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Dentifibula viburni (Felt) (Diptera: Cecidomyiidae), type species of Dentifibula Felt, is redescribed with illustrations and compared with its congeners. The genus is differentiated from the generally similar Lestodiplosis Kieffer. As far as presently known, larvae of Dentifibula prey on Diaspididae (Hemiptera: Coccoidea). The nine other species of Dentifibula are diagnosed. Dentifibula ceylanica Felt is removed from synonymy under Dentifibula obtusiloba Felt.

Supplemental feeding can increase the overall health of animals but also can have variable effects on animals dealing with parasites. Furthermore, the spatial and temporal effects of food supplementation on host-parasite interactions remain poorly understood, likely because large-scale, coordinated efforts are difficult. Here, we introduce the Nest Parasite Community Science Project (hereafter “Project”), which is a community-based science project that coordinates studies with bird nest box “landlords” from the public and scientific community. The Project was established to understand broad ecological patterns between box-nesting birds and their nest parasite community. The goal of this study was to use data collected by the Project to determine the effect of food supplementation on Eastern Bluebirds (Sialia sialis) and their nest parasite community across the geographic range of the bluebirds from 2018-2020. Nest box landlords either fed their bluebirds mealworms or not, then followed the nesting success of the birds (number of eggs, percent hatched, percent fledged). Nests were sent to our lab where we identified and quantified all nest parasite taxa. Birds from the western range laid more eggs than the eastern range, but this spatial effect was not observed for hatching success. However, food supplementation increased hatching success across years and the study area. We primarily found parasitic blow flies and fleas in the nests, which were restricted to northern latitudes. Within this geographic area, food supplementation affected blow fly abundance, but this effect varied across years, with abundances increasing, decreasing, or not changing, depending on the year. Interestingly, more landlords at southern latitudes fed bluebirds compared to northern latitudes, which contradicted the results of other community-based science projects. Our study demonstrates the importance of coordinated studies across years and locations to understand the effects of environmental heterogeneity on host-parasite dynamics.

The Christmas tree industry is mainly supplied with seeds from natural stands and genetically improved germplasm from clonal seed orchards (CSOs). Chalcid wasps (Megastigmus spp.) have a negative e...

The plant genus Corylus is an economically important crop, valued especially for its nuts. Numerous pathogens and harmful phytophagous arthropods are known to damage hazelnuts. We report on a new eriophyoid mite, Rhyncaphytoptus corylivagrans n. sp., and the first record of Coptophylla lamimani both collected from leaves of American hazelnut (Corylus americana) in North Carolina, USA. Including our new data, the complex of eriophyoids from Corylus comprises 15 species from three families: Phytoptidae (2 spp.), Eriophyidae (11 spp.), and Diptilomiopidae (2 spp.). We obtained sequences of three genes (Cox1, D1-D5 28S, and ITS1-5.8S-ITS2), applied BLAST and tree-based approaches for identification of R. corylivagrans n. sp., and performed the first molecular phylogenetic analysis focused on Rhyncaphytoptinae. Among the three genes, Cox1 showed better power when used for BLAST searches. Combined molecular phylogenetic analyses inferred R. corylivagrans n. sp. as sister to R. betulae, determined several moderately supported host-specific lineages of rhyncaphytoptines, and indicated a close relationship of the new species with members of the genus Rhinotergum. In two Rhinotergum spp. from Rosaceae, confocal microscopy revealed a new structure, the needle-like anterior process of the prodorsal shield, which is absent in R. corylivagrans n. sp. Additionally, the elements of the anal secretory apparatus presumably associated with silk-production and hypothesized as a synapomorphy of Eriophyoidea, were detected in the new species, providing the first documented report of this structure in Diptilomiopidae. Our study contributes to knowledge on the biodiversity of phytoparasites associated with hazelnuts and calls for future comparative phylogenetics of Diptilomiopidae.

The invasive woodboring beetle emerald ash borer Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) first was detected in the US in 2002 near Detroit, Michigan, USA. Since then it has continued to expand its range into the southern and midwestern sections of the country. Emerald ash borer was discovered in North Carolina, USA, in 2013, and is currently reported from more than 60 counties in the state. The present study was undertaken to begin determining the phenology of emerald ash borer in its newly expanded range below 40 °N latitude in North America. Here we report 4 yr of data on the overwintering stages of emerald ash borer in north-central North Carolina, obtained by debarking infested trees harvested from a single site near the northern border with Virginia, USA, with 1-yr additional data from a site about 39 km away. Results indicated that most emerald ash borers overwinter as fourth instars, with a small proportion advancing to the J-larva stage. Parasitoids that emerged from cocoons collected from emerald ash borer galleries and from logs held in emergence cages also were identified. These Hymenoptera include specimens of Xorides (Exomus) humeralis (Say) (Ichneumonidae), Atanycolus cf. cappaerti Marsh and Strazanac (Braconidae), Balcha indica (Mani & Kau) (Eupelmidae), Spathius sp. Nees (Braconidae), and Wroughtonia sp. Cameron (Braconidae).

In this brief, we report the observation of downy mildew caused by Peronospora chenopodii-ambrosioidis on epazote (Dysphania ambrosioides) in North Carolina, U.S.A. We performed morphological characterization of the sporangia and sporangiophores for identification. We also confirmed the identity of the pathogen by performing an alignment and generating a maximum likelihood phylogeny of the concatenated internal transcribed spacer region and cytochrome c oxidase subunit I sequences.

The purpose of mounting an immune response is to destroy pathogens, but this response comes at a physiological cost, including the generation of oxidative damage. However, many studies on the effects of immune challenges employ a single high dose of a simulated infection, meaning that the consequences of more mild immune challenges are poorly understood. We tested whether the degree of immunological challenge in tree swallows (Tachycineta bicolor) affects oxidative physiology and body mass, and whether these metrics correlate with parasitic nest mite load. We injected 14 day old nestlings with 0, 0.01, 0.1 or 1 mg lipopolysaccharide (LPS) per kg body mass, then collected a blood sample 24 h later to quantify multiple physiological metrics, including oxidative damage (i.e. d-ROMs), circulating amounts of triglyceride and glycerol, and levels of the acute phase protein haptoglobin. After birds had fledged, we identified and counted parasitic nest mites (Dermanyssus spp. and Ornithonyssus spp.). We found that only nestlings injected with 1 mg LPS kg−1 body mass, which is a common dosage in ecoimmunological studies, lost more body mass than individuals from other treatment groups. However, every dose of LPS resulted in a commensurate increase in oxidative damage. Parasitic mite abundance had no effect on oxidative damage across treatments. The amount of oxidative damage correlated with haptoglobin levels, suggesting compensatory mechanisms to limit self-damage during an immune response. We conclude that while only the highest-intensity immune challenges resulted in costs related to body mass, even low-intensity immune challenges result in detectable increases in oxidative damage.

Haemaphysalis longicornis is a tick indigenous to eastern Asia and an important vector of human and animal disease agents, resulting in such outcomes as human hemorrhagic fever and reduction of production in dairy cattle by 25%. H. longicornis was discovered on a sheep in New Jersey in August 2017 (1). This was the first detection in the United States outside of quarantine. In the spring of 2018, the tick was again detected at the index site, and later, in other counties in New Jersey, in seven other states in the eastern United States, and in Arkansas. The hosts included six species of domestic animals, six species of wildlife, and humans. To forestall adverse consequences in humans, pets, livestock, and wildlife, several critical actions are indicated, including expanded surveillance to determine the evolving distribution of H. longicornis, detection of pathogens that H. longicornis currently harbors, determination of the capacity of H. longicornis to serve as a vector for a range of potential pathogens, and evaluation of effective agents and methods for the control of H. longicornis. Published version

Because carnivorous plants rely on arthropods as pollinators and prey, they risk consuming would-be mutualists. We examined this potential conflict in the Venus flytrap (Dionaea muscipula), whose pollinators were previously unknown. Diverse arthropods from two classes and nine orders visited flowers; 56% of visitors carried D. muscipula pollen, often mixed with pollen of coflowering species. Within this diverse, generalized community, certain bee and beetle species appear to be the most important pollinators, on the basis of their abundance, pollen load size, and pollen fidelity. Dionaea muscipula prey spanned four invertebrate classes and 11 orders; spiders, beetles, and ants were most common. At the family and species levels, few taxa were shared between traps and flowers, yielding a near-zero value of niche overlap for these potentially competing structures. Spatial separation of traps and flowers may contribute to partitioning the invertebrate community between nutritional and reproductive functions in D. muscipula.

The history of people living with insects, spiders and their relatives is long, probably as long as humans have been using fixed domiciles (e.g., caves). Studies of caves inhabited by prehistoric people 26,000 years ago suggest arthropod pests already lived alongside our ancestors in those caves (Araújo et al. 2009). Arthropods are also both abundant and diverse in domestic archaeological sites from agricultural civilizations in Egypt, Israel and Europe (Switzerland and Greenland). Arthropods are especially common in association with stored food products and livestock (Panagiotakopulu 2001; Overgaard Nielsen, Mahler, and Rasmussen 2000; Kislev, Hartmann, and Galili 2004).

Increased market viability of harvest residues as forest bioenergy feedstock may escalate removal of coarse woody debris in managed forests. Meanwhile, many forest invertebrates use coarse woody debris for cover, food, and reproduction. Few studies have explicitly addressed effects of operational-scale woody biomass harvesting on invertebrates following clearcutting. Therefore, we measured invertebrate community response to large-scale harvest residue removal and micro-site manipulations of harvest residue availability in recently clearcut, intensively managed loblolly pine (Pinus taeda) forests in North Carolina (NC; n = 4) and Georgia (GA; n = 4), USA. We captured 39,794 surface-active invertebrates representing 171 taxonomic groups using pitfall traps situated among micro-site locations (i.e., purposefully retained piles of hardwood stems and piles of conifer stems and areas without coarse woody debris in NC; windrows and no windrows in GA). Micro-site locations were located within six, large-scale treatments (7.16-14.3 ha) in clearcuts. Large-scale treatments represented intensive harvest residue removal, 15% and 30% harvest residue retention, and no harvest residue removal. In NC, ground beetles (Coleoptera: Carabidae) and crickets (Orthoptera: Gryllidae) were three times more abundant in treatments with no harvest residue removal than those with the most intensive harvest residue removal and were reduced in treatments that retained 15% or 30% of harvest residues, although not significantly. Invertebrate taxa richness was greater at micro-site locations with retained hardwood and pine (Pinus spp.) harvest residues than those with minimal amounts of coarse woody debris. In both states, relative abundances of several invertebrate taxa, including cave crickets (Orthoptera: Rhaphidophoridae), fungus gnats (Diptera: Mycetophilidae and Sciaridae), millipedes (Diplopoda), and wood roaches (Blattodea: Ectobiidae), were greater at micro-site locations with retained harvest residues than those with minimal coarse woody debris. Intensified woody biomass harvesting without retention of ≥15% of harvest residue volume may reduce invertebrate taxa richness and abundances of some key invertebrate taxa in regenerating stands. Further, harvest residue management during and after woody biomass harvesting may be an important consideration for maintaining invertebrate diversity and conserving invertebrates that are influential in the maintenance of ecosystem function and integrity in young forests.

The indoor biome is a novel habitat which recent studies have shown exhibit not only high microbial diversity, but also high arthropod diversity. Here, we analyze findings from a survey of 50 houses (southeastern USA) within the context of additional survey data concerning house and room features, along with resident behavior, to explore how arthropod diversity and community composition are influenced by physical aspects of rooms and their usage, as well as the lifestyles of human residents. We found that indoor arthropod diversity is strongly influenced by access to the outdoors and carpeted rooms hosted more types of arthropods than non-carpeted rooms. Arthropod communities were similar across most room types, but basements exhibited more unique community compositions. Resident behavior such as house tidiness, pesticide usage, and pet ownership showed no significant influence on arthropod community composition. Arthropod communities across all rooms in houses exhibit trophic structure—with both generalized predators and scavengers included in the most frequently found groups. These findings suggest that indoor arthropods serve as a connection to the outdoors, and that there is still much yet to be discovered about their impact on indoor health and the unique ecological dynamics within our homes.

Diaphania costata (Fabricius) (Lepidoptera: Crambidae) is a leaffolder on Apocynaceae in the Neotropics and the southeastern United States. Adult moths photographed or captured at light in the United States have been commonly misidentified, and the species’ Nearctic distribution has been poorly documented. We provide diagnostic characters for larval and adult stages, compare it to related Old World taxa, and describe the damage to ornamental Vinca L. The species is not restricted to Texas but occurs as far east as the Atlantic Coast and as far north as Virginia. Specimens collected in daily surveys over many years indicate that the range of D. costata spread naturally from Louisiana and Texas from the 1980s into the 2000s.

The cypress weevil, Eudociminus mannerheimii (Boheman, 1836), is reported from northwestern Arkansas (new state record). The suspected host in this area is eastern red cedar (Juniperus virginiana L.), which represents a new host record. Additional new host records from arborvitae (Thuja L.) in North Carolina are reported. A brief redescription of the adult that expands upon the original description and photographs are included. Although cypress weevils are not generally considered pestiferous, a case of landscape trees likely killed by this species is included.

The indoor environment created by the construction of homes and other buildings is often considered to be uniquely different from other environments. It is composed of organisms that are less diverse than those of the outdoors and strongly sourced by, or dependent upon, human bodies. Yet, no one has ever compared the composition of species found in contemporary human homes to that of other structures built by mammals, including those of non-human primates. Here we consider the microbes and arthropods found in chimpanzee beds, relative to the surrounding environment ( n = 41 and 15 beds, respectively). Based on the study of human homes, we hypothesized that the microbes found in chimpanzee beds would be less diverse than those on nearby branches and leaves and that their beds would be primarily composed of body-associated organisms. However, we found that differences between wet and dry seasons and elevation above sea level explained nearly all of the observed variation in microbial diversity and community structure. While we can identify the presence of a chimpanzee based on the assemblage of bacteria, the dominant signal is that of environmental microbes. We found just four ectoparasitic arthropod specimens, none of which appears to be specialized on chimpanzees or their structures. These results suggest that the life to which chimpanzees are exposed while in their beds is predominately the same as that of the surrounding environment.

Madagascar is an island known for its richness of endemic species, including flies. Only eight genera of bee flies (Bombyliidae), including 17 described species (the majority of which are in the subfamily Anthracinae), are known from Madagascar. Here we describe a new species of Bombyliidae from Madagascar, Thevenetimyia spinosavus Maass & Bertone sp. nov. This fly represents the first record of the genus Thevenetimyia from Madagascar and the second species recorded in the Afrotropical Region. A revised checklist of Bombyliidae in Madagascar is provided, along with an appendix of associated literature. The known bee fly species likely represent only a fraction of the true diversity on the island, which has been relatively well sampled through extensive Malaise trapping of flies in Madagascar over the past decade.

We spend most of our lives inside homes, surrounded by arthropods that impact our property as pests and our health as disease vectors and producers of sensitizing allergens. Despite their relevance to human health and well-being, we know relatively little about the arthropods that exist in our homes and the factors structuring their diversity. As previous work has been limited in scale by the costs and time associated with collecting arthropods and the subsequent morphological identification, we used a DNA-based method for investigating the arthropod diversity in homes via high-throughput marker gene sequencing of home dust. Settled dust samples were collected by citizen scientists from both inside and outside more than 700 homes across the United States, yielding the first continental-scale estimates of arthropod diversity associated with our residences. We were able to document food webs and previously unknown geographic distributions of diverse arthropods - from allergen producers to invasive species and nuisance pests. Home characteristics, including the presence of basements, home occupants and surrounding land use, were more useful than climate parameters in predicting arthropod diversity in homes. These noninvasive, scalable tools and resultant findings not only provide the first continental-scale maps of household arthropod diversity, but our analyses also provide valuable baseline information on arthropod allergen exposures and the distributions of invasive pests inside homes.

Tipuloidea, the crane flies, are a diverse lineage of true flies (Insecta: Diptera) whose phylogenetic classification and taxonomy remain a challenge. Here we present the results of a quantitative phylogenetic analysis of Tipuloidea based on combined morphological characters (adult, larvae and pupae) and nuclear gene sequence data (28S rDNA and CAD). Forty-five species, from 44 genera and subgenera, were sampled, representing the four putative families of Tipuloidea (Cylindrotomidae, Limoniidae, Pediciidae and Tipulidae sensu stricto). Analyses of individual datasets, although differing in overall topology, support the monophyly of several major lineages within Tipuloidea. Parsimony and Bayesian analyses using individual morphological and molecular datasets resulted in incongruent topologies. Increased resolution and tree support was obtained when both datasets (morphology and genes) were combined, in both combined evidence parsimony and Bayesian analyses, than when analysed separately. The recovered consensus phylogeny was not consistent with any previously proposed Tipuloidea classification, with previous importance assigned to character states shown here to represent losses and reversals seen as a major factor influencing erroneous classification. The results provided here, together with evidence from previous analyses, were used to append the Tipuloidea classification to supported evolutionary lineages. Tipuloidea is presented as two families: Pediciidae and Tipulidae. Pediciidae is recovered as the sister group to all remaining Tipuloidea. Our current phylogenetic hypothesis is not consistent with the existing subfamilial classification of the ‘Limoniidae’, which is paraphyletic with respect to a well-supported Tipulinae + Cylindrotominae clade, whereas the three ‘limoniid’ subfamilies are para- or polyphyletic. The recognition of ‘Limoniidae’ as a valid monophyletic family is discouraged and the subfamilies of ‘Limoniidae’ are amended and placed within Tipulidae. A revised phylogenetic classification is proposed for the crane flies based on a synthesis of evidence from multiple genes and morphology.

Relationships among families of the lower Diptera (formerly suborder 'Nematocera') have been exceptionally difficult to resolve. Multiple hypotheses based on morphology have been proposed to identify the earliest lineages of flies and place the phylogenetic origin of the higher flies (Brachycera), but convincing support is limited. Here we resolve relationships among the major groups of lower Diptera using sequence data from four nuclear markers, including both ribosomal (28S rDNA) and protein-coding (CAD, TPI and PGD) genes. Our results support both novel and traditional arrangements. Most unexpectedly, the small, highly-specialized family Deuterophlebiidae appears to be sister to all remaining Diptera. Other results include the resolution of the traditional infra-orders Culicomorpha (including a novel superfamily Simulioidea ¼ Thaumaleidae þ Simuliidae), Tipulomorpha (Tipulidae sensu lato þ Trichoceridae) and Bibionomorpha sensu lato. We find support for a limited Psychodomorpha (Blephariceridae, Tanyderidae and Psycho- didae) and Ptychopteromorpha (Ptychopteridae), whereas the placement of several enigmatic families (Nymphomyiidae, Axymyiidae and Perissommatidae) remains ambiguous. According to genetic data, the infra-order Bibionomorpha is sister to the Brachycera. Much of the phylogenetic signal for major lineages was found in the 28S rDNA gene, whereas protein-coding genes performed variably at different levels. In addition to elucidating relationships, we also estimate the age of major lower dipteran clades, based on molecular divergence time estimates using relaxed-clock Bayesian methods and fossil calibration points.

The effect of dung beetle activity on soil nutrition was studied in three distinct soil types under laboratory conditions. Two tunneling dung beetles, Onthophagus gazella (Fabricius) and Onthophagus taurus (Schreber), were allowed to incorporate cattle dung, for brood production, into a piedmont Cecil clay soil, a coastal plain sandy-loam soil and commercially available play sand. Controlled treatments included soil alone and soil exposed to dung only. Soils were tested for primary nutrients (P and K), secondary nutrients (Ca and Mg), and micronutrients (Mn, Zn, and Cu), as well as other soil characteristics (pH, exchangeable acidity, etc.). Both O. gazella and O. taurus produced the most offspring in the piedmont clay soil; variable numbers of brood were produced in other soil types. Soils exposed to dung beetle activity were generally higher in nutrient content than both soils left untreated, and those that had been exposed to cattle dung only. In this manner, tunneling dung beetles can be considered vital to nutrient recycling and plant health in pasture systems.

Species composition and seasonal distribution of dung beetles were studied on dairy and beef cattle pastures in North Carolina. Study sites included a dairy located in the piedmont region (North Carolina Department of Agriculture Piedmont Research Station, Salisbury, NC) and a combined dairy/beef facility in the coastal plain (North Carolina Department of Agriculture Center for Environmental Farming Systems, Goldsboro, NC). Dung beetles were trapped in cattle pastures from March 2002 through September 2003 by using dung-baited pitfall traps. Trapping yielded 4,111 beetles representing 14 species from the piedmont dairy, including Aphodius prodromus Brahm, a new record for North Carolina. Totals of 57,026 beetles representing 28 species and 28,857 beetles representing 26 species were trapped from the dairy unit and beef unit in the coastal plain site, respectively. Onthophagus gazella (F.), a second new record for North Carolina, was collected from the coastal plain. Beetles common to all collection sites include Aphodius erraticus (L.), Aphodius fimetarius (L.), Aphodius granarius (L.), Aphodius pseudolividus Balthasar, Onthophagus taurus Schreber, Onthophagus hecate hecate Panzer, and Onthophagus pennsylvanicus Harold. The introduced beetle O. taurus dominated the dung beetle population, accounting for >50% of the total beetles caught at either site. Beetle activity was greatest from March until November, with activity declining during the winter. Nine exotic species in the genera Onthophagus and Aphodius represented nearly 95% of the beetles trapped.

Hymenoptera, the insect order that includes sawflies, bees, wasps, and ants, exhibits an incredible diversity of phenotypes, with over 145,000 species described in a corpus of textual knowledge since Carolus Linnaeus. In the absence of specialized training, often spanning decades, however, these articles can be challenging to decipher. Much of the vocabulary is domain-specific (e.g., Hymenoptera biology), historically without a comprehensive glossary, and contains much homonymous and synonymous terminology. The Hymenoptera Anatomy Ontology was developed to surmount this challenge and to aid future communication related to hymenopteran anatomy, as well as provide support for domain experts so they may actively benefit from the anatomy ontology development. As part of HAO development, an active learning, dictionary-based, natural language recognition tool was implemented to facilitate Hymenoptera anatomy term discovery in literature. We present this tool, referred to as the 'Proofer', as part of an iterative approach to growing phenotype-relevant ontologies, regardless of domain. The process of ontology development results in a critical mass of terms that is applied as a filter to the source collection of articles in order to reveal term occurrence and biases in natural language species descriptions. Our results indicate that taxonomists use domain-specific terminology that follows taxonomic specialization, particularly at superfamily and family level groupings and that the developed Proofer tool is effective for term discovery, facilitating ontology construction.

In urban ecosystems, socioeconomics contribute to patterns of biodiversity. The ‘luxury effect’, in which wealthier neighbourhoods are more biologically diverse, has been observed for plants, birds, bats and lizards. Here, we used data from a survey of indoor arthropod diversity (defined throughout as family-level richness) from 50 urban houses and found that house size, surrounding vegetation, as well as mean neighbourhood income best predict the number of kinds of arthropods found indoors. Our finding, that homes in wealthier neighbourhoods host higher indoor arthropod diversity (consisting of primarily non-pest species), shows that the luxury effect can extend to the indoor environment. The effect of mean neighbourhood income on indoor arthropod diversity was particularly strong for individual houses that lacked high surrounding vegetation ground cover, suggesting that neighbourhood dynamics can compensate for local choices of homeowners. Our work suggests that the management of neighbourhoods and cities can have effects on biodiversity that can extend from trees and birds all the way to the arthropod life in bedrooms and basements.

Pinned insect specimens stored in museum collections are a fragile and valuable resource for entomological research. As such, they are usually kept away from viewing by the public and hard to access by experts. Here we present a method for mass imaging insect specimens, using GigaPan technology to achieve highly explorable, many-megapixel panoramas of insect museum drawers. We discuss the advantages and limitations of the system, and describe future avenues of collections research using this technology.

A spectacular hypothesis was published recently, which suggested that the "helmet" (a dorsal thoracic sclerite that obscures most of the body) of treehoppers (Insecta: Hemiptera: Membracidae) is connected to the 1st thoracic segment (T1; prothorax) via a jointed articulation and therefore was a true appendage. Furthermore, the "helmet" was interpreted to share multiple characteristics with wings, which in extant pterygote insects are present only on the 2nd (T2) and 3rd (T3) thoracic segments. In this context, the "helmet" could be considered an evolutionary novelty. Although multiple lines of morphological evidence putatively supported the "helmet"-wing homology, the relationship of the "helmet" to other thoracic sclerites and muscles remained unclear. Our observations of exemplar thoraces of 10 hemipteran families reveal multiple misinterpretations relevant to the "helmet"-wing homology hypothesis as originally conceived: 1) the "helmet" actually represents T1 (excluding the fore legs); 2) the "T1 tergum" is actually the anterior dorsal area of T2; 3) the putative articulation between the "helmet" and T1 is actually the articulation between T1 and T2. We conclude that there is no dorsal, articulated appendage on the membracid T1. Although the posterior, flattened, cuticular evagination (PFE) of the membracid T1 does share structural and genetic attributes with wings, the PFE is actually widely distributed across Hemiptera. Hence, the presence of this structure in Membracidae is not an evolutionary novelty for this clade. We discuss this new interpretation of the membracid T1 and the challenges of interpreting and representing morphological data more broadly. We acknowledge that the lack of data standards for morphology is a contributing factor to misinterpreted results and offer an example for how one can reduce ambiguity in morphology by referencing anatomical concepts in published ontologies.

Hymenoptera exhibit an incredible diversity of phenotypes, the result of ~240 million years of evolution and the primary subject of more than 250 years of research. Here we describe the history, development, and utility of the Hymenoptera Anatomy Ontology (HAO) and its associated applications. These resources are designed to facilitate accessible and extensible research on hymenopteran phenotypes. Outreach with the hymenopterist community is of utmost importance to the HAO project, and this paper is a direct response to questions that arised from project workshops. In a concerted attempt to surmount barriers of understanding, especially regarding the format, utility, and development of the HAO, we discuss the roles of homology, “preferred terms”, and “structural equivalency”. We also outline the use of Universal Resource Identifiers (URIs) and posit that they are a key element necessary for increasing the objectivity and repeatability of science that references hymenopteran anatomy. Pragmatically, we detail a mechanism (the “URI table”) by which authors can use URIs to link their published text to the HAO, and we describe an associated tool (the “Analyzer”) to derive these tables. These tools, and others, are available through the HAO Portal website (http://portal.hymao.org). We conclude by discussing the future of the HAO with respect to digital publication, cross-taxon ontology alignment, the advent of semantic phenotypes, and community-based curation.

Flies are one of four superradiations of insects (along with beetles, wasps, and moths) that account for the majority of animal life on Earth. Diptera includes species known for their ubiquity (Musca domestica house fly), their role as pests (Anopheles gambiae malaria mosquito), and their value as model organisms across the biological sciences (Drosophila melanogaster). A resolved phylogeny for flies provides a framework for genomic, developmental, and evolutionary studies by facilitating comparisons across model organisms, yet recent research has suggested that fly relationships have been obscured by multiple episodes of rapid diversification. We provide a phylogenomic estimate of fly relationships based on molecules and morphology from 149 of 157 families, including 30 kb from 14 nuclear loci and complete mitochondrial genomes combined with 371 morphological characters. Multiple analyses show support for traditional groups (Brachycera, Cyclorrhapha, and Schizophora) and corroborate contentious findings, such as the anomalous Deuterophlebiidae as the sister group to all remaining Diptera. Our findings reveal that the closest relatives of the Drosophilidae are highly modified parasites (including the wingless Braulidae) of bees and other insects. Furthermore, we use micro-RNAs to resolve a node with implications for the evolution of embryonic development in Diptera. We demonstrate that flies experienced three episodes of rapid radiation—lower Diptera (220 Ma), lower Brachycera (180 Ma), and Schizophora (65 Ma)—and a number of life history transitions to hematophagy, phytophagy, and parasitism in the history of fly evolution over 260 million y.

Hymenoptera is an extraordinarily diverse lineage, both in terms of species numbers and morphotypes, that includes sawflies, bees, wasps, and ants. These organisms serve critical roles as herbivores, predators, parasitoids, and pollinators, with several species functioning as models for agricultural, behavioral, and genomic research. The collective anatomical knowledge of these insects, however, has been described or referred to by labels derived from numerous, partially overlapping lexicons. The resulting corpus of information--millions of statements about hymenopteran phenotypes--remains inaccessible due to language discrepancies. The Hymenoptera Anatomy Ontology (HAO) was developed to surmount this challenge and to aid future communication related to hymenopteran anatomy. The HAO was built using newly developed interfaces within mx, a Web-based, open source software package, that enables collaborators to simultaneously contribute to an ontology. Over twenty people contributed to the development of this ontology by adding terms, genus differentia, references, images, relationships, and annotations. The database interface returns an Open Biomedical Ontology (OBO) formatted version of the ontology and includes mechanisms for extracting candidate data and for publishing a searchable ontology to the Web. The application tools are subject-agnostic and may be used by others initiating and developing ontologies. The present core HAO data constitute 2,111 concepts, 6,977 terms (labels for concepts), 3,152 relations, 4,361 sensus (links between terms, concepts, and references) and over 6,000 text and graphical annotations. The HAO is rooted with the Common Anatomy Reference Ontology (CARO), in order to facilitate interoperability with and future alignment to other anatomy ontologies, and is available through the OBO Foundry ontology repository and BioPortal. The HAO provides a foundation through which connections between genomic, evolutionary developmental biology, phylogenetic, taxonomic, and morphological research can be actualized. Inherent mechanisms for feedback and content delivery demonstrate the effectiveness of remote, collaborative ontology development and facilitate future refinement of the HAO.

Although humans and arthropods have been living and evolving together for all of our history, we know very little about the arthropods we share our homes with apart from major pest groups. Here we surveyed, for the first time, the complete arthropod fauna of the indoor biome in 50 houses (located in and around Raleigh, North Carolina, USA). We discovered high diversity, with a conservative estimate range of 32–211 morphospecies, and 24–128 distinct arthropod families per house. The majority of this indoor diversity (73%) was made up of true flies (Diptera), spiders (Araneae), beetles (Coleoptera), and wasps and kin (Hymenoptera, especially ants: Formicidae). Much of the arthropod diversity within houses did not consist of synanthropic species, but instead included arthropods that were filtered from the surrounding landscape. As such, common pest species were found less frequently than benign species. Some of the most frequently found arthropods in houses, such as gall midges (Cecidomyiidae) and book lice (Liposcelididae), are unfamiliar to the general public despite their ubiquity. These findings present a new understanding of the diversity, prevalence, and distribution of the arthropods in our daily lives. Considering their impact as household pests, disease vectors, generators of allergens, and facilitators of the indoor microbiome, advancing our knowledge of the ecology and evolution of arthropods in homes has major economic and human health implications.

Background Phylogenetic analyses provide a framework for examining the evolution of morphological and molecular diversity, interpreting patterns in biogeography, and achieving a stable classification. The generic and suprageneric relationships within mosquitoes (Diptera: Culicidae) are poorly resolved, making these subjects difficult to address. Results We carried out maximum parsimony and maximum likelihood, including Bayesian, analyses on a data set consisting of six nuclear genes and 80 morphological characters to assess their ability to resolve relationships among 25 genera. We also estimated divergence times based on sequence data and fossil calibration points, using Bayesian relaxed clock methods. Strong support was recovered for the basal position and monophyly of the subfamily Anophelinae and the tribes Aedini and Sabethini of subfamily Culicinae. Divergence times for major culicid lineages date to the early Cretaceous. Conclusions Deeper relationships within the family remain poorly resolved, suggesting the need for additional taxonomic sampling. Our results support the notion of rapid radiations early in the diversification of mosquitoes.

Background Evolutionary relationships among the 11 extant orders of insects that undergo complete metamorphosis, called Holometabola, remain either unresolved or contentious, but are extremely important as a context for accurate comparative biology of insect model organisms. The most phylogenetically enigmatic holometabolan insects are Strepsiptera or twisted wing parasites, whose evolutionary relationship to any other insect order is unconfirmed. They have been controversially proposed as the closest relatives of the flies, based on rDNA, and a possible homeotic transformation in the common ancestor of both groups that would make the reduced forewings of Strepsiptera homologous to the reduced hindwings of Diptera. Here we present evidence from nucleotide sequences of six single-copy nuclear protein coding genes used to reconstruct phylogenetic relationships and estimate evolutionary divergence times for all holometabolan orders. Results Our results strongly support Hymenoptera as the earliest branching holometabolan lineage, the monophyly of the extant orders, including the fleas, and traditionally recognized groupings of Neuropteroidea and Mecopterida. Most significantly, we find strong support for a close relationship between Coleoptera (beetles) and Strepsiptera, a previously proposed, but analytically controversial relationship. Exploratory analyses reveal that this relationship cannot be explained by long-branch attraction or other systematic biases. Bayesian divergence times analysis, with reference to specific fossil constraints, places the origin of Holometabola in the Carboniferous (355 Ma), a date significantly older than previous paleontological and morphological phylogenetic reconstructions. The origin and diversification of most extant insect orders began in the Triassic, but flourished in the Jurassic, with multiple adaptive radiations producing the astounding diversity of insect species for which these groups are so well known. Conclusion These findings provide the most complete evolutionary framework for future comparative studies on holometabolous model organisms and contribute strong evidence for the resolution of the 'Strepsiptera problem', a long-standing and hotly debated issue in insect phylogenetics.