A molecular phylogeny of the Petaluridae (Odonata: Anisoptera): A 160-Million-Year-Old story of drift and extinction.

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.

Genomic data provide insights into the classification of extant termites.

The higher classification of termites requires substantial revision as the Neoisoptera, the most diverse termite lineage, comprise many paraphyletic and polyphyletic higher taxa. Here, we produce an updated termite classification using genomic-scale analyses. We reconstruct phylogenies under diverse substitution models with ultraconserved elements analyzed as concatenated matrices or within the multi-species coalescence framework. Our classification is further supported by analyses controlling for rogue loci and taxa, and topological tests. We show that the Neoisoptera are composed of seven family-level monophyletic lineages, including the Heterotermitidae Froggatt, Psammotermitidae Holmgren, and Termitogetonidae Holmgren, raised from subfamilial rank. The species-rich Termitidae are composed of 18 subfamily-level monophyletic lineages, including the new subfamilies Crepititermitinae, Cylindrotermitinae, Forficulitermitinae, Neocapritermitinae, Protohamitermitinae, and Promirotermitinae; and the revived Amitermitinae Kemner, Microcerotermitinae Holmgren, and Mirocapritermitinae Kemner. Building an updated taxonomic classification on the foundation of unambiguously supported monophyletic lineages makes it highly resilient to potential destabilization caused by the future availability of novel phylogenetic markers and methods. The taxonomic stability is further guaranteed by the modularity of the new termite classification, designed to accommodate as-yet undescribed species with uncertain affinities to the herein delimited monophyletic lineages in the form of new families or subfamilies.

An unusually large genome from an unusually large stonefly: a chromosome-length genome assembly for the giant salmonfly, Pteronarcys californica (Plecoptera: Pteronarcyidae).

Pteronarcys californica (Newport 1848) is commonly referred to as the giant salmonfly and is the largest species of stonefly (Insecta: Plecoptera) in the western United States. Historically, it was widespread and abundant in western rivers, but populations have experienced a substantial decline in the past few decades, becoming locally extirpated in numerous rivers in Utah, Colorado, and Montana. Although previous research has explored the ecological variables conducive to the survivability of populations of the giant salmonfly, a lack of genomic resources hampers exploration of how genetic variation is spread across extant populations. To accelerate research on this imperiled species, we present a de novo chromosomal-length genome assembly of P. californica generated from PacBio HiFi sequencing and Hi-C chromosome conformation capture. Our assembly includes 14 predicted pseudo chromosomes and 98.8% of Insecta universal core orthologs. At 2.40 gigabases, the P. californica assembly is the largest of available stonefly assemblies, highlighting at least 9.5-fold variation in assembly size across the order. Repetitive elements (REs) account for much of the genome size increase in P. californica relative to other stonefly species, with the content of Class I retroelements alone exceeding the entire assembly size of all but two other species studied. We also observed preliminary suborder-specific trends in genome size that merit testing with more robust taxon sampling.

Species distribution models predict genetic isolation of Hetaerina vulnerata Hagen in Selys, 1853 (Odonata, Calopterygidae).

Understanding how past and current environmental conditions shape the demographic and genetic distributions of organisms facilitates our predictions of how future environmental patterns may affect populations. The Canyon Rubyspot damselfly (Odonata: Zygoptera: Hetaerina vulnerata) is an insect with a range distribution from Colombia to the arid southwestern United States, where it inhabits shaded mountain streams in the arid southwestern United States. Past spatial fragmentation of habitat and limited dispersal capacity of H. vulnerata may cause population isolation and genetic differentiation, and projected climate change may exacerbate isolation by further restricting the species' distribution. We constructed species distribution models (SDMs) based on occurrences of H. vulnerata and environmental variables characterizing the species' niche. We inferred seven current potential population clusters isolated by unsuitable habitat. Paleoclimate models indicated habitat contiguity in past conditions; projected models indicated some habitat fragmentation in future scenarios. Seventy-eight H. vulnerata individuals from six of the current clusters were sequenced via ddRADseq and processed with Stacks. Principal components and phylogeographic analyses resolved three subpopulations; Structure resolved four subpopulations. F ST values were low (<0.05) for nearby populations and >0.15 for populations separated by expanses of unsuitable habitat. Isolation by distance was an existing but weak factor in determining genomic structure; isolation by environment and the intervening landscape explained a significant proportion of genetic distance. Hetaerina vulnerata populations were shown to be isolated by a lack of tree canopy coverage, an important habitat predictor for oviposition and territoriality. Thus, H. vulnerata populations are likely separated and are genetically isolated. Integrating SDMs with landscape genetics allowed us to identify populations separated by distance and unsuitable habitat, explaining population genetic patterns and probable fates for populations under future climate scenarios.

Collection methods and distribution modeling for Strepsiptera in the United States.

The twisted-wing parasite order (Strepsiptera Kirby, 1813) is difficult to study due to the complexity of strepsipteran life histories, small body sizes, and a lack of accessible distribution data for most species. Here, we present a review of the strepsipteran species known from New York State. We also demonstrate successful collection methods and a survey of species carried out in an old-growth deciduous forest dominated by native New York species (Black Rock Forest, Cornwall, NY) and a private site in the Catskill Mountains (Shandaken, NY). Additionally, we model suitable habitats for Strepsiptera in the United States with species distribution modeling. We base our models on host distribution and climatic variables to inform predictions of where these twisted-wing parasites are likely to be found. This work provides a useful reference for the future collection of Strepsiptera.

Stolleagrion foghnielseni (Odonata, Cephalozygoptera, Dysagrionidae) gen. et sp. nov.: a new odonatan from the PETM recovery phase of the earliest Ypresian Fur Formation, Denmark.

We describe the new genus and species Stolleagrion foghnielseni n. gen. et sp. from the Fur Formation in northwestern Denmark based on a single fossil wing. This is the first odonatan described from the earliest part of the PETM recovery phase of the early Eocene. A combination of nine wing character states are considered to be diagnostic of the Dysagrionidae Cockrell only together with the cephalozygopteran head; however, the combination of these nine plus the presence of Ax0 is also diagnostic without the head. By this, we assign Stolleagrion foghnielseni to the Dysagrionidae and reassess the position of other odonates previously treated as cf. Dysagrionidae.

Chewing through challenges: Exploring the evolutionary pathways to wood-feeding in insects.

Decaying wood, while an abundant and stable resource, presents considerable nutritional challenges due to its structural rigidity, chemical recalcitrance, and low nitrogen content. Despite these challenges, certain insect lineages have successfully evolved saproxylophagy (consuming and deriving sustenance from decaying wood), impacting nutrient recycling in ecosystems and carbon sequestration dynamics. This study explores the uneven phylogenetic distribution of saproxylophagy across insects and delves into the evolutionary origins of this trait in disparate insect orders. Employing a comprehensive analysis of gut microbiome data, from both saproxylophagous insects and their non-saproxylophagous relatives, including new data from unexplored wood-feeding insects, this Hypothesis paper discusses the broader phylogenetic context and potential adaptations necessary for this dietary specialization. The study proposes the "Detritivore-First Hypothesis," suggesting an evolutionary pathway to saproxylophagy through detritivory, and highlights the critical role of symbiotic gut microbiomes in the digestion of decaying wood.

Phylogenomics of Tetraopes longhorn beetles unravels their evolutionary history and biogeographic origins.

Tetraopes longhorn beetles are known for their resistance to milkweed plant toxins and their coevolutionary dynamics with milkweed plants (Asclepias). This association is considered a textbook example of coevolution, in which each species of Tetraopes is specialized to feed on one or a few species of Asclepias. A major challenge to investigating coevolutionary hypotheses and conducting molecular ecology studies lies in the limited understanding of the evolutionary history and biogeographical patterns of Tetraopes. By integrating genomic, morphological, paleontological, and geographical data, we present a robust phylogeny of Tetraopes and their relatives, using three inference methods with varying subsets of data, encompassing 2-12 thousand UCE loci. We elucidate the diversification patterns of Tetraopes species across major biogeographical regions and their colonization of the American continent. Our findings suggest that the genus originated in Central America approximately 21 million years ago during the Miocene and diversified from the Mid-Miocene to the Pleistocene. These events coincided with intense geological activity in Central America. Additionally, independent colonization events in North America occurred from the Late Miocene to the early Pleistocene, potentially contributing to the early diversification of the group. Our data suggest that a common ancestor of Tetraopini migrated into North America, likely facilitated by North Atlantic land bridges, while closely related tribes diverged in Asia and Europe during the Paleocene. Establishing a robust and densely sampled phylogeny of Tetraopes beetles provides a foundation for investigating micro- and macroevolutionary phenomena, including clinal variation, coevolution, and detoxification mechanisms in this ecologically important group.

Tropical Origin, Global Diversification, and Dispersal in the Pond Damselflies (Coenagrionoidea) Revealed by a New Molecular Phylogeny.

The processes responsible for the formation of Earth's most conspicuous diversity pattern, the latitudinal diversity gradient (LDG), remain unexplored for many clades in the Tree of Life. Here, we present a densely sampled and dated molecular phylogeny for the most speciose clade of damselflies worldwide (Odonata: Coenagrionoidea) and investigate the role of time, macroevolutionary processes, and biome-shift dynamics in shaping the LDG in this ancient insect superfamily. We used process-based biogeographic models to jointly infer ancestral ranges and speciation times and to characterize within-biome dispersal and biome-shift dynamics across the cosmopolitan distribution of Coenagrionoidea. We also investigated temporal and biome-dependent variation in diversification rates. Our results uncover a tropical origin of pond damselflies and featherlegs ~105 Ma, while highlighting the uncertainty of ancestral ranges within the tropics in deep time. Even though diversification rates have declined since the origin of this clade, global climate change and biome-shifts have slowly increased diversity in warm- and cold-temperate areas, where lineage turnover rates have been relatively higher. This study underscores the importance of biogeographic origin and time to diversify as important drivers of the LDG in pond damselflies and their relatives, while diversification dynamics have instead resulted in the formation of ephemeral species in temperate regions. Biome-shifts, although limited by tropical niche conservatism, have been the main factor reducing the steepness of the LDG in the last 30 Myr. With ongoing climate change and increasing northward range expansions of many damselfly taxa, the LDG may become less pronounced. Our results support recent calls to unify biogeographic and macroevolutionary approaches to improve our understanding of how latitudinal diversity gradients are formed and why they vary across time and among taxa.

Long-term monitoring and analysis of Brood X cicada activity by distributed fiber optic sensing technology.

Brood X is the largest of the 15 broods of periodical cicadas, and individuals from this brood emerged across the Eastern United States in spring 2021. Using distributed acoustic sensing (DAS) technology, the activity of Brood X cicadas was monitored in their natural environment in Princeton, NJ. Critical information regarding their acoustic signatures and activity level is collected and analyzed using standard outdoor-grade telecommunication fiber cables. We believe these results have the potential to be a quantitative baseline for regional Brood X activity and pave the way for more detailed monitoring of insect populations to combat global insect decline. We also show that it is possible to transform readily available fiber optic networks into environmental sensors with no additional installation costs. To our knowledge, this is the first reported use case of a distributed fiber optic sensing system for entomological sciences and environmental studies.

Exploring chromosome evolution in 250 million year old groups of dragonflies and damselflies (Insecta:Odonata).

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.

The damselfly genus Furagrion Petrulevicius et al. (Odonata, Zygoptera) from the early Eocene Fur Formation of Denmark and the dysagrionoid grade.

The earliest Eocene odonate genus Furagrion Petrulevicius et al. from the Danish Fur Formation is revised based on eighteen specimens, two of which apparently have been lost since their publication. The holotype of Phenacolestes jutlandicus Henriksen, type species of Furagrion, is incomplete and lacks the characters currently used to differentiate species, genera and higher taxa in Odonata. We, therefore, propose that the holotype is set aside and a recently discovered nearly complete Fur Formation fossil is designated as neotype. Furagrion possesses all of the nine wing character states currently used along with head shape for diagnosing the Dysagrionidae; however, Furagrion has a characteristically zygopteran head, not the distinctive head shape of the suborder Cephalozygoptera. We, therefore, treat it as a zygopteran unassigned to family. These nine wing character states appear in different combinations not only in various Zygoptera and Cephalozygoptera, but also in the Frenguelliidae, an Eocene family of Argentina that may represent an unnamed suborder. We recognise these taxa as constituting a dysagrionoid grade, in which these character states appear either convergently or as symplesiomorphies. Furagrion morsi Zessin is synonymized with Phenacolestes jutlandicus Henriksen, syn. nov. and Morsagrion Zessin with Furagrion Petrulevicius, Wappler, Wedmann, Rust, and Nel, syn. nov.

Evolution of Odonata: genomic insights.

Odonata is an order of insects that comprises ∼6500 species. They are among the earliest flying insects, and one of the first diverging lineages in the Pterygota. Odonate evolution has been a topic of research for over 100 years, with studies focusing primarily on their flight behavior, color, vision, and aquatic juvenile lifestyles. Recent genomics studies have provided new interpretations about the evolution of these traits. In this paper, we look at how high-throughput sequence data (i.e. subgenomic and genomic data) have been used to answer long-standing questions in Odonata ranging from evolutionary relationships to vision evolution to flight behavior. Additionally, we evaluate these data at multiple taxonomic levels (i.e. ordinal, familial, generic, and population) and provide comparative analysis of genomes across Odonata, identifying features of these new data. Last, we discuss the next two years of Odonata genomic study, with context about what questions are currently being tackled.

A Chromosome-length Assembly of the Black Petaltail (Tanypteryx hageni) Dragonfly.

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.

Taxonomic revision of the Neotropical genus Heteragrion Selys, 1862 (Zygoptera: Heteragrionidae): male morphology, new species and illustrated key.

In this study, we have comprehensively presented taxonomic information on all 62 known species of Heteragrion Selys, 1862, which includes illustrations, photographs, diagnostic characters, and a key to males. Our analysis is based on the examination of over 900 specimens from 19 different collections worldwide, encompassing the type material for at least 42 species. Furthermore, we have described a new species, Heteragrion corderoi sp. nov. (, BRAZIL, So Paulo state, Campos do Jordo, Condomnio Paradise, 24.i.1999, (Coordinates: -22.7072, -45.5894, 1796 m asl), F.A.A. Lencioni leg., LABECO), which we identified from a male that was previously considered to be a paratype of H. mantiqueirae Machado, 2006 and additional specimens collected in Campos do Jordo, So Paulo state.

Review of Nasuconia Sakakibara, 2006 (Hemiptera: Membracidae) with description of three new species.

The treehopper genus Nasuconia Sakakibara, 2006 previously included four species and was recorded only from Brazil. Here we provide a revised diagnosis of the genus and describe three new species: Nasuconia ellenfutterae sp. nov. from Ecuador, Nasuconia guianensis sp. nov. from French Guiana and Nasuconia yasuni sp. nov. from Ecuador. The genus can be distinguished by the following combination of characters: frontoclypeus conical with transverse grooves, obliquely projected forward at least 1/3 of its length beyond suprantennal margin; pronotum navicular, low, punctate, with longitudinal elevated lines or nodes; first valvulae with ventralinterlocking device distinctively sinuate. Two informal species groups are recognized based on characters of the head, forewing and leg chaetotaxy. A key to species, photographs, updated morphological descriptions, and the first descriptions of the female and male genitalia of Nasuconia species are provided. Comparisons of cucullate setae and fine abdominal integument structures are also made using scanning electron microscopy.

Review of Cyphotes Burmeister, 1835 (Hemiptera: Membracidae) with the description of a related new genus.

The Neotropical treehopper genus Cyphotes Burmeister, 1835 (= Aspona Stal, 1862 syn. nov.) is redefined. Cyphotes contains only two species, the type species Cyphotes nodosa Burmeister, 1835 (= Aspona bullata Stl, 1862 syn. nov.) and Cyphotes quadrinodosa (Fonseca & Diringshofen, 1969) reinstated. comb. Allocyphotes gen. nov. (type species Cyphotes insolita Goding, 1929) is proposed to accommodate two other species previously placed in Cyphotes, Allocyphotes pompanoni (Boulard, 2011) comb. nov. and Allocyphotes colombiensis (Gonzlez-Mozo, 2017) comb. nov., and three new species from Ecuador: A. flavus sp. nov., A. waoraniorum sp. nov and A. robertoi sp. nov. for a total of six species. Illustrations, including genitalia images, new locality records and keys to genera and species are provided.

Danowhetaksa gen. nov. with two species from the early Eocene lst Formation from Denmark, the first Palearctic Whetwhetaksidae (Odonata: Cephalozygoptera).

We propose Danowhetaksa n. gen. (Odonata: Whetwhetaksidae) with two new species: D. birgitteae n. gen. et sp. and D. rusti n. gen. et sp. from the earliest Ypresian Stolleklint clay of the lst Formation in northwestern Denmark. Whetwhetaksidae has previously been known only from the Ypresian Okanagan Highlands of far-western North America, the new records are, therefore, the first from the Palearctic Region.

Taxonomic note on the species status of Epiophlebiadiana (Insecta, Odonata, Epiophlebiidae), including remarks on biogeography and possible species distribution.

The species included in the genus Epiophlebia Calvert, 1903 represent an exception within Recent lineages - they do not belong to either dragonflies (Anisoptera) nor damselflies (Zygoptera). Nowadays, the genus is solely known from the Asian continent. Due to their stenoecious lifestyle, representatives of Epiophlebia are found in often very small relict populations in Nepal, Bhutan, India, Vietnam, China, North Korea, and Japan. We here present a taxonomic re-evaluation on the species status of Epiophlebiadiana Carle, 2012, known from the Sichuan province in China, supplemented with a morphological character mapping on a genetic tree to highlight synapomorphies of E.diana and E.laidlawi Tillyard, 1921. We conclude that E.diana is a junior synonym of E.laidlawi. Furthermore, we discuss the Recent distribution of the group, allowing for predictions of new habitats of representatives of this group.

Evolutionary history and divergence times of Odonata (dragonflies and damselflies) revealed through transcriptomics.

Dragonflies and damselflies are among the earliest flying insects with extant representatives. However, unraveling details of their long evolutionary history, such as egg laying (oviposition) strategies, is impeded by unresolved phylogenetic relationships, particularly in damselflies. Here we present a transcriptome-based phylogenetic reconstruction of Odonata, analyzing 2,980 protein-coding genes in 105 species representing nearly all the order's families. All damselfly and most dragonfly families are recovered as monophyletic. Our data suggest a sister relationship between dragonfly families of Gomphidae and Petaluridae. According to our divergence time estimates, both crown-Zygoptera and -Anisoptera arose during the late Triassic. Egg-laying with a reduced ovipositor apparently evolved in dragonflies during the late Jurassic/early Cretaceous. Lastly, we also test the impact of fossil choice and placement, particularly, of the extinct fossil species, †Triassolestodes asiaticus, and †Proterogomphus renateae on divergence time estimates. We find placement of †Proterogomphus renateae to be much more impactful than †Triassolestodes asiaticus.