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Petaluridae (Odonata: Anisoptera) is a relict dragonfly family, having diverged from its sister family in the Jurassic, of eleven species that are notable among odonates (dragonflies and damselflies) for their exclusive use of fen and bog habitats, their burrowing behavior as nymphs, large body size as adults, and extended lifespans. To date, several nodes within this family remain unresolved, limiting the study of the evolution of this peculiar family. Using an anchored hybrid enrichment dataset of over 900 loci we reconstructed the species tree of Petaluridae. To estimate the temporal origin of the genera within this family, we used a set of well-vetted fossils and a relaxed molecular clock model in a divergence time estimation analysis. We estimate that Petaluridae originated in the early Cretaceous and confirm the existence of monophyletic Gondwanan and Laurasian clades within the family. Our relaxed molecular clock analysis estimated that these clades diverged from their MRCA approximately 160 mya. Extant lineages within this family were identified to have persisted from 6 (Uropetala) to 120 million years (Phenes). Our biogeographical analyses focusing on a set of key regions suggest that divergence within Petaluridae is largely correlated with continental drift, the exposure of land bridges, and the development of mountain ranges. Our results support the hypothesis that species within Petaluridae have persisted for tens of millions of years, with little fossil evidence to suggest widespread extinction in the family, despite optimal conditions for the fossilization of nymphs. Petaluridae appear to be a rare example of habitat specialists that have persisted for tens of millions of years., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Inc. All rights reserved.)

Nikoulabasis roseosticta sp. nov. (holotype , Fiji, Vanua Levu Island, Navuturerega creek, 6 August 2009, Christopher Beatty leg.) is erected as a new taxon. Illustrations of key characters and a distribution map are provided.

Using recently published chromosome-length genome assemblies of two damselfly species, Ischnura elegans and Platycnemis pennipes, and two dragonfly species, Pantala flavescens and Tanypteryx hageni, we demonstrate that the autosomes of Odonata have undergone few fission, fusion, or inversion events, despite 250 million years of separation. In the four genomes discussed here, our results show that all autosomes have a clear ortholog in the ancestral karyotype. Despite this clear chromosomal orthology, we demonstrate that different factors, including concentration of repeat dynamics, GC content, relative position on the chromosome, and the relative proportion of coding sequence all influence the density of syntenic blocks across chromosomes. However, these factors do not interact to influence synteny the same way in any two pairs of species, nor is any one factor retained in all four species. Furthermore, it was previously unknown whether the micro-chromosomes in Odonata are descended from one ancestral chromosome. Despite structural rearrangements, our evidence suggests that the micro-chromosomes in the sampled Odonata do indeed descend from an ancestral chromosome, and that the micro-chromosome in P. flavescens was lost through fusion with autosomes., (© 2023 The Authors. Molecular Ecology published by John Wiley & Sons Ltd.)

We present a chromosome-length genome assembly and annotation of the Black Petaltail dragonfly (Tanypteryx hageni). This habitat specialist diverged from its sister species over 70 million years ago, and separated from the most closely related Odonata with a reference genome 150 million years ago. Using PacBio HiFi reads and Hi-C data for scaffolding we produce one of the most high-quality Odonata genomes to date. A scaffold N50 of 206.6 Mb and a single copy BUSCO score of 96.2% indicate high contiguity and completeness., (© The Author(s) 2023. Published by Oxford University Press on behalf of Society for Molecular Biology and Evolution.)

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.

Background: The New World Tropics has experienced a dynamic landscape across evolutionary history and harbors a high diversity of flora and fauna. While there are some studies addressing diversification in Neotropical vertebrates and plants, there is still a lack of knowledge in arthropods. Here we examine temporal and spatial diversification patterns in the damselfly family Polythoridae, which comprises seven genera with a total of 58 species distributed across much of Central and South America., Results: Our time-calibrated phylogeny for 48 species suggests that this family radiated during the late Eocene (~ 33 Ma), diversifying during the Miocene. As with other neotropical groups, the Most Recent Common Ancestor (MRCA) of most of the Polythoridae genera has a primary origin in the Northern Andes though the MRCA of at least one genus may have appeared in the Amazon Basin. Our molecular clock suggests correlations with some major geographical events, and our biogeographical modeling (with BioGeoBEARS and RASP) found a significant influence of the formation of the Pebas and Acre systems on the early diversification of these damselflies, though evidence for the influence of the rise of the different Andean ranges was mixed. Diversification rates have been uniform in all genera except one-Polythore-where a significant increase in the late Pliocene (~ 3 mya) may have been influenced by recent Andean uplift., Conclusion: The biogeographical models implemented here suggest that the Pebas and Acre Systems were significant geological events associated with the diversification of this damselfly family; while diversification in the tree shows some correlation with mountain building events, it is possible that other abiotic and biotic changes during our study period have influenced diversification as well. The high diversification rate observed in Polythore could be explained by the late uplift of the Northern Andes. However, it is possible that other intrinsic factors like sexual and natural selection acting on color patterns could be involved in the diversification of this genus.

Wolbachia is one of the most common endosymbionts found infecting arthropods. Theory predicts symbionts like Wolbachia will be more common in species radiations, as host shift events occur with greatest frequency between closely related species. Further, the presence of Wolbachia itself may engender reproductive isolation, and promote speciation of their hosts. Here we screened 178 individuals belonging to 30 species of the damselfly genera Nesobasis and Melanesobasis - species radiations endemic to the Fiji archipelago in the South Pacific - for Wolbachia, using multilocus sequence typing to characterize bacterial strains. Incidence of Wolbachia was 71% in Nesobasis and 40% in Melanesobasis, and prevalence was also high, with an average of 88% in the Nesobasis species screened. We identified a total of 25 Wolbachia strains, belonging to supergroups A, B and F, with some epidemic strains present in multiple species. The occurrence of Wolbachia in both males and females, and the similar global prevalence found in both sexes rules out any strong effect of Wolbachia on the primary sex-ratio, but are compatible with the phenotype of cytoplasmic incompatibility. Nesobasis has higher species richness than most endemic island damselfly genera, and we discuss the potential for endosymbiont-mediated speciation within this group.

The study of island fauna has greatly informed our understanding of the evolution of diversity. We here examine the phylogenetics, biogeography, and diversification of the damselfly genera Nesobasis and Melanesobasis , endemic to the Fiji Islands, to explore mechanisms of speciation in these highly speciose groups. Using mitochondrial (COI, 12S) and nuclear (ITS) replicons, we recovered garli-part maximum likelihood and mrbayes Bayesian phylogenetic hypotheses for 26 species of Nesobasis and eight species/subspecies of Melanesobasis . Biogeographical patterns were explored using lagrange and bayes-lagrange and interpreted through beast relaxed clock dating analyses. We found that Nesobasis and Melanesobasis have radiated throughout Fiji, but are not sister groups. For Nesobasis , while the two largest islands of the archipelago-Viti Levu and Vanua Levu-currently host two distinct species assemblages, they do not represent phylogenetic clades; of the three major groupings each contains some Viti Levu and some Vanua Levu species, suggesting independent colonization events across the archipelago. Our beast analysis suggests a high level of species diversification around 2-6 Ma. Our ancestral area reconstruction (rasp-lagrange) suggests that both dispersal and vicariance events contributed to the evolution of diversity. We thus conclude that the evolutionary history of Nesobasis and Melanesobasis is complex; while inter-island dispersal followed by speciation (i.e., peripatry) has contributed to diversity, speciation within islands appears to have taken place a number of times as well. This speciation has taken place relatively recently and appears to be driven more by reproductive isolation than by ecological differentiation: while species in Nesobasis are morphologically distinct from one another, they are ecologically very similar, and currently are found to exist sympatrically throughout the islands on which they are distributed. We consider the potential for allopatric speciation within islands, as well as the influence of parasitic endosymbionts, to explain the high rates of speciation in these 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.

In damselflies, sexual colour dimorphism is commonly explained as a consequence of selection on traits that increase male attractiveness to females. However, while many species in the damselfly family Coenagrionidae (Insecta: Odonata) are sexually dimorphic, the males do not engage in displays, and male competition for mates resembles a "scramble". An alternative explanation for the sexual differences in coloration within these species is that sexual dimorphism has evolved as a sex-related warning signal, with males signalling their uprofitability as mates to other males, thereby avoiding harassment from conspecifics. We evaluated an underlying assumption of the theory that male-male harassment rate is influenced by colour by comparing harassment of males of the species Nehalennia irene that had been painted to make them appear: (i) similar to an unaltered male (blue), (ii) different from a male (orange) and (iii) more similar to a female (black). When caged together we found that blue-painted males experienced significantly lower harassment than black-painted males. When unpainted males were caged with each type of painted male we found that blue-painted males and the unpainted males housed in the same cages experienced lower rates of harassment than males housed in cages where some males were painted black, suggesting that a single, reliable signal of unprofitability may benefit the individuals that carry it. While our results do not in themselves demonstrate that sexual colour dimorphism originally evolved as an intra-specific warning signal, they do show that harassment is influenced by coloration, and that such selection could conceivably maintain male coloration as a warning signal.

The study of color polymorphisms (CP) has provided profound insights into the maintenance of genetic variation in natural populations. We here offer the first evidence for an elaborate wing polymorphism in the Neotropical damselfly genus Polythore, which consists of 21 described species, distributed along the eastern slopes of the Andes in South America. These damselflies display highly complex wing colors and patterning, incorporating black, white, yellow, and orange in multiple wing bands. Wing colors, along with some components of the male genitalia, have been the primary characters used in species description; few other morphological traits vary within the group, and so there are few useful diagnostic characters. Previous research has indicated the possibility of a cryptic species existing in P. procera in Colombia, despite there being no significant differences in wing color and pattern between the populations of the two putative species. Here we analyze the complexity and diversity of wing color patterns of individuals from five described Polythore species in the Central Amazon Basin of Peru using a novel suite of morphological analyses to quantify wing color and pattern: geometric morphometrics, chromaticity analysis, and Gabor wavelet transformation. We then test whether these color patterns are good predictors of species by recovering the phylogenetic relationships among the 5 species using the barcode gene (COI). Our results suggest that, while highly distinct and discrete wing patterns exist in Polythore, these "wingforms" do not represent monophyletic clades in the recovered topology. The wingforms identified as P. victoria and P. ornata are both involved in a polymorphism with P. neopicta; also, cryptic speciation may have taking place among individuals with the P. victoria wingform. Only P. aurora and P. spateri represent monophyletic species with a single wingform in our molecular phylogeny. We discuss the implications of this polymorphism, and the potential evolutionary mechanisms that could maintain it.

We report here the prevalence of parasitism by water mites (Arrenurus sp.) and terrestrial mites (Leptus killingtoni) on parthenogenetic Ischnura hastata (Odonata: Coenagrionidae) from the Azores islands. Leptus killingtoni was only found on the island of Pico, and the prevalence of infestation was highly variable among the different ponds studied, ranging from 0 to 41%. Leptus killingtoni was observed on three of the four odonate species from the archipelago: I. hastata, I. pumilio, and Sympetrum fonscolombii, all of them new hosts for this species. Aquatic mites have been found parasitizing I. hastata females on the island of São Miguel. The prevalence of mite parasitism by Arrenurus sp. on I. hastata was very low, ranging from 12% (2003) to 1% (2008), and in most of the studied ponds, no mites were found attached to females. Although I. hastata coexists with a sexual congener species in the Azores (I. pumilio), they are syntopic in only a small fraction of ponds. Therefore, a comparison between I. hastata and I. pumilio was insufficient to test the predictions of the Red Queen Hypothesis, and further research on parasitism rates in both species needs to be done. In any case, the low prevalence of mite parasitism found in the Azores, coupled with the fact that most of the populations in the archipelago are almost free from competitors and predators, could explain the persistence of these I. hastata parthenogenetic populations, despite their low levels of genetic variation.

Although the principles of disruptive colouration are widely believed to explain a variety of animal colour patterns, there has been no field evidence that it works to reduce the detection rates of natural prey. In a recent paper, Cuthill et al. successfully address this shortfall, separating the benefits of background matching from those of disruptive colouration. Their results provide the first definitive field support for this long-recognized phenomenon and suggest several new avenues of research.

Prey species that are unprofitable to attack often share conspicuous colours and patterns with other coexisting defended species. This phenomenon, termed müllerian mimicry, has long been explained as a consequence of selection on defended prey to adopt a common way of advertising their unprofitability. However, studies using two unpalatable prey types have not always supported this theory. Here we show, using a system of humans hunting for computer-generated prey, that predators do not always generate strong selection for mimicry when there are two unprofitable prey types. By contrast, we demonstrate that when predators are faced with a range of different prey species, selection on unprofitable prey to resemble one another can be intense. Here the primary selective force is not one in which predators evaluate the profitabilities of distinct prey types independently, but one in which predators learn better to avoid unprofitable phenotypes that share traits distinguishing them from profitable prey. This need to simplify decision making readily facilitates the spread of imperfect mimetic forms from rarity, and suggests that müllerian mimicry is more likely to arise in multispecies communities.

Prey that are unprofitable to attack (for example, those containing noxious chemicals) frequently exhibit slower and more predictable movement than species that lack these defenses. Possible explanations for the phenomenon include a lack of selection pressure on unprofitable prey to avoid predators and active selection on unprofitable prey to advertise their noxiousness. We explicitly tested these and other hypotheses using a novel "artificial world" in which the locomotory characteristics (step size, waiting time, and angular direction) of artificial profitable and unprofitable computer-generated prey were subject to continued selection by humans over a number of generations. Unprofitable prey evolved significantly slower movement behavior than profitable prey when they were readily recognized as unprofitable, and also when they frequently survived predatory attacks. This difference arose primarily as a consequence of more intense selection on profitable prey to avoid capture. When unprofitable prey were very similar (but not identical) in morphological appearance to profitable prey, unprofitable prey evolved particularly slow movement behavior, presumably because when they were slow-moving they could be more readily recognized as being unprofitable. When unprofitable prey were constrained to move slowly, a morphologically identical profitable prey species evolved locomotor mimicry only when it had no more effective means of avoiding predation. Overall, our results provide some of the first empirical support for a number of earlier hypotheses for differences in movement between unprofitable and profitable prey and demonstrate that locomotor mimicry is not an inevitable outcome of selection even in morphologically similar prey.

It is widely argued that defended prey have tended to evolve conspicuous traits because predators more readily learn to avoid defended prey when they are conspicuous. However, a rival theory proposes that defended prey have evolved such characters because it allows them to be distinguished from undefended prey. Here we investigated how the attributes of defended (unprofitable) and undefended (profitable) computer-generated prey species tended to evolve when they were subject to selection by foraging humans. When cryptic forms of defended and undefended species were similar in appearance but their conspicuous forms were not, defended prey became conspicuous while undefended prey remained cryptic. Indeed, in all of our experiments, defended prey invariably evolved any trait that enabled them to be distinguished from undefended prey, even if such traits were cryptic. When conspicuous mutants of defended prey were extremely rare, they frequently overcame their initial disadvantage by chance. When Batesian mimicry of defended species was possible, defended prey evolved unique traits or characteristics that would make undefended prey vulnerable. Overall, our work supports the contention that warning signals are selected for their reliability as indicators of defense rather than to capitalize on any inherent educational biases of predators.