The great evolutionary success of angiosperms has traditionally been explained, in part, by the partnership of these plants with insect pollinators [1-6]. The main approach to understanding the origins of this pervasive relationship has been study of the pollinators of living cycads, gnetaleans, and basal angiosperms . Among the most morphologically specialized living pollinators are diverse, long-proboscid flies. Early such flies include the brachyceran family Zhangsolvidae, previously known only as compression fossils from the Early Cretaceous of China and Brazil. It belongs to the infraorder Stratiomyomorpha, a group that includes the flower-visiting families Xylomyidae and Stratiomyidae. New zhangsolvid specimens in amber from Spain (ca. 105 mega-annum [Ma]) and Myanmar (100 Ma) reveal a detailed proboscis structure adapted to nectivory. Pollen clumped on a specimen from Spain is Exesipollenites, attributed to a Mesozoic gymnosperm, most likely the Bennettitales. Late Mesozoic scorpionflies with a long proboscis have been proposed as specialized pollinators of various extinct gymnosperms, but pollen has never been observed on or in their bodies . The new discovery is a very rare co-occurrence of pollen with its insect vector and provides substantiating evidence that other long-proboscid Mesozoic insects were gymnosperm pollinators. Evidence is thus now gathering that visitors and probable pollinators of early anthophytes, or seed plants, involved some insects with highly specialized morphological adaptations, which has consequences for interpreting the reproductive modes of Mesozoic gymnosperms and the significance of insect pollination in angiosperm success.
The Chironomidae (Insecta: Diptera) type collection at the Laboratory of Ecology of Aquatic Insects (LEIA - UFSCar) is reviewed. It comprises 103 primary types, as well as 95 paratypes, mostly resulting from research by S. Trivinho-Strixino and G. Strixino. Notes updating the taxonomic status are provided for some species.
Chironomidae (Diptera) are among the most diverse and widespread aquatic insects, with roughly 5,500 described species. However, prior to the present work, no species of Chironomidae had been documented from the island of Hispaniola. Collections of non-biting midges, with emphasis on the lotic fauna, were made in the Dominican Republic during July of 2015. In total, 578 specimens belonging to 27 genera and at least 44 species within the subfamilies Chironominae (20 taxa), Orthocladiinae (16 taxa) and Tanypodinae (8 taxa) were found. The subfamilies Chironominae and Orthocladiinae predominated. Polypedilum was the most widespread and diverse genus of Chironominae. Metriocnemus were collected in bromeliad tanks. The chironomid fauna in Dominican Republic includes multiple genera with worldwide distributions, including Holarctic and Neotropical components.
Here we present a phylogeny of beetles (Insecta: Coleoptera) based on DNA sequence data from eight nuclear genes, including six single-copy nuclear protein-coding genes, for 367 species representing 172 of 183 extant families. Our results refine existing knowledge of relationships among major groups of bee- tles. Strepsiptera was confirmed as sister to Coleoptera and each of the suborders of Coleoptera was recovered as monophyletic. Interrelationships among the subor- ders, namely Polyphaga (Adephaga (Archostemata, Myxophaga)), in our study differ from previous studies. Adephaga comprised two clades corresponding to Hydrade- phaga and Geadephaga. The series and superfamilies of Polyphaga were mostly monophyletic. The traditional Cucujoidea were recovered in three distantly related clades. Lymexyloidea was recovered within Tenebrionoidea. Several of the series and superfamilies of Polyphaga received moderate to maximal clade support in most analyses, for example Buprestoidea, Chrysomeloidea, Coccinelloidea, Cucujiformia, Curculionoidea, Dascilloidea, Elateroidea, Histeroidea and Hydrophiloidea. However, 836 D. D. McKenna et al. many of the relationships within Polyphaga lacked compatible resolution under maximum-likelihood and Bayesian inference, and/or lacked consistently strong nodal support. Overall, we recovered slightly younger estimated divergence times than pre- vious studies for most groups of beetles. The ordinal split between Coleoptera and Strepsiptera was estimated to have occurred in the Early Permian. Crown Coleoptera appeared in the Late Permian, and only one or two lineages survived the end-Permian mass extinction, with stem group representatives of all four suborders appearing by the end of the Triassic. The basal split in Polyphaga was estimated to have occurred in the Triassic, with the stem groups of most series and superfamilies originating during the Triassic or Jurassic. Most extant families of beetles were estimated to have Creta- ceous origins. Overall, Coleoptera experienced an increase in diversification rate com- pared to the rest of Neuropteroidea. Furthermore, 10 family-level clades, all in suborder Polyphaga, were identified as having experienced significant increases in diversification rate.These include most beetle species with phytophagous habits, but also several groups not typically or primarily associated with plants. Most of these groups originated in the Cretaceous, which is also when a majority of the most species-rich beetle families first appeared. An additional 12 clades showed evidence for significant decreases in diversi- fication rate. These clades are species-poor in theModern fauna, but collectively exhibit diverse trophic habits. The apparent success of beetles, as measured by species numbers, may result from their associations with widespread and diverse substrates – especially plants, but also including fungi, wood and leaf litter – but what facilitated these associ- ations in the first place or has allowed these associations to flourish likely varies within and between lineages.Our results provide a uniquelywell-resolved temporal and phylo- genetic framework for studying patterns of innovation and diversification in Coleoptera, and a foundation for further sampling and resolution of the beetle tree of life.
Stag beetles (family Lucanidae Latreille, 1804) are one of the earliest branching lineages of scarab beetles that are characterized by the striking development of the male mandibles. Despite stag beetles’ popularity among traditional taxonomists and amateur collectors, there has been almost no study of lucanid relationships and evolution. Entomologists, including Jeannel (1942), have long recognized resemblance between the austral stag beetles of the tribes Chiasognathini, Colophonini, Lamprimini, Pholidotini, Rhyssonotini, and Streptocerini, but this hypothesis of their close relationship across the continents has never been tested. To gain further insight into lucanid phylogeny and biogeography, we reconstructed the first molecular phylogeny of world stag beetles using DNA sequences from mitochondrial 16S rDNA, nuclear 18S and 28S rDNA, and the nuclear protein-coding (NPC) gene wingless for 93 lucanid species representing all extant subfamilies and 24 out of the 27 tribes, together with 14 representative samples of other early branching scarabaeoid families and two staphyliniform beetle families as outgroups. Both Bayesian inference (BI) and maximum likelihood inference (MLI) strongly supported the monophyly of Lucanidae sensu lato that includes Diphyllostomatidae. Within Lucanidae sensu stricto, the subfamilies Lucaninae and Lampriminae appeared monophyletic under both methods of phylogenetic inferences; however, Aesalinae and Syndesinae were found to be polyphyletic. A time-calibrated phylogeny based on five fossil data estimated the origin of crown group Lucanidae as circa 160 million years ago (MYA). Divergence between the Neotropical and Australasian groups of the Chiasognathini was estimated to be circa 47 MYA, with the South African Colophonini branching off from the ancient Chiasognathini lineage around 87 MYA. Another Gondwanan relationship was recovered between the Australasian Eucarteria and the Neotropical Casignetus, which diverged circa 58 MYA. Lastly, as Jeannel’s hypothesis predicted, divergence within Lampriminae between the Australasian Lamprima and the Neotropical Streptocerus was estimated to be circa 37 MYA. The split of these lineages were generally concordant with the pattern of continental break-up of the super-continent Gondwana, and our biogeographic reconstructions based on the dispersal-extinction-cladogenesis model (DEC) corroborate our view that the divergences in these austral lineages were caused by vicariance events following the Gondwanan break-up. In addition, the phylogenetic position and geographic origin of the Hawaiian genus Apterocyclus was revealed for the first time. Overall, our results provide the framework toward studying lucanid relationships and divergence time estimates, which allowed for more accurate biogeographic explanations and discussions on ancestral lucanids and the evolutionary origin of the enlarged male mandibles.
The family Buprestidae (jewel beetles or metallic wood-boring beetles), contains nearly 15000 species in 522 genera. Together with the small family Schizopodidae (seven species, three genera), they form the superfamily Buprestoidea. Adult Buprestoidea feed on flowers or foliage, whereas larvae are mostly internal feeders, boring in roots or stems, or mining the leaves of woody or herbaceous plants. The subfamilial and tribal classification of Buprestoidea remains unsettled, with substantially different schemes proposed by different workers based on morphology. Here we report the first large-scale molecular phylogenetic study of the superfamily Buprestoidea based on data from four genes for 141 ingroup species. We used these data to reconstruct higher-level relationships and to assess the current classification and the origins of the larval leaf-mining habit within Buprestoidea. In our analyses, the monophyly of Buprestoidea was strongly supported, as was the monophyly of Schizopodidae and its placement sister to Buprestidae. Our results are largely consistent with the generally accepted major lineages of buprestoids, including clearly-defined agrilines, buprestines-chrysochroines and early-branching julodines-polycestines. In addition to Schizopodidae, three of the six subfamilies were monophyletic in our study: Agrilinae, Julodinae and the monogeneric Galbellinae (Galbella). Polycestinae was monophyletic with the exception of the enigmatic Haplostethini. Chrysochroinae and Buprestinae were not monophyletic, but were recovered together in a large mixed clade along with Galbella. The interrelationships of Chrysochroinae and Buprestinae were not well resolved; however they were clearly polyphyletic, with chrysochroine genera falling into several different well-supported clades otherwise comprising buprestine genera. All Agrilinae were contained in a single strongly supported clade. Coraebini were dispersed throughout Agrilinae, with strong nodal support for several clades representing subtribes. Neither Agrilini nor Tracheini were monophyletic. The leaf-mining genus Paratrachys (Paratracheini) was recovered within the Acmaeoderioid clade, consistent with the current classification, and confirming the independent origins of leaf-mining within Polycestinae and Agrilinae. Additionally, our results strongly suggest that the leaf-mining agriline tribe Tracheini is polyphyletic, as are several of its constituent subtribes. External root feeding was likely the ancestral larval feeding habit in Buprestoidea. The apparent evolutionary transitions to internal feeding allowed access to a variety of additional plant tissues, including leaves. Interestingly, the several genera of leaf-mining agrilines do not form a monophyletic group. Many of these genera are diverse and highly specialized, possibly indicating adaptive radiations.
The beetle series Staphyliniformia exhibits extraordinary taxonomic, ecological and morphological diversity. To gain further insight into staphyliniform relationships and evolution, we reconstructed the phylogeny of Staphyliniformia using DNA sequences from nuclear 28S rDNA and the nuclear protein-coding gene CAD for 282 species representing all living families and most subfamilies, a representative sample of Scarabaeiformia serving as a near outgroup, and three additional beetles as more distant outgroups. Under both Bayesian inference (BI) and maximum likelihood inference (MLI), the major taxa within Staphyliniformia are each monophyletic: (i) Staphylinoidea, (ii) Hydrophiloidea s.l., and the contained superfamilies (iii) Hydrophiloidea s.s. and (iv) Histeroidea, although Staphylinoidea and Hydrophiloidea s.l. are not strongly supported by MLI bootstrap. Scarabaeiformia is monophyletic under both methods of phylogenetic inference. However, the relative relationships of Staphylinoidea, Hydrophiloidea s.l. and Scarabaeiformia differ between BI and MLI: under BI, Staphyliniformia and Scarabaeiformia were sister groups; under MLI, Hydrophiloidea s.l. and Scarabaeiformia were sister groups and these together were sister to Staphylinoidea. The internal relationships in Scarabaeiformia were similar under both methods of phylogenetic inference, with Cetoniinae, Dynastinae+Rutelinae, Hybosoridae, Passalidae, Scarabaeidae and Scarabaeinae recovered as monophyla. Histeridae comprised two major clades: (1) Abraeinae, Trypanaeine and Trypeticinae; and (2) Chlamydopsinae, Dendrophilinae, Haeteriinae, Histerinae, Onthophilinae, Saprininae and Tribalinae. The relationships among early-divergent Hydrophiloidea differed between BI and MLI, and overall were unresolved or received only moderate to low nodal support. The staphylinoid families Agyrtidae, Hydraenidae and Ptiliidae were recovered as monophyletic; the latter two were sister taxa, and Staphylinidae+Silphidae was also monophyletic. Silphidae was placed within Staphylinidae in close relation to a subset of Tachyporinae. Pselaphinae and Scydmaeninae were both recovered within Staphylinidae, in accordance with recent analyses of morphological characters, although not always with recently proposed sister taxa. None of the four major groups of Staphylinidae proposed by Lawrence and Newton (1982) was recovered as monophyletic. Certain highly specialized staphyliniform habits and morphologies, such as abdominal defensive glands and reduced elytra, have arisen in parallel in separate lineages. Further, our analyses support two major transitions to an aquatic lifestyle within Staphyliniformia: once within Staphylinoidea (Hydraenidae), and once within Hydrophiloidea s.l. (Hydrophiloidea s.s.). On a smaller scale, the most common transition is from litter to subcortical or to periaquatic microhabitats and the next most common is from litter to carrion and to fungi. Overall, transitions to periaquatic microhabitats were the most numerous. The broad picture in Staphyliniformia seems to be a high level of evolutionary plasticity, with multiple possible pathways to and from many microhabitat associations, and litter as a major source microhabitat for diversification. In Scarabaeiformia, the most common transitions were from litter to foliage, with flowers to litter, litter to flowers, and litter to dung being next, and then litter to roots, logs or carrion. Litter is again the largest overall source microhabitat. The most common transitions were to foliage and flowers.It thus seems that the litter environment presents ecological and evolutionary opportunities/challenges that facilitate entry of Staphyliniformia and Scarabaeiformia into new' and different ecological adaptive zones.
Weevils are one of the largest groups of living organisms, with more than 60,000 species feeding mostly on plants. With only one exception, their described larvae are typical plant-feeders, with mouthparts adapted to chewing plant material. Here we describe the second case of a weevil with early-instar larvae adapted to killing conspecifics. We have studied the life history of Anchylorhynchus eriospathae G. G. Bondar, 1943 (Curculioninae: Derelomini sensu Caldara, Franz & Oberprieler (2014)), a species whose immatures feed internally on palm flowers and fruits. We provide detailed descriptions of all immature stages, including the extremely modified first-instar larva. Unlike other weevils and later instars, this stage exhibits a flat body with very long ventropedal lobe setae, a large and prognathous head with a gula, and falciform mandibles, each with a serrate retinaculum, that are used to fight with and eventually kill other first-instar larvae. We also provide biological notes on all stages and the results of behavioral tests that showed that larval aggression occurs only among early life stages. Finally we show that adult size is highly dependent on timing of oviposition. This specialized killer first instar probably evolved independently from the one other case known in weevils, in Revena rubiginosa (Conoderinae: Bariditae sensu Prena, Colonnelli & Hespenheide (2014)). Interestingly, both lineages inhabit the same hosts, raising the possibility that both intra-and inter-specific competition shaped those phenotypes. Given the scarcity of knowledge on early larval stages of concealed insect herbivores, it is possible that our findings represent an instance of a much broader phenomenon. Our observations also allowed us to conclude that Anchylorhynchus eriospathae and A. hatschbachi G. G. Bondar, 1943 are actually the same species, which we synonymize here by considering the latter as a junior synonym(new synonymy).
The Boston Harbor Islands Partnership (BHIP) and the Museum of Comparative Zoology (MCZ) at Harvard University collaborated to conduct the first phase of an All Taxa Biodiversity Inventory (ATBI) in Boston Harbor Islands national park area, Boston, Massachusetts. This six-year phase of the ATBI focused on the “microwilderness” of the park, a vast realm of invertebrate animals. The main objectives of this phase of the ATBI were threefold: (1) to document the diversity and distribution of arthropod and mollusk taxa in the park; (2) to engage and educate students and the public about local invertebrate biodiversity; (3) to use biodiversity data to inform park management. The project relied on the participation of more than fifty students, interns, citizen scientists, and volunteers to collect invertebrates on nineteen islands, and to sort, prepare, image, and database more than 75,000 specimens in the lab. Additionally, more than forty taxonomists from North America and Europe donated their time and expertise to identify 1,732 species and morphospecies. All specimens are permanently housed in the collections at the MCZ. Exploratory analyses of selected taxa suggested that, as predicted by the theory of island biogeography (MacArthur and Wilson 1967), island size was a reasonable predictor of species richness. However, an island’s distance from the mainland was correlated with species richness for only two of six taxa. Additional factors such as habitat diversity and habitat type (i.e., presence of freshwater or mature forests), as well as human-aided transport of species, appeared to be significant drivers of invertebrate distribution patterns across the islands. Numbers of introduced species were proportionally higher on the islands than on the coastal mainland for several beetle families, but comparable data for the mainland were lacking for most taxa, pointing to the lack of basic species distribution data that exist for invertebrates even in temperate, accessible locales. A large proportion of introduced species also had extremely high population counts and widespread distributions across islands, relative to native species.
Anchylorhynchus vanini sp. nov. from the Amazon is described, including the mouthparts and male genitalia. The new species is compared with similar species within the genus and the key to the species of Anchylorhynchus provided by Vaurie (1954) is modified to include the new species. Adult specimens were collected in flowers of the palm Syagrus vermicularis Noblick and additional collections in other palms species suggest that this association is specific. This is the first record of the palm Syagrus vermicularis as host for a species of Anchylorhynchus.
Sicoderus robini sp. nov. (type locality: Bolivia, Santa Cruz de la Sierra, Buena Vista, El Cairo) is described and illustrated. The new species is assigned to the Sicoderus appendiculatus species group, compared with similar species of the group and with the two other species of Sicoderus that occur in Bolivia. The previously published key for species identification of the S. appendiculatus group is updated to include the new species.
Three new species of the genus Anchylorhynchus from Colombia, are described: Anchylorhynchus pinocchio sp. nov., A. centrosquamatus sp. nov. and A. luteobrunneus sp. nov.. A morphological description, including the male genitalia, is provided for each species as well as a comparison with similar species within the genus. All three species are found in inflorescences of species of Syagrus Mart. (Arecaceae). The adults are pollinators and the larvae develop inside fruits and feed on the endosperm, interrupting seed formation and causing fruit abortion.
Light pollution due to exterior lighting is a rising concern. While glare, light trespass and general light pollution have been well described, there are few reported studies on the impact of light pollution on insects. By studying insect behavior in relation to artificial lighting, we suggest that control of the UV component of artificial lighting can significantly reduce its attractiveness, offering a strong ability to control the impact on insects. Traditionally, the attractiveness of a lamp to insects is calculated using the luminous efficiency spectrum of insect rhodopsin. This has enabled the development of lamps that emit radiation with wavelengths that are less visible to insects (that is, yellow lamps). We tested the assumption that the degree of visibility of a lamp to insects can predict its attractiveness by means of experimental collections. We found that the expected lamp's visibility is indeed related to the extent to which it attracts insects. However, the number of insects attracted to a lamp is disproportionally affected by the emission of ultraviolet radiation. UV triggers the behavior of approaching lights more or less independently of the amount of UV radiation emitted. Thus, even small amounts of UV should be controlled in order to develop bug-free lamps.
As part of an All Taxa Biodiversity Inventory in Boston Harbor Islands national park area, an inventory of carabid beetles on 13 islands was conducted. Intensive sampling on ten of the islands, using an assortment of passive traps and limited hand collecting, resulted in the capture of 6,194 specimens, comprising 128 species. Among these species were seven new state records for Massachusetts (Acupalpus nanellus, Amara aulica, Amara bifrons, Apenes lucidulus, Bradycellus tantillus, Harpalus rubripes and Laemostenus tern cola terricola-the last also a new country record; in passing we report also new state records for Happalus rubripes from New York and Pennsylvania, Amara ovata from Pennsylvania, and the first mainland New York records for Asaphidion curtum). For most islands, there was a clear relationship between species richness and island area. Two islands, however, Calf and Grape, had far more species than their relatively small size would predict. Freshwater marshes on these islands, along with a suite of hygrophilous species, suggested that habitat diversity plays an important role in island species richness. Introduced species (18) comprised 14.0% of the total observed species richness, compared to 5.5% (17 out of 306 species) documented for Rhode Island. We surmise that the higher proportion of introduced species on the islands is, in part, due to a higher proportion of disturbed and open habitats as well as high rates of human traffic. We predict that more active sampling in specialized habitats would bring the total carabid fauna of the Boston Harbor Islands closer to that of Rhode Island or eastern Massachusetts in richness and composition; however, isolation, human disturbance and traffic, and limited habitat diversity all contribute to reducing the species pool on the islands relative to that on the mainland.
Many studies have examined how island biogeography affects diversity on the scale of island systems. In this study, we address how diversity varies over very short periods of time on individual islands. To do this, we compile an inventory of the ants living in the Boston Harbor Islands National Recreation Area, Boston, Massachusetts, USA using data from a five-year All Taxa Biodiversity Inventory of the region's arthropods. Consistent with the classical theory of island biogeography, species richness increased with island size, decreased with island isolation, and remained relatively constant over time. Additionally, our inventory finds that almost half of the known Massachusetts ant fauna can be collected in the BHI, and identifies four new species records for Massachusetts, including one new to the United States, Myrmica scabrinodis. We find that the number of species actually active on islands depended greatly on the timescale under consideration. The species that could be detected during any given week of sampling could by no means account for total island species richness, even when correcting for sampling effort. Though we consistently collected the same number of species over any given week of sampling, the identities of those species varied greatly between weeks. This variation does not result from local immigration and extinction of species, nor from seasonally-driven changes in the abundance of individual species, but rather from weekly changes in the distribution and activity of foraging ants. This variation can be upwards of 50% of ant species per week. This suggests that numerous ant species on the BHI share the same physical space at different times. This temporal partitioning could well explain such unexpectedly high ant diversity in an isolated, urban site.
The ability to predict patterns of species dispersal across habitat edges takes on increasing conservation relevance as landscapes become more fragmented. We assessed edge responses for four ground-dwelling arthropod taxa by measuring their distribution and movement patterns across a forest edge associated with a forested riparian buffer and an adjacent upslope clearcut at a single site. We used Spatial Analysis by Distance Indices to describe the spatial distribution of arthropods, habitat variables, and the associations between them across a 49 x 63 m pitfall trapping grid with 80 trapping stations. We used mark-release-recapture techniques to measure the distance, rate, and direction of arthropod movement within and between habitat types. Scaphinotus angusticollis (Coleoptera; Carabidae) was restricted to the cool, moist portion of the riparian buffer and was not observed to cross the forest edge. The carabids Pterostichus lattini and Scaphinotus marginatus had more eurytopic distributions across all habitats. Higher overall mobility for S. marginauts may explain, in part, the greater frequency of movement for this species across the forest/clearcut edge. Lycosid spiders, strongly associated with the clearcut, also moved into one portion of the riparian buffer that was warmer and drier. They had a relatively high frequency of movement across the clearcut/forest boundary. These results suggest that ground-dwelling arthropod taxa are likely to respond differently to habitat edges, and that their habitat affinities and mobility strongly influence movement patterns across the landscape. Understanding these patterns of distribution and movement will aid the sustainable management of forest and riparian arthropod taxa in fragmented landscapes.
Bembidion (Lymnaeum) nigropiceum (Marsham) (=puritanum Hayward), a European species introduced into Massachusetts but presumed not to have become established, has been rediscovered during the Boston Harbor Islands All Taxa Biodiversity Inventory undertaken by the Museum of Comparative Zoology and the National Park Service. A summary is presented of treatment of this species in North America. Data on specimens collected are presented, along with observations on habitat and biology. Some speculations are presented about its highly specialized habitat in the gravel pushed up by high tide, which may act as a food-trapping sieve. A few words are included about future actions needed to resolve questions of distribution and behavior.
Background: The extraordinary morphology, reproductive and developmental biology, and behavioral ecology of twisted wing parasites (order Strepsiptera) have puzzled biologists for centuries. Even today, the phylogenetic position of these enigmatic "insects from outer space''  remains uncertain and contentious. Recent authors have argued for the placement of Strepsiptera within or as a close relative of beetles (order Coleoptera), as sister group of flies (order Diptera), or even outside of Holometabola.Methodology/Principal Findings: Here, we combine data from several recent studies with new data (for a total of 9 nuclear genes and similar to 13 kb of aligned data for 34 taxa), to help clarify the phylogenetic placement of Strepsiptera. Our results unequivocally support the monophyly of Neuropteroidea (= Neuropterida + Coleoptera) + Strepsiptera, but recover Strepsiptera either derived from within polyphagan beetles (order Coleoptera), or in a position sister to Neuropterida. All other supra-ordinal- and ordinal-level relationships recovered with strong nodal support were consistent with most other recent studies.Conclusions/Significance: These results, coupled with the recent proposed placement of Strepsiptera sister to Coleoptera, suggest that while the phylogenetic neighborhood of Strepsiptera has been identified, unequivocal placement to a specific branch within Neuropteroidea will require additional study.