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Concept: Pterygota


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.

Concepts: Evolution, Nematocera, Insect, Arthropod, Flies, Ant, Spider, Pterygota


The evolution of flight is a key innovation that may enable the extreme diversification of insects. Nonetheless, many species-rich, winged insect groups contain flightless lineages. The loss of flight may promote allopatric differentiation due to limited dispersal power and may result in a high speciation rate in the flightless lineage. Here we show that loss of flight accelerates allopatric speciation using carrion beetles (Coleoptera: Silphidae). We demonstrate that flightless species retain higher genetic differentiation among populations and comprise a higher number of genetically distinct lineages than flight-capable species, and that the speciation rate with the flightless state is twice that with the flight-capable state. Moreover, a meta-analysis of 51 beetle species from 15 families reveals higher genetic differentiation among populations in flightless compared with flight-capable species. In beetles, which represent almost one-fourth of all described species, repeated evolution of flightlessness may have contributed to their steady diversification since the Mesozoic era.

Concepts: Evolution, Biology, Insect, Beetle, Speciation, Insects, Pterygota, Endopterygota


The cross-pollination of most alpine plants depends on insects, whose altitudinal distribution is limited by temperature. However, although global warming is causing shifts in temporal and spatial species distribution, we are still largely unaware of how plant-pollinator interactions change with elevation and time along altitudinal gradients. This makes the detection of endangered interactions and species challenging. In this study, we aimed at providing such a reference, and tested if and how the major flower-visiting insect orders and families segregated by altitude, phenology and foraging preferences along an elevational gradient from 970 m to 2700 m in the Alps. Flies were the main potential pollinators from 1500 m, as bees and beetles decreased rapidly above that limit. Diptera, Coleoptera and Hymenoptera differed significantly in the angiosperm assemblages visited. Within Diptera, the predominant group, major families segregated by both phenology and foraging preferences along the gradient. Empidids, muscids and anthomyiids, whose role in pollination has never been investigated, dominated the upper part of the gradient. Our results thus suggest that flies and the peculiar plants they visit might be particularly at risk under global warming, and highlight the blatant lack of studies about critical components of these rich, yet fragile mountain ecosystems.

Concepts: Insect, Larva, Ecology, Pollination, Insects, Pterygota, Endopterygota, Neoptera


During the mid-Cretaceous, angiosperms diversified from several nondiverse lineages to their current global domination [1], replacing earlier gymnosperm lineages [2]. Several hypotheses explain this extensive radiation [3], one of which involves proliferation of insect pollinator associations in the transition from gymnosperm to angiosperm dominance. However, most evidence supports gymnosperm-insect pollinator associations, buttressed by direct evidence of pollen on insect bodies, currently established for four groups: Thysanoptera (thrips), Neuroptera (lacewings), Diptera (flies), and now Coleoptera (beetles). Each group represents a distinctive pollination mode linked to a unique mouthpart type and feeding guild [4-9]. Extensive indirect evidence, based on specialized head and mouthpart morphology, is present for one of these pollinator types, the long-proboscid pollination mode [10], representing minimally ten family-level lineages of Neuroptera, Mecoptera (scorpionflies), and Diptera [8, 10, 11]. A recurring feature uniting these pollinator modes is host associations with ginkgoalean, cycad, conifer, and bennettitalean gymnosperms. Pollinator lineages bearing these pollination modes were categorized into four evolutionary cohorts during the 35-million-year-long angiosperm radiation, each defined by its host-plant associations (gymnosperm or angiosperm) and evolutionary pattern (extinction, continuation, or origination) during this interval [12]. Here, we provide the first direct evidence for one cohort, exemplified by the beetle Darwinylus marcosi, family Oedemeridae (false blister beetles), that had an earlier gymnosperm (most likely cycad) host association, later transitioning onto angiosperms [13]. This association constitutes one of four patterns explaining the plateau of family-level plant lineages generally and pollinating insects specifically during the mid-Cretaceous angiosperm radiation [12].

Concepts: Insect, Plant, Pollination, Pollen, Insects, Pterygota, Gymnosperm, Endopterygota


The red flour beetle Tribolium castaneum is an emerging insect model organism representing the largest insect order, Coleoptera, which encompasses several serious agricultural and forest pests. Despite the ecological and economic importance of beetles, most insect olfaction studies have so far focused on dipteran, lepidopteran, or hymenopteran systems.

Concepts: Biology, Insect, Larva, Antenna, Insects, Pterygota, Tenebrionidae, Endopterygota


Explaining the taxonomic richness of the insects, comprising over half of all described species, is a major challenge in evolutionary biology. Previously, several evolutionary novelties (key innovations) have been posited to contribute to that richness, including the insect bauplan, wings, wing folding and complete metamorphosis, but evidence over their relative importance and modes of action is sparse and equivocal. Here, a new dataset on the first and last occurrences of fossil hexapod (insects and close relatives) families is used to show that basal families of winged insects (Palaeoptera, e.g. dragonflies) show higher origination and extinction rates in the fossil record than basal wingless groups (Apterygota, e.g. silverfish). Origination and extinction rates were maintained at levels similar to Palaeoptera in the more derived Polyneoptera (e.g. cockroaches) and Paraneoptera (e.g. true bugs), but extinction rates subsequently reduced in the very rich group of insects with complete metamorphosis (Holometabola, e.g. beetles). Holometabola show evidence of a recent slow-down in their high net diversification rate, whereas other winged taxa continue to diversify at constant but low rates. These data suggest that wings and complete metamorphosis have had the most effect on family-level insect macroevolution, and point to specific mechanisms by which they have influenced insect diversity through time.

Concepts: Evolution, Insect, Charles Darwin, Insects, Pterygota, Insect wing, Endopterygota, Pupa


Insects exhibit a wide diversity of anatomical specializations in their adult and immature stages associated with particular aspects of their biology. The order Neuroptera (lacewings, antlions, and their relatives) are a moderately diverse lineage of principally predatory animals, at least in their immature stages, as all have a modified piercing-sucking mandible-maxillary complex that allows them to drain fluids from their prey. As such, the larvae of various groups have evolved unique anatomical and behavioral specializations for approaching and subduing their prey, particularly the green lacewings (Chrysopidae), where immatures are also adept at camouflage [1-4]. Here we report the discovery of a unique mode of life among mid-Cretaceous mesochrysopids, an early stem group to modern green lacewings [5-7] exhibiting a combination of morphological modifications in both adults and larvae unknown among living and fossil Neuroptera, even across winged insects. The new mesochrysopids exhibit a uniquely prolonged thorax, elongate legs, and dramatically reduced hind wings in adults, and larvae have extremely elongate, slender legs with pectinate pretarsal claws and lacking trumpet-shaped empodia. The peculiarities of the larvae include features principally found in spider-associated insect groups, implying that these lacewings were early specialists on web-spinning spiders, either as active predators or kleptoparasites. This reveals a dramatic and ancient degree of ecological refinement in a major lineage of insect predators, for a food resource otherwise not utilized by most lacewings.

Concepts: Evolution, Insect, Ecology, Biological pest control, Pterygota, Neuroptera, Chrysopidae, Hemerobiidae


The position of the Zoraptera remains one of the most challenging and uncertain concerns in ordinal-level phylogenies of the insects. Zoraptera have been viewed as having a close relationship with five different groups of Polyneoptera, or as being allied to the Paraneoptera or even Holometabola. Although rDNAs have been widely used in phylogenetic studies of insects, the application of the complete 28S rDNA are still scattered in only a few orders. In this study, a secondary structure model of the complete 28S rRNAs of insects was reconstructed based on all orders of Insecta. It was found that one length-variable region, D3-4, is particularly distinctive. The length and/or sequence of D3-4 is conservative within each order of Polyneoptera, but it can be divided into two types between the different orders of the supercohort, of which the enigmatic order Zoraptera and Dictyoptera share one type, while the remaining orders of Polyneoptera share the other. Additionally, independent evidence from phylogenetic results support the clade (Zoraptera+Dictyoptera) as well. Thus, the similarity of D3-4 between Zoraptera and Dictyoptera can serve as potentially valuable autapomorphy or synapomorphy in phylogeny reconstruction. The clades of (Plecoptera+Dermaptera) and ((Grylloblattodea+Mantophasmatodea)+(Embiodea+Phasmatodea)) were also recovered in the phylogenetic study. In addition, considering the other studies based on rDNAs, this study reached the highest congruence with previous phylogenetic studies of Holometabola based on nuclear protein coding genes or morphology characters. Future comparative studies of secondary structures across deep divergences and additional taxa are likely to reveal conserved patterns, structures and motifs that can provide support for major phylogenetic lineages.

Concepts: Biology, Insect, Phylogenetic nomenclature, Phylogenetics, Cladistics, Insects, Pterygota, Neoptera


Many extant insects have developed pad structure, euplantulae or arolia on their tarsi to increase friction or enhance adhesion for better mobility. Many polyneopteran insects with euplantulae, e.g., Grylloblattodea, Mantophasmatodea and Orthoptera, have been described from the Mesozoic. However, the origin and evolution of stick insect’s euplantulae are poorly understood due to rare fossil records. Here, we report the earliest fossil records of Timematodea hitherto, Tumefactipes prolongates gen. et sp. nov. and Granosicorpes lirates gen. et sp. nov., based on three specimens from the mid-Cretaceous Burmese amber. Specimens of Tumefactipes prolongates gen. et sp. nov. have extremely specialized and expanded euplantulae on their tarsomere II. These new findings are the first known and the earliest fossil records about euplantula structure within Phasmatodea, demonstrating the diversity of euplantulae in Polyneoptera during the Mesozoic. Such tarsal pads might have increased friction and helped these mid-Cretaceous stick insects to climb more firmly on various surfaces, such as broad leaves, wetted tree branches or ground. These specimens provide more morphological data for us to understand the relationships of Timematodea, Euphasmatodea, Orthoptera and Embioptera, suggesting that Timematodea might be monophyletic with Euphasmatodea rather than Embioptera and Phasmatodea should have a closer relationship with Orthoptera rather than Embioptera. This article is protected by copyright. All rights reserved.

Concepts: Insect, Phasmatodea, Fossil, Copyright, Insects, Pterygota, Neoptera, Exopterygota


Agroforestry systems are environment-friendly production systems which help to preserve biodiversity while providing people with a way of earning a living. Cacao is a historically important crop in Venezuela that traditionally has been produced in agroforestry systems. However, few studies have evaluated how different trees used in those systems affect the dynamics and abundance of insects. The present study evaluated the entomofauna assemblages associated with different combinations of four timber-yielding trees and four Criollo cacao cultivars established in a lowland tropical ecosystem in Venezuela. A randomized block design with two replicates was used, each block having 16 plots which included all 16 possible combinations of four native timber trees (Cordia thaisiana, Cedrela odorata, Swietenia macrophylla, and Tabebuia rosea) and four Criollo cacao cultivars (Porcelana, Guasare, Lobatera and Criollo Merideño). Insects were collected with yellow pan traps and sorted to order. Coleoptera and parasitoid Hymenoptera were determined to the family level. In total, 49,538 individuals of seven orders were collected, with Hymenoptera, Diptera, and Hemiptera being the most abundant, although only Lepidoptera and Coleoptera abundances were significantly influenced by the timber tree species. Twenty-three families of parasitoid Hymenoptera and 26 of Coleoptera were found. Significant differences in insects’ assemblages were found both in parasitoid Hymenoptera and Coleoptera families associated to every shade tree, with the families Eulophidae and Lycidae being indicators for Cordia, and Chalcididae for Swietenia. The entomofauna relationship with the cacao cultivar was barely significant, although Scydmaenidae and Scarabaeidae were indicators for Lobatera and Merideño, respectively. No significant effects were found for interaction with cacao cultivars and native trees. We concluded that the particular insect assemblages found in Cedrela odorata and Cordia thaisiana, together with their high growing rates, make these two species an optimal choice for cacao agroforestry systems.

Concepts: Insect, Forestry, Beetle, Hymenoptera, Apocrita, Insects, Pterygota, Endopterygota