Demography and Behaviour of Teinopodagrion oscillans (Odonata: Megapodagrionidae) in a Protected Area of the Colombian Andean Region.

The demography and behaviour of Teinopodagrion oscillans was studied in a protected area in the Andean region of Colombia. Adult damselflies were individually marked, and using their recapture histories, we estimated survival, longevity, sex ratio, and population size using Cormack-Jolly-Seber models. Other aspects of their behaviour were recorded. Survival, recapture, and lifespan (14.1 ± 0.59 days) were similar for both sexes and all age groups. Mature males were larger, and the distance from the water was similar for all individuals. The most supported model was the time-dependent model for survival and recapture. This suggests that weather variations affect the demography of this population in a significant way. Individuals exhibited high fidelity to their site perch, perching with open wings near water on a variety of perches. Mature males make short flights from the perch to intercept conspecific and interspecific males and to hunt prey. The tandem position was formed on macrophytes, and then the pair flew away. Oviposition lasted for 11.23 min on average, with the females ovipositing by abdomen submersion. Our results offer insights into the demographic characteristics and behaviour of this species, providing crucial information for the short- and long-term, from the demography of one species to the conservation of ecosystems of the Andean region.

Direct and cross-generational effects of reproduction on fitness and behavioral variability in male-biased environments.

Population structure determines individuals' interactions and trade-offs with evolutionary consequences. Male-biased populations increase intrasexual competition and intersexual harassment, reducing female resource acquisition, and thus, resources availability for the following generation. We analyzed direct and cross-generational effects of male harassment in two generations of damselflies (Odonata). We exposed adult females to treatments with different sex-ratio and density (balanced and male-biased) to modify the male harassment level. We analyzed female fecundity, fertility, and number of faecal deposits as an indirect measure of resources acquisition. We studied female flight performance after repeated exposures to males. We analyzed survivorship, development, exploration, thigmotaxis, and feeding latency of larvae produced by the experimental females. In both generations, we analyzed four metrics of behavior: mean value, interindividual differences in plasticity, intra-individual unpredictability, and repeatability. Mating duration increased in male-biased treatment, whereas female resources acquisition and fertility decreased. Females that mated longer showed higher fecundity when they were exposed to balanced treatment, but not if they were exposed to male-biased treatment. Females from the male-biased treatment showed interindividual differences in plasticity and no repeatability in flight performance. Offspring showed balanced sex-ratio and similar survivorship, development, and feeding latency independently of the parental treatment; however, females exposed to male-biased treatment produced offspring with higher differences in exploration plasticity and daughters less explorative and with higher unpredictable thigmotaxis. We propose prolonged copulation as courtship at balanced sex-ratio but a cost to females under male-biased sex-ratio. Cross-generational effects in behavioral variability may be a mechanism to cope with predicted future environments.

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.

Nikoulabasis roseosticta, sp. nov. from Fiji (Odonata: Coenagrionidae).

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.

Detecting Darwinian Shortfalls in the Amazonian Odonata.

Among the oldest winged insects, odonates are a monophyletic order that have become important models for ecological studies because of their highly diverse reproductive behaviors and their role as top predators and bioindicators. However, knowledge on evolutionary relationships within the order is still scarce compared to other taxa, and this situation is even more complicated in areas with high biodiversity, such as in the Amazon. Here, we sought to identify knowledge gaps on Amazonian Odonata regarding three main aspects: (i) how the inclusion of Amazonian taxa affects our interpretation of the evolutionary relationships of Zygoptera and Anisoptera; (ii) the position of Amazonian taxa in the existing supertree of the Odonata; (iii) dating evolutionary divergence between nodes using fossil records; (iv) assessing whether more species-rich basins (e.g., Amazon basin) have a larger phylogenetic gap when compared to basins with lower richness in South and Central America; and (v) in the light of our knowledge, we discuss diversification patterns found in the most predominant clades of Amazonian taxa. We built a supertree from currently available phylogenetic information of Odonata. The results show that there is no genetic information for 85% (n: 503) of the Amazonian species and that family level relationships are unknown for 17 genera. After compiling the data, we observed that clades belonging to Neotropical lineages are the most poorly resolved, with large polytomies. This problem was identified in many Anisoptera genera, such as Macrothemis, Dasythemis, Elasmothemis, and Erythrodiplax. Our results also suggest that not always the richest basins have the greatest phylogenetic gaps. As expected, we found important gaps in the existing Odonata phylogenies, especially in clades that include Amazonian representatives, that are also those less known from ecological and conservation perspectives.

Ischnura praematura sp. nov. (Odonata: Zygoptera: Coenagrionidae): a species from Yunnan (China) whose females mate in the teneral state.

Ischnura praematura sp. nov. (Holotype , China, Yunnan, Lijiang, 263103.54N, 1001338.89E, 2396 m, 04 xii 2015, I. Sanmartn-Villar H. Zhang leg.) is morphologically described, illustrated and compared with close species of the genus. Ischnura praematura can be mainly distinguished from its congeners I. aurora, I. rubilio and I. asiatica by its abdominal and thoracic morphology and colouration. The posterior lobe of the prothorax is elevated in I. praematura and the mesostigmal plates possess dorsal triangular protuberances. Ischnura praematura shows pointed paraprocts, internalised wide cerci and lacks a dorsal tuberculum in the tenth abdominal segment. The blue abdominal colouration is present in the last three segments (incomplete for segment eight and ten in some individuals). No female polychromatism was detected and all females observed possessed different colouration than male (gynochrome). Morphological distinctiveness of the species is supported by genetic analyses, which show that I. praematura forms a well-supported, monophyletic clade, with I. asiatica, I. ezoin and I. pumilio as the most closely related species. In the field, mature females show strong reluctance to mate, and males were observed mating with newly emerged females.

Blood parasite diversity (Apicomplexa: Haemogregarinidae) within the western populations of the European pond turtle Emys orbicularis.

Molecular tools have revolutionized assessments of blood parasites in freshwater turtles. In the Iberian Peninsula and North Africa, two native species of terrapins occur, Emys orbicularis (Linnaeus) and Mauremys leprosa (Schweigger). Both have been identified as hosts for the blood parasite Haemogregarina stepanowi Danilewsky, 1885, which has also been found in related species. However, recent assessments of M. leprosa have identified several distinct genetic lineages of these parasites in this host, while only three haemogregarine lineages were identified in E. orbicularis in Tunisia. Here, we screened 215 individuals of E. orbicularis from the Iberian Peninsula, Menorca Island and Morocco for haemogregarine parasites using partial 18S rRNA gene sequences to estimate relationships. Three unrelated lineages of parasites were detected, one presumed H. stepanowi and two lineages previously known from M. leprosa. A considerable undescribed diversity of parasites exists within these vertebrate host species, while mixed infection and host-sharing is also widespread. Considering that E. orbicularis is near threatened in this region, it is of great importance to identify the parasites infecting it, and to further assess the potential deleterious impact of these diverse parasites on their hosts.

Biosociological ethodiversity in the social system.

A comprehensive understanding of human sociality needs to embrace the coevolution of genes and culture. Recent advances in biological research about niche construction by organisms, and the development of the concepts of social niche and ethodiversity, can be integrated into a common approach to understand this coevolution, which implies the interaction between sociology and ecology in an integrative framework of knowledge. In this paper the authors propose such inclusive biosociological and heuristic framework to improve the understanding of the evolution of social niche construction. In addition, it allows a better understanding of the concept of sociotype in non-human organisms and explains some aspects of the social or presocial behavior through the concept of ethodiversity.

Impact of environmental changes on the behavioral diversity of the Odonata (Insecta) in the Amazon.

The odonates are insects that have a wide range of reproductive, ritualized territorial, and aggressive behaviors. Changes in behavior are the first response of most odonate species to environmental alterations. In this context, the primary objective of the present study was to assess the effects of environmental alterations resulting from shifts in land use on different aspects of the behavioral diversity of adult odonates. Fieldwork was conducted at 92 low-order streams in two different regions of the Brazilian Amazon. To address our main objective, we measured 29 abiotic variables at each stream, together with five morphological and five behavioral traits of the resident odonates. The results indicate a loss of behaviors at sites impacted by anthropogenic changes, as well as variation in some morphological/behavioral traits under specific environmental conditions. We highlight the importance of considering behavioral traits in the development of conservation strategies, given that species with a unique behavioral repertoire may suffer specific types of extinction pressure.

Phylogeny and classification of Odonata using targeted genomics.

Dragonflies and damselflies are a charismatic, medium-sized insect order (~6300 species) with a unique potential to approach comparative research questions. Their taxonomy and many ecological traits for a large fraction of extant species are relatively well understood. However, until now, the lack of a large-scale phylogeny based on high throughput data with the potential to connect both perspectives has precluded comparative evolutionary questions for these insects. Here, we provide an ordinal hypothesis of classification based on anchored hybrid enrichment using a total of 136 species representing 46 of the 48 families or incertae sedis, and a total of 478 target loci. Our analyses recovered the monophyly for all three suborders: Anisoptera, Anisozygoptera and Zygoptera. Although the backbone of the topology was reinforced and showed the highest support values to date, our genomic data was unable to stronglyresolve portions of the topology. In addition, a quartet sampling approach highlights the potential evolutionary scenarios that may have shaped evolutionary phylogeny (e.g., incomplete lineage sorting and introgression) of this taxon. Finally, in light of our phylogenomic reconstruction and previous morphological and molecular information we proposed an updated odonate classification and define five new families (Amanipodagrionidae fam. nov., Mesagrionidae fam. nov., Mesopodagrionidae fam. nov., Priscagrionidae fam. nov., Protolestidae fam. nov.) and reinstate another two (Rhipidolestidae stat. res., Tatocnemididae stat. res.). Additionally, we feature the problematic taxonomic groupings for examination in future studies to improve our current phylogenetic hypothesis.

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.

DISPERSE, a trait database to assess the dispersal potential of European aquatic macroinvertebrates.

Dispersal is an essential process in population and community dynamics, but is difficult to measure in the field. In freshwater ecosystems, information on biological traits related to organisms' morphology, life history and behaviour provides useful dispersal proxies, but information remains scattered or unpublished for many taxa. We compiled information on multiple dispersal-related biological traits of European aquatic macroinvertebrates in a unique resource, the DISPERSE database. DISPERSE includes nine dispersal-related traits subdivided into 39 trait categories for 480 taxa, including Annelida, Mollusca, Platyhelminthes, and Arthropoda such as Crustacea and Insecta, generally at the genus level. Information within DISPERSE can be used to address fundamental research questions in metapopulation ecology, metacommunity ecology, macroecology and evolutionary ecology. Information on dispersal proxies can be applied to improve predictions of ecological responses to global change, and to inform improvements to biomonitoring, conservation and management strategies. The diverse sources used in DISPERSE complement existing trait databases by providing new information on dispersal traits, most of which would not otherwise be accessible to the scientific community.

Revision of the genus Minagrion Santos, 1965 (Odonata: Coenagrionidae).

Endemic to Brazil, the genus Minagrion Santos, 1965 is revised for the first time. The genus contains five species, and here we establish diagnostic characters, keys and drawings for both sexes. Furthermore, we propose the synonymy of M. franciscoi Machado Bedê, 2015 new syn. with M. caldense Santos, 1965.

Darwin Returns to the Galapagos: Genetic and Morphological Analyses Confirm the Presence of Tramea darwini at the Archipelago (Odonata, Libellulidae).

The status of the Tramea species present in the Galapagos Islands (Odonata, Libellulidae) has been the subject of a long-standing debate among odonatologists. Here, we use molecular and morphological data to analyze a series of specimens from this genus collected in 2018 from the Islands of San Cristobal, Isabela, and Santa Cruz, with the aim of determining their relationship with Tramea calverti Muttkowski and with their currently considered senior synonym T. cophysa Hagen. We combined sequencing of mitochondrial and nuclear DNA with morphological examination of several specimens of Tramea, including representatives of continental T. cophysa and T. calverti. Our molecular analyses place the Tramea from Galapagos in the same clade as T. calverti, with T. cophysa as a closely related species. The morphological analyses found only one consistent difference between T. cophysa and T. calverti: the presence of an accessory lobe in the male vesica spermalis of T. cophysa that is absent in T. calverti and in the Tramea from Galapagos. In agreement with our genetic results, the overall morphological differences documented by us indicate that the Galapagos material examined is conspecific with T. calverti. In light of this, and following the principle of priority in taxonomic nomenclature, Tramea calverti Muttkowski, 1910 should hereafter be considered a junior synonym of Tramea darwini Kirby, 1889.

The female of Heteragrion icterops Selys, 1862 (Odonata: Heteragrionidae) with comments and key on the Brazilian females of Heteragrion group B.

The female of Heteragrion icterops Selys, 1862, incompletely mentioned in 1886, is described, illustrated, keyed and diagnosed based on specimens collected in Pará State, North Brazil. We also provide an identification key and diagnostic comments on other two Group B females occurring in Brazil: H. angustipenne Selys, 1886 and H. bariai De Marmels, 1989. Heteragrion icterops females can be distinguished from other female congeners by having the genital valves tips leveling the posterior margin of S10 and by the morphology of the intersternites, presenting a well-developed dorsal plate and lacking a posterior plate.

Philogenia gaiae sp. nov. (Zygoptera: Philogeniidae) and description of the female of P. macuma Dunkle, 1986, two species from the Ecuadorean lowland rainforest.

Philogenia gaiae sp. nov. (Holotype ♂, Ecuador, Orellana, Tiputini Biodiversity Station, -0.6349, -76.1501, 241 m, 13 xii 2012, A. Cordero-Rivera M. Sánchez-Herrera leg., in MUAE) from the helena group is described, illustrated, diagnosed and compared with morphologically close species of the genus. Philogenia gaiae can be distinguished from its most similar congener P. minteri Dunkle, 1986 by the enlarged cerci and the club-like paraprocts. We also describe the female of P. macuma Dunkle, 1986, from a pair collected at Jatún Sacha Biological Reserve, which is also similar to P. gaiae and can be distinguished by the intersternite morphology, ovipositor length and vulvar lamina shape.

A further study on Franciscobasis Machado & Bedê, 2016 (Odonata: Coenagrionidae), a newly described genus from Minas Gerais, Brazil.

The genus Franciscobasis Machado & Bedê, 2016 is endemic to the Serra da Canastra National Park in Minas Gerais state, Brazil. Two species of Franciscobasis were described simultaneously with the genus description: F. franciscoi and F. sonia, the latter described only from females. Through morphological and molecular analysis, we investigated if F. sonia may represent the young female of F. franciscoi. Resulting data did not present adequate differences between females to characterize them as different species. Therefore, we suggest that F. sonia is a junior synonym of F. franciscoi, and the female of F. franciscoi goes through a complex ontogenetic color change.

Mechanisms of allopatric speciation in an Antillean damselfly genus (Odonata, Zygoptera): Vicariance or long-distance dispersal?

We have examined divergence times of the Antillean damselfly genus Hypolestes, to elucidate which mechanism of allopatric speciation, vicariance or long-distance dispersal, could better explain the currently observed disjunct distributions of this genus. Samples of the three extant species of the genus, Hypolestes clara (Jamaica), H. hatuey (Hispaniola) and H. trinitatis (Cuba), were collected. Mitochondrial and nuclear DNA gene fragments were amplified to reconstruct phylogenetic relationships and estimate divergence times in this genus. Hypolestes comprises currently three species, which consist in four geographically and genetically isolated lineages located in Jamaica, Hispaniola, Eastern Cuba and Central Cuba. Results of our analyses suggest that the three species diverged between ∼5.91 and 1.69 mya, and that the separation between the lineages from Central Cuba and Eastern Cuba occurred between ∼2.0 and 0.62 mya. Disjunct distributions in the genus Hypolestes can be better explained by a long-distance dispersal mechanism, since the divergence times of the three species do not coincide with the timeline formation of the geographic barriers between Cuba, Hispaniola and Jamaica. The Cuban lineages of H. trinitatis constitute different molecular operational taxonomic units (MOTU). The elevation of these MOTU to the species category requires the analysis of additional characters.

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.

Females of two species of Argia from Chapada dos Guimarães National Park, Brazil (Odonata: Coenagrionidae).

The female of Argia tupi Calvert, 1909 (BRAZIL, Mato Grosso, Chapada dos Guimarães National Park, Cachoeira do Marimbondo (15.4330° S, 55.7198° W, 370 m), 01 xi 2015) is described, illustrated and diagnosed based on comparison with sympatric species of Argia Rambur, 1842. We also augmented the description of Argia bicellulata (Calvert, 1909) female (BRAZIL, Mato Grosso, Chapada dos Guimarães National Park, Rio Paciencia (15.3438° S, 55.8322° W, 280 m), 25 x 2015).