can use genital adaptations against males. Females of the damselfly Enallagma cyathigerum have a conspicuous vulvar spine on the eighth abdominal segment, which contacts with the male during copulation. We tested 3 hypotheses for its function: it (a) inflicts damage to the male during copulation; (b) facilitates endophytic oviposition; and (c) stimulates males during copulation to increase their investment. We found that males mated on average for 54 min with control females, but increased copulation to 99 min with females without spine. There was no evidence of physical harm of the spine on the male's seminal vesicle, which shows 8- to 18-folds, exactly where the spine contacts during copulation. Females with and without spine exhibited the same egg-laying rates and showed similar fecundity and fertility. Longevity was also similar in males mated to control and spineless females. In contrast to many species where females resist male harassment by behavioral responses, the morphological adaptation observed in E. cyathigerum appears to act as a sexual weapon, allowing females to control copulation duration. We suggest that the spine has evolved because of sexual conflict over mating duration.

Nesobasis rito sp. nov. (Zygoptera: Coenagrionidae), a new species of forest damselfly from Vanua Levu, Fiji.

Nesobasis rito sp. nov. (Holotype , Fiji, Vanua Levu, Drawa, 31 v 2018, A. Rivas-Torres leg.) from the comosa group is here described, illustrated, diagnosed, and compared with morphologically close species of the genus. Nesobasis rito can be distinguished from its related congeners by the shape of the caudal appendages and the ligula. The most similar species are N. comosa and N. heteroneura, which, like N. rito, have the caudal appendages covered by dense setae (especially the first species), but the shape differs clearly in lateral view, with N. rito having longer and more slender appendages, and a basal tooth clearly seen in dorsal view, absent in other members of the comosa group. The specific status of the collected specimens is also supported by the results of genetic analyses, where N. rito appears as a well-supported monophyletic clade. Nesobasis rito also has a distinct distribution from its most similar congeners: it is found on Vanua Levu, while N. comosa is found on Viti Levu and the closely related N. heteroneura is found on Viti Levu and Ovalau. All species of this group are found in streams with native forest riparian vegetation on their respective islands.

Alternative reproductive strategies in black-winged territorial males of Paraphlebia zoe (Odonata, Thaumatoneuridae).

Alternative reproductive strategies are commonly associated with male dimorphism. In Paraphlebia zoe, a species of damselfly whose males are dimorphic in wing coloration, black-and-white-winged (BW) males defend territories, while hyaline-winged (HW) males usually play the role of satellites. We found that several BW males can sometimes share a territory, and we hypothesized that within this morph there are two alternative tactics: submissive and dominant. We conducted an experiment to test whether dominant and submissive roles are plastic or stable and fixed on each individual. To this end, we manipulated black and white spots of BW males in four treatments: (i) painting over white and black spots without changing their size, (ii) erasing the white spot using black painting, (iii) increasing the black spot and moving the white spot maintaining its size and (iv) control males. Additionally, we investigated the correlation between some phenotypic variables (wing asymmetry, survival and recapture probabilities) and male behaviour (in terms of quality of the territory). We found that the two behavioural roles (submissive and dominant) were not affected by the manipulative experiments, therefore suggesting that they are stable and fixed. Additionally, we found a positive correlation between body size and survival in both sexes, and a positive effect of territory quality and lifespan on mating success. Moreover, the largest and youngest BW males were the most symmetrical. We conclude that Paraphlebia zoe holds high behavioural diversity, with two types of strategies in BW males, dominant and submissive. The occurrence of this intra-morph behavioural diversity might depend on demographic factors such as population density and/or the relative frequency of the different morphs.

Alternative reproductive strategies and the maintenance of female color polymorphism in damselflies.

Genetic polymorphisms are powerful model systems to study the maintenance of diversity in nature. In some systems, polymorphisms are limited to female coloration; these are thought to have arisen as a consequence of reducing male mating harassment, commonly resulting in negative frequency-dependent selection on female color morphs. One example is the damselfly Ischnura elegans, which shows three female color morphs and strong sexual conflict over mating rates. Here, we present research integrating male tactics, and female evolutionary strategies (female mating behavior and morph-specific female fecundity) in populations with different morph-specific mating frequencies, to obtain an understanding of mating rates in nature that goes beyond the mere measure of color frequencies. We found that female morph behavior differed significantly among but not within morphs (i.e., female morph behavior was fixed). In contrast, male tactics were strongly affected by the female morph frequency in the population. Laboratory work comparing morph-specific female fecundity revealed that androchrome females have lower fecundity than both of the gynochrome female morphs in the short term (3-days), but over a 10-day period one of the gynochrome female morphs became more fecund than either of the other morphs. In summary, our study found sex-specific dynamics in response to different morph frequencies and also highlights the importance of studying morph-specific fecundities across different time frames to gain a better understanding of the role of alternative reproductive strategies in the maintenance of female-limited color polymorphism.