We present an updated classification for extant dragonflies and damselflies (Odonata) and summarize new insights gained over the past two decades. Our focus is on taxa of family-level and higher and we indicate subfamilies only when their monophyly is currently undisputed and well-supported by phylogenetic analyses. The superfamily Calopterygoidea was known to be polyphyletic and based on recent phylogenomic data is divided into nine superfamilies, of which eight are recognized for the first time (Amphipterygoidea stat. nov., Euphaeoidea stat. nov., Megapodagrionoidea stat. nov., Mesopodagrionoidea stat. nov., Philogangoidea stat. nov., Polythoroidea stat. nov., Priscagrionoidea stat. nov., Tatocnemidoidea stat. nov.). At present, midway through 2025, odonates are divided into three suborders, 17 superfamilies, 55 families and 687 genera containing 6447 species (May 1, 2025). We give an overview of the distribution of the families across major biogeographical realms. Except for Amanipodagrionidae, nymphs of at least some of the species of all families are known. Our understanding of the classification of dragonflies and damselflies has greatly improved in the past two decades largely due to phylogenetic inferences based on molecular studies. We expect that in the next few years the last remaining issues regarding the higher-level phylogeny and classification, including the position of the South American genus Sciotropis and the division into subfamilies of families such as Coenagrionidae, Aeshnidae, Gomphidae and Libellulidae, will be better refined by the acquisition of additional morphological and genomic data.
Mountains and islands provide an opportunity for studying the biogeography of diversification and population fragmentation. Aotearoa (New Zealand) is an excellent location to investigate both phenomena due to alpine emergence and oceanic separation. While it would be expected that separation across oceanic and elevation gradients are major barriers to gene flow in animals, including aquatic insects, such hypotheses have not been thoroughly tested in these taxa. By integrating population genomic from subgenomic Anchored-Hybrid Enrichment sequencing, ecological niche modeling, and morphological analyses from scanning-electron microscopy, we show that tectonic uplift and oceanic vicariance are implicated in speciation and population structure in Kapokapowai (Uropetala) dragonflies. Although Te Moana o Raukawa (Cook Strait) is likely responsible for some of the genetic structure observed, speciation has not yet occurred in populations separated by the strait. We find that the altitudinal gradient across Kā Tiritiri-o-te-Moana (the Southern Alps) is not impervious, but it significantly restricts gene flow between the aforementioned species. Our data support the hypothesis of an active colonization of Kā Tiritiri-o-te-Moana by the ancestral population of Kapokapowai, followed by a recolonization of the lowlands. These findings provide key foundations for the study of lineages endemic to Aotearoa.
Odonata is an insect order that comprises ~6420 described species distributed among three suborders. Here we review the dragonflies, or Anisoptera, a suborder which has 10 described families, and forms a sister group relationship with the ‘Anisozygoptera’ [Epiprocta: Anisoptera + ‘Anisozygoptera’], a paraphyletic grouping containing just one extant family and many extinct taxa. Dragonflies are charismatic, ancient, fast flying, predatory insects whose natural history is woven into the fabric of human culture. As objects of systematic study, their relative position in the insect tree of life relates to the origin of flight, of freshwater juvenile lifestyles, and the evolution of color and vision. In this paper, we will review the past, present, and future of Odonatology, highlighting the work being done all across the globe to revise and define the suborder Anisoptera. In particular, we focus on ‘problematic’ areas in the tree, the Gomphidae and Libellulidae, with an emphasis on how taxon and data sampling are solving these problems in the current global work.
Mountains and islands are both model systems for studying the biogeography of diversification and population fragmentation. Aotearoa is an excellent location to study both phenomena due to alpine emergence and oceanic separation. While it would be expected that separation across Te Moana o Raukawa and elevation gradients are major barriers to gene flow in aquatic insects, such hypotheses have not been thoroughly explored in these taxa. Here, we show that mountains and oceanic separation function as semi-permeable barriers for Kapokapowai dragonflies. We show that, although Te Moana o Raukawa, is likely responsible for some of the genetic structure observed, speciation has not yet occurred in populations separated by the strait. Although there is no evidence that they are an impervious barrier, Kā Tiritiri-o-te-Moana do represent a major barrier to gene flow between named species. The distribution of alpine Kapokapowai can also not be explained by the ancestral populations simply rising with the uplifted tectonic plates. Although further research is needed, our findings suggest that the ancestral Kapokapowai colonized alpine habitats after their formation, and then radiated back out to lowlands. These findings suggest that aquatic insects could be an exciting new frontier in the study of the biogeography of Aotearoa. ### Competing Interest Statement The authors have declared no competing interest.
Environmental context Pesticides are critical to agriculture and food production but require ecological risk assessments. Although most risk assessments require data from vertebrate animal testing, we have developed an approach to assess risk to fish, birds and mammals using other means. This approach could help to ensure protection of the environment while minimising animal testing.Rationale Recent directives to reduce animal testing have implications for ecological risk assessment, as several vertebrate tests are used to support these assessments. Therefore, a modern approach was devised to address these key knowledge needs without the use of chemical-specific vertebrate testing.Methodology An ecological risk assessment for a novel acetyl-coenzyme A carboxylase (ACCase) inhibitor herbicide was conducted using alternative lines of evidence. For fish, chemical toxicity distributions were constructed to quantify the probability of effects, and these distributions were compared with exposure estimates for a representative use in soybeans. The effect distributions were further refined based on invertebrate toxicity and partitioning behaviour. For birds and mammals, a joint probability curve was constructed by integrating chemical toxicity distributions and Kenaga exposure distributions.Results The lines of evidence presented in this predictive risk assessment suggest the intended use of a new ACCase inhibitor is unlikely to affect fish, birds, or mammals. Exposure was unlikely to exceed effect estimates, regardless of whether they were derived based on chemical-read across, invertebrate toxicity, or partitioning behaviour.Discussion Key knowledge needs for ecological risk assessment can be informed by lines of evidence that do not require animal testing. The present study demonstrates such an approach by comparing predicted exposure and effects, which are expected to be protective. This predictive approach can be extended to other active ingredients and chemical classes, as well as other taxonomic groups of interest. Future research should aim to integrate new approach methods in a predictive risk assessment framework.
Insect coloration has evolved in response to multiple pressures, and in Odonata (dragonflies and damselflies) a body of work supports a role of wing color in a variety of visual signals and potentially in thermoregulation. Previous efforts have focused primarily on melanistic coloration even though wings are often multicolored, and there has yet to be comprehensive comparative analyses of wing color across broad geographic regions and phylogenetic groups. Percher vs. flier flight-style, a trait with thermoregulatory and signaling consequences, has not yet been studied with regard to color. We used a new color clustering approach to quantify color across a dataset of over 8,000 odonate wing images representing 343 Nearctic species. We then utilized phylogenetically informed Bayesian zero-inflated mixture models to test how color varies with mean ambient temperature, body size, sex and flight-style. We found that wing coloration clustered into two groups across all specimens - light brown-yellow and black-dark brown - with black-dark brown being a much more cohesive grouping. Male perchers have a greater proportion of black-dark brown color on their wings as do species with longer wings. In colder climates, odonates were more likely to have black-dark brown color present, but we found no relationship between the proportion of black and temperature. Light brown-yellow showed similar scaling with wing length, but no relationship with temperature. Our results suggest that black-dark brown coloration may have a limited role in thermoregulation, while light brown-yellow does not have such a role. We also find that the odonate sexes are divergent in wing color in percher species only, suggesting a strong role for color in signaling in more territorial males. Our research contributes to an understanding of complex interactions driving ecological and evolutionary dynamics of color in animals.
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.
More than tools for managing physical and digital objects, museum collection management systems (CMS) serve as platforms for structuring, integrating, and making accessible the rich data embodied by natural history collections. Here we describe Arctos, a scalable community solution for managing and publishing global biological, geological, and cultural collections data for research and education. Specific goals are to: (1) Describe the core features and implementation of Arctos for a broad audience with respect to the biodiversity informatics principles that enable high quality research; (2) Highlight the unique aspects of Arctos; (3) Illustrate Arctos as a model for supporting and enhancing the Digital Extended Specimen concept; and (4) Emphasize the role of the Arctos community for improving data discovery and enabling cross-disciplinary, integrative studies within a sustainable governance model. In addition to detailing Arctos as both a community of museum professionals and a collection database platform, we discuss how Arctos achieves its richly annotated data by creating a web of knowledge with deep connections between catalog records and derived or associated data. We also highlight the value of Arctos as an educational resource. Finally, we present the financial model of fiscal sponsorship by a nonprofit organization, implemented in 2022, to ensure the long-term success and sustainability of Arctos.
Museum collections house millions of objects and associated data records that document biological and cultural diversity. In recent decades, digitization efforts have greatly increased accessibility to these data, thereby revolutionizing interdisciplinary studies in evolutionary biology, biogeography, epidemiology, cultural change, and human-mediated environmental impacts. Curators and collection managers can make museum data as accessible as possible to scientists and learners by using a collection management system. However, selecting a system can be a challenging task. Here, we describe Arctos, a community solution for managing and accessing collections data for research and education. Specific goals are to: (1) Describe the core elements of Arctos for a broad audience with respect to the biodiversity informatics principles that enable high quality research; (2) Highlight the unique aspects of Arctos; (3) Illustrate Arctos as a model for supporting and enhancing the Digital Extended Specimen; and (4) Emphasize the role of the Arctos community for improving data discovery and enabling cross-disciplinary, integrative studies within a sustainable governance model. In addition to detailing Arctos as both a community of museum professionals and a collection database platform, we discuss how Arctos achieves its richly annotated data by creating a web of knowledge with deep connections between catalog records and derived or associated data. We also highlight the value of Arctos as an educational resource. Finally, we present a financial model of fiscal sponsorship by a non-profit organization, implemented in 2022, to ensure the long-term success and sustainability of Arctos. We attribute Arctos’ longevity of nearly three decades to its core development principles of standardization, flexibility, interdisciplinarity, and connectivity within a nimble development model for addressing novel needs and information types in response to changing technology, workflows, ethical considerations, and regulations.### Competing Interest StatementThe authors have declared no competing interest.
The use of gDNAs isolated from museum specimens for high throughput sequencing, especially targeted sequencing in the context of phylogenetics, is a common practice. Yet, little understanding has been focused on comparing the quality of DNA and results of sequencing museum DNAs. Dragonflies and damselflies are ubiquitous in freshwater ecosystems and are commonly collected and preserved insects in museum collections hence their use in this study. However, the history of odonate preservation across time and museums has resulted in wide variability in the success of viable DNA extraction, necessitating an assessment of their usefulness in genetic studies. Using Anchored Hybrid Enrichment probes, we sequenced DNA from samples at 2 museums, 48 from the American Museum of Natural History (AMNH) in NYC, USA and 46 from the Naturalis Biodiversity Center (RMNH) in Leiden, Netherlands ranging from global collection localities and across a 120-year time span. We recovered at least 4 loci out of an >1,000 locus probe set for all samples, with the average capture being ~385 loci (539 loci on average when a clade of ambiguous taxa omitted). Neither specimen age nor size was a good predictor of locus capture, but recapture rates differed significantly between museums. Samples from the AMNH had lower overall locus capture than the RMNH, perhaps due to differences in specimen storage over time.
In recent decades, a lack of available knowledge about the magnitude, identity and distribution of biodiversity has given way to a taxonomic impediment where species are not being described as fast as the rate of extinction. Using Machine Learning methods based on seven different algorithms (LR, CART, KNN, GNB, LDA, SVM and RFC) we have created an automatic identification approach for odonate genera, through images of wing contours. The training population is composed of the collected specimens that have been digitized in the framework of the NSF funded Odomatic and TOWD projects. Each contour was pre-processed, and 80 coefficients were extracted for each specimen. These form a database with 4656 rows and 80 columns, which was divided into 70% for training and 30% for testing the classifiers. The classifier with the best performance was a Linear Discriminant Analysis (LDA), which discriminated the highest number of classes (100) with an accuracy value of 0.7337, precision of 0.75, recall of 0.73 and a F1 score of 0.73. Additionally, two main confusion groups are reported, among genera within the suborders of Anisoptera and Zygoptera. These confusion groups suggest a need to include other morphological characters that complement the wing information used for the classification of these groups thereby improving accuracy of classification. Likewise, the findings of this work open the door to the application of machine learning methods for the identification of species in Odonata and in insects more broadly which would potentially reduce the impact of the taxonomic impediment.
Despite claims of an insect decline worldwide, our understanding of extinction risk in insects is incomplete. Using bionomic data of all odonate (603 dragonflies and damselflies) North American species, we assessed (i) regional extinction risk and whether this is related to local extirpation; (ii) whether these two patterns are similar altitudinally and latitudinally; and (iii) the areas of conservation concern. We used geographic range size as a predictor of regional extinction risk and body size, thermal limits and habitat association as predictors of local extirpation. We found that (i) greater regional extinction risk is related to narrow thermal limits, lotic habitat use and large body size (this in damselflies but not dragonflies); (ii) southern species are more climate tolerant but with more limited geographic range size than northern species; and (iii) two priority areas for odonate conservation are the cold temperate to sub-boreal northeastern USA and the transversal neo-volcanic system. Our approach can be used to estimate insect extinction risk as it compensates for the lack of abundance data.
Insects are reportedly experiencing widespread declines, but we generally have sparse data on their abundance. Correcting this shortfall will take more effort than professional entomologists alone can manage. Volunteer nature enthusiasts can greatly help to monitor the abundance of dragonflies and damselflies (Odonata), iconic freshwater sentinels and one of the few nonpollinator insect groups appreciated by the public and amenable to citizen science. Although counting individual odonates is common in some locations, current data will not enable a global perspective on odonate abundance patterns and trends. Borrowing insight from butterfly monitoring efforts, we outline basic plans for a global volunteer network to count odonates, including organizational structure, advertising and recruiting, and data collection, submission, and synthesis. We hope our proposal serves as a catalyst for richer coordinated efforts to understand population trends of odonates and other insects in the Anthropocene.
A recently introduced, ecologically dominant, exotic ant species, Nylanderia fulva, is invading the Southeastern United States and Texas. We evaluate how this invader impacts diversity and abundance of co-occurring ants and other arthropods in two grasslands. N. fulva rapidly attains densities up to 2 orders of magnitude greater than the combined abundance of all other ants. Overall ant biomass increases in invaded habitat, indicating that N. fulva exploits resources not fully utilized by the local ant assemblage. At high density, as N. fulva spreads, it eliminates the current ecologically dominant invasive ant, red imported fire ants (Solenopsis invicta). Compared to imported fire ant dominated habitat, N. fulva invasion zones have lower non-ant arthropod species richness and abundance with impacts differing by trophic category. Further, N. fulva reduces abundance and species richness of the remainder of the ant assemblage and does so in a non-random manner: impacting species with small sized workers much less than species with larger workers. In these and other ant assemblages with a large exotic component, the exotics tend to be small bodied species. As a result, N. fulva almost completely eliminates regionally distributed species, but leaves globally distributed species largely unaffected, thereby systematically favoring introduced over native diversity. S. invicta impacts wildlife and arthropod assemblage structure and is nearly ubiquitous in non-forested habitats of the Southeastern United States and Texas. Its displacement by N. fulva has critical implications for the natural systems of this region.
A dramatic increase in interest in the North American Odonata (dragonfly and damselfly) fauna in the last few years has lead to many new discoveries, particularly in southern areas where subtropical species seem to be expanding their range northward. I report the occurrence of eight Odonata species previously unknown from Texas: Argia oenea Hagen in Selys [Coenagrionidae], Enallagma antennaturn (Say) [Coenagrionidae], Leptobasis melinogaster Gonzales-Soriano [Coenagrionidae], Aeshna persephone Donnelly [Aeshnidae), Anax concolor Bauer [Aeshnidael, Phyllocycla breviphylla Belle [Gomphidae], Erythemis attala (Selys) [Libellulidael, and Erythemis mithroides (Brauer) [Libellulidael. These discoveries include four species previously unknown from the United States and the first occurrence of the genus Leptobasis in the country. Additionally, I discuss recent records of several other species rarely reported from Texas.
Previously, no Odonata have been reported from 44 Texas counties (17%), mainly from the northern Panhandle. Adult dragonflies and damselflies collected since September of 1999 are reported from 24 sites in 14 counties throughout the Texas Panhandle. A total of 35 species is discussed, representing 73 new county records and 4 new state records. First records of Odonata are included for 6 counties.