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The humble and industrious dung beetle is a marvelous beast: the 6,000 species identified so far are intricately entwined with human history and scientific endeavor. these night-soil collectors of the planet have been worshiped as gods, worn as jewelery, and painted by artists. More practically, they saved Hawaii from ecological blight, and rescued Australia from plagues of flies. They fertilize soil, cleanse pastures, steer by the stars, and have a unique relationship with the African elephant (along with many other ungulates). Above all, they are the ideal subject for biological study in an evolving world. This entertaining outline of the development of science from the beetle's perspective will enchant general readers and entomologists alike.

The order Coleoptera (beetles) is arguably the most speciose group of animals, but the evolutionary history of beetles, including the impacts of plant feeding (herbivory) on beetle diversification, remain poorly understood. We inferred the phylogeny of beetles using 4,818 genes for 146 species, estimated timing and rates of beetle diversification using 89 genes for 521 species representing all major lineages and traced the evolution of beetle genes enabling symbiont-independent digestion of lignocellulose using 154 genomes or transcriptomes. Phylogenomic analyses of these uniquely comprehensive datasets resolved previously controversial beetle relationships, dated the origin of Coleoptera to the Carboniferous, and supported the codiversification of beetles and angiosperms. Moreover, plant cell wall-degrading enzymes (PCWDEs) obtained from bacteria and fungi via horizontal gene transfers may have been key to the Mesozoic diversification of herbivorous beetles—remarkably, both major independent origins of specialized herbivory in beetles coincide with the first appearances of an arsenal of PCWDEs encoded in their genomes. Furthermore, corresponding (Jurassic) diversification rate increases suggest that these novel genes triggered adaptive radiations that resulted in nearly half of all living beetle species. We propose that PCWDEs enabled efficient digestion of plant tissues, including lignocellulose in cell walls, facilitating the evolution of uniquely specialized plant-feeding habits, such as leaf mining and stem and wood boring. Beetle diversity thus appears to have resulted from multiple factors, including low extinction rates over a long evolutionary history, codiversification with angiosperms, and adaptive radiations of specialized herbivorous beetles following convergent horizontal transfers of microbial genes encoding PCWDEs.

Significance Many suggest we are approaching a sixth mass extinction event, and yet estimates of how many species exist, and thus how many might become extinct, vary by as much as an order of magnitude. There are few statistically robust methods to estimate global species richness, and here we introduce several new methods, including one that builds on the observation that larger species are often described before smaller species. We combine these, giving equal weight to each, to provide mean global species estimates for the most speciose order, class, and phylum on Earth, beetles, insects, and arthropods (terrestrial). We attempt to aid conservation planning by broadening the range of methods used and bringing greater stability to global estimates for these taxa. It has been suggested that we do not know within an order of magnitude the number of all species on Earth [May RM (1988) Science 241(4872):1441–1449]. Roughly 1.5 million valid species of all organisms have been named and described [Costello MJ, Wilson S, Houlding B (2012) Syst Biol 61(5):871–883]. Given Kingdom Animalia numerically dominates this list and virtually all terrestrial vertebrates have been described, the question of how many terrestrial species exist is all but reduced to one of how many arthropod species there are. With beetles alone accounting for about 40% of all described arthropod species, the truly pertinent question is how many beetle species exist. Here we present four new and independent estimates of beetle species richness, which produce a mean estimate of 1.5 million beetle species. We argue that the surprisingly narrow range (0.9–2.1 million) of these four autonomous estimates—derived from host-specificity relationships, ratios with other taxa, plant:beetle ratios, and a completely novel body-size approach—represents a major advance in honing in on the richness of this most significant taxon, and is thus of considerable importance to the debate on how many species exist. Using analogous approaches, we also produce independent estimates for all insects, mean: 5.5 million species (range 2.6–7.8 million), and for terrestrial arthropods, mean: 6.8 million species (range 5.9–7.8 million), which suggest that estimates for the world’s insects and their relatives are narrowing considerably.

Flight speed is positively correlated with body size in animals 1 . However, miniature featherwing beetles can fly at speeds and accelerations of insects three times their size 2 . Here we show that this performance results from a reduced wing mass and a previously unknown type of wing-motion cycle. Our experiment combines three-dimensional reconstructions of morphology and kinematics in one of the smallest insects, the beetle Paratuposa placentis (body length 395 μm). The flapping bristled wings follow a pronounced figure-of-eight loop that consists of subperpendicular up and down strokes followed by claps at stroke reversals above and below the body. The elytra act as inertial brakes that prevent excessive body oscillation. Computational analyses suggest functional decomposition of the wingbeat cycle into two power half strokes, which produce a large upward force, and two down-dragging recovery half strokes. In contrast to heavier membranous wings, the motion of bristled wings of the same size requires little inertial power. Muscle mechanical power requirements thus remain positive throughout the wingbeat cycle, making elastic energy storage obsolete. These adaptations help to explain how extremely small insects have preserved good aerial performance during miniaturization, one of the factors of their evolutionary success. Three-dimensional reconstructions of morphology and flight mechanics of the beetle Paratuposa placentis reveal adaptations that enable extremely small insects to fly at speeds similar to those of much larger insects.

Beetles represent almost one-fourth of all described species, and knowledge about their relationships and evolution adds to our understanding of biodiversity. We performed a comprehensive phylogenetic analysis of Coleoptera inferred from three genes and nearly 1900 species, representing more than 80% of the world's recognized beetle families. We defined basal relationships in the Polyphaga supergroup, which contains over 300,000 species, and established five families as the earliest branching lineages. By dating the phylogeny, we found that the success of beetles is explained neither by exceptional net diversification rates nor by a predominant role of herbivory and the Cretaceous rise of angiosperms. Instead, the pre-Cretaceous origin of more than 100 present-day lineages suggests that beetle species richness is due to high survival of lineages and sustained diversification in a variety of niches.

Beetles (Coleoptera) are the most diverse and species-rich group of insects, and a robust, time-calibrated phylogeny is fundamental to understanding macroevolutionary processes that underlie their diversity. Here we infer the phylogeny and divergence times of all major lineages of Coleoptera by analyzing 95 protein-coding genes in 373 beetle species, including ~67% of the currently recognized families. The subordinal relationships are strongly supported as Polyphaga (Adephaga (Archostemata, Myxophaga)). The series and superfamilies of Polyphaga are mostly monophyletic. The species-poor Nosodendridae is robustly recovered in a novel position sister to Staphyliniformia, Bostrichiformia, and Cucujiformia. Our divergence time analyses suggest that the crown group of extant beetles occurred ~297 million years ago (Mya) and that ~64% of families originated in the Cretaceous. Most of the herbivorous families experienced a significant increase in diversification rate during the Cretaceous, thus suggesting that the rise of angiosperms in the Cretaceous may have been an ‘evolutionary impetus’ driving the hyperdiversity of herbivorous beetles. The phylogeny of beetles, which represent ~25% of known extant animal species, has been a challenge to resolve. Here, Zhang et al. infer a time-calibrated phylogeny for Coleoptera based on 95 protein-coding genes in 373 species and suggest an association between the hyperdiversification of beetles and the rise of angiosperms.

To investigate the characteristics of compression, buffering and energy dissipation in beetle elytron plates (BEPs), compression experiments were performed on BEPs and honeycomb plates (HPs) with the same wall thickness in different core structures and using different molding methods. The results are as follows: 1) The compressive strength and energy dissipation capacity in the BEP are 2.44 and 5.0 times those in the HP, respectively, when the plates are prepared using the full integrated method (FIM). 2) The buckling stress is directly proportional to the square of the wall thickness (t). Thus, for core structures with equal wall thicknesses, although the core volume of the BEP is 42 percent greater than that of the HP, the mechanical properties of the BEP are several times higher than those of the HP. 3) It is also proven that even when the single integrated method (SIM) is used to prepare BEPs, the properties discussed above remain superior to those of HPs by a factor of several; this finding lays the foundation for accelerating the commercialization of BEPs based on modern manufacturing processes.

Many male animals wield ornaments or weapons of exaggerated proportions. We propose that increased cellular sensitivity to signaling through the insulin/insulin-like growth factor (IGF) pathway may be responsible for the extreme growth of these structures. We document how rhinoceros beetle horns, a sexually selected weapon, are more sensitive to nutrition and more responsive to perturbation of the insulin/IGF pathway than other body structures. We then illustrate how enhanced sensitivity to insulin/IGF signaling in a growing ornament or weapon would cause heightened condition sensitivity and increased variability in expression among individuals—critical properties of reliable signals of male quality. The possibility that reliable signaling arises as a by-product of the growth mechanism may explain why trait exaggeration has evolved so many different times in the context of sexual selection.

Joining dissimilar materials such as plastics and metals in engineered structures remains a challenge 1 . Mechanical fastening, conventional welding and adhesive bonding are examples of techniques currently used for this purpose, but each of these methods presents its own set of problems 2 such as formation of stress concentrators or degradation under environmental exposure, reducing strength and causing premature failure. In the biological tissues of numerous animal and plant species, efficient strategies have evolved to synthesize, construct and integrate composites that have exceptional mechanical properties 3 . One impressive example is found in the exoskeletal forewings (elytra) of the diabolical ironclad beetle, Phloeodes diabolicus . Lacking the ability to fly away from predators, this desert insect has extremely impact-resistant and crush-resistant elytra, produced by complex and graded interfaces. Here, using advanced microscopy, spectroscopy and in situ mechanical testing, we identify multiscale architectural designs within the exoskeleton of this beetle, and examine the resulting mechanical response and toughening mechanisms. We highlight a series of interdigitated sutures, the ellipsoidal geometry and laminated microstructure of which provide mechanical interlocking and toughening at critical strains, while avoiding catastrophic failure. These observations could be applied in developing tough, impact- and crush-resistant materials for joining dissimilar materials. We demonstrate this by creating interlocking sutures from biomimetic composites that show a considerable increase in toughness compared with a frequently used engineering joint. A jigsaw-style configuration of interlocking structures identified in the elytra of the remarkably tough diabolical ironclad beetle, Phloeodes diabolicus , is used to inspire crush-resistant multilayer composites for engineering joints.

Phosphine is the most widely used fumigant for stored grain insect pests, and resistance to phosphine has evolved in several species worldwide. This study was designed to determine the presence of phosphine resistance in 34 populations of Rhyzopertha dominica (F.) collected from the United States and Canada. Adult R. dominica were sampled and subjected to a discriminatory dose toxicity assay of exposure to 20 ppm of phosphine for 20 h of exposure to distinguish a susceptible R. dominica adult by death from a resistant beetle that survives the treatment. All but two of the 34 geographic populations surveyed had some beetles that were resistant to phosphine, and the frequency of resistance varied from 97% in a population from Parlier, California to 0% in beetles from both Carnduff, Saskatchewan and Starbuck, Manitoba. Probit analyses of dose-mortality bioassays with beetles from a laboratory-susceptible strain and those from five of the populations sampled were used to calculate resistance ratio factors (RRs) based on the ratio of LC50 (estimate for the concentration to kill 50% of a test group) in the sampled population to the LC50 for the susceptible strain.The highest RR for the five resistant populations was nearly 596-fold in beetles from Belle Glade, Florida, whereas the lowest RR in that group was 9-fold in Wamego, Kansas.This study revealed that phosphine resistance in R. dominica is common across North America and some populations have levels of resistance that may pose challenges for continued use of phosphine for their management.