Arthropods

Traditionally, measurement of the acute toxicity of pesticides to beneficial arthropods has relied largely on the determination of an acute median lethal dose or concentration. However, the estimated lethal dose during acute toxicity tests may only be a partial measure of the deleterious effects. In addition to direct mortality induced by pesticides, their sublethal effects on arthropod physiology and behavior must be considered for a complete analysis of their impact. An increasing number of studies and methods related to the identification and characterization of these effects have been published in the past 15 years. Review of sublethal effects reported in published literature, taking into account recent data, has revealed new insights into the sublethal effects of pesticides including effects on learning performance, behavior, and neurophysiology. We characterize the different types of sublethal effects on beneficial arthropods, focusing mainly on honey bees and natural enemies, and we describe the methods used in these studies. Finally, we discuss the potential for developing experimental approaches that take into account these sublethal effects in integrated pest management and the possibility of integrating their evaluation in pesticide registration procedures. 81 Annu. Rev. Entomol. 2007.52:81-106. Downloaded from arjournals.annualreviews.org by Dr. Axel Decourtye on 12/15/06. For personal use only. Pollinator: an organism that carries pollen from one flower to another IPM: integrated pest management LD 50 : lethal dose 50% Sublethal effect: an effect (physiological or behavioral) on individuals that survive an exposure to a pesticide (the pesticide dose/concentration can be sublethal or lethal)

Key Words cytoplasmic incompatibility, parthenogenesis, male killing, feminization, α-proteobacteria s Abstract The α-proteobacterium Wolbachia pipientis is a very common cytoplasmic symbiont of insects, crustaceans, mites, and filarial nematodes. To enhance its transmission, W. pipientis has evolved a large scale of host manipulations: parthenogenesis induction, feminization, and male killing. W. pipientis's most common effect is a crossing incompatibility between infected males and uninfected females. Little is known about the genetics and biochemistry of these symbionts because of their fastidious requirements. The affinity of W. pipientis for the microtubules associated with the early divisions in eggs may explain some of their effects. Such inherited microorganisms are thought to have been major factors in the evolution of sex determination, eusociality, and speciation. W. pipientis isolates are also of interest as vectors for the modification of wild insect populations, in the improvement of parasitoid wasps in biological pest control, and as a new method for interfering with diseases caused by filarial nematodes.

Wolbachia are intracellular bacteria found in many species of arthropods and nematodes. They manipulate the reproduction of their arthropod hosts in various ways, may play a role in host speciation and have potential applications in biological pest control. Estimates suggest that at least 20% of all insect species are infected with Wolbachia. These estimates result from several Wolbachia screenings in which numerous species were tested for infection; however, tests were mostly performed on only one to two individuals per species. The actual percent of species infected will depend on the distribution of infection frequencies among species. We present a meta-analysis that estimates percentage of infected species based on data on the distribution of infection levels among species. We used a beta-binomial model that describes the distribution of infection frequencies of Wolbachia, shedding light on the overall infection rate as well as on the infection frequency within species. Our main findings are that (1) the proportion of Wolbachia-infected species is estimated to be 66%, and that (2) within species the infection frequency follows a 'most-orfew' infection pattern in a sense that the Wolbachia infection frequency within one species is typically either very high ( 4 90%) or very low ( o 10%).

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Currently, the use of synthetic chemicals to control insects and arthropods raises several concerns related to environment and human health. An alternative is to use natural products that possess good efficacy and are environmentally friendly. Among those chemicals, essential oils from plants belonging to several species have been extensively tested to assess their repellent properties as a valuable natural resource. The essential oils whose repellent activities have been demonstrated, as well as the importance of the synergistic effects among their components are the main focus of this review. Essential oils are volatile mixtures of hydrocarbons with a diversity of functional groups, and their repellent activity has been linked to the presence of monoterpenes and sesquiterpenes. However, in some cases, these chemicals can work synergistically, improving their effectiveness. In addition, the use of other natural products in the mixture, such as vanillin, could increase the protection time, potentiating the repellent effect of some essential oils. Among the plant families with promising essential oils used as repellents, Cymbopogon spp., Ocimum spp. and Eucalyptus spp. are the most cited. Individual compounds present in these mixtures with high repellent activity include a-pinene, limonene, citronellol, citronellal, camphor and thymol. Finally, although from an economical point of view synthetic chemicals are still more frequently used as repellents than essential oils, these natural products have the potential to provide efficient, and safer repellents for humans and the environment.

The remarkable antiquity, diversity and ecological significance of arthropods have inspired numerous attempts to resolve their deep phylogenetic history, but the results of two decades of intensive molecular phylogenetics have been mixed 1-7 . The discovery that terrestrial insects (Hexapoda) are more closely related to aquatic Crustacea than to the terrestrial centipedes and millipedes 2,8 (Myriapoda) was an early, if exceptional, success. More typically, analyses based on limited samples of taxa and genes have generated results that are inconsistent, weakly supported and highly sensitive to analytical conditions 7,9,10 . Here we present strongly supported results from likelihood, Bayesian and parsimony analyses of over 41 kilobases of aligned DNA sequence from 62 single-copy nuclear protein-coding genes from 75 arthropod species. These species represent every major arthropod lineage, plus five species of tardigrades and onychophorans as outgroups. Our results strongly support Pancrustacea (Hexapoda plus Crustacea) but also strongly favour the traditional morphology-based Mandibulata 11 (Myriapoda plus Pancrustacea) over the molecule-based Paradoxopoda (Myriapoda plus Chelicerata) 2,5,12 . In addition to Hexapoda, Pancrustacea includes three major extant lineages of 'crustaceans', each spanning a significant range of morphological disparity. These are Oligostraca (ostracods, mystacocarids, branchiurans and pentastomids), Vericrustacea (malacostracans, thecostracans, copepods and branchiopods) and Xenocarida (cephalocarids and remipedes). Finally, within Pancrustacea we identify Xenocarida as the long-sought sister group to the Hexapoda, a result confirming that 'crustaceans' are not monophyletic. These results provide a statistically well-supported phylogenetic framework for the largest animal phylum and represent a step towards ending the often-heated, century-long debate on arthropod relationships.

engrailed is a homeobox gene that has an important role in Drosophila segmentation. Genes homologous to engrailed have been identified in several other organisms. Here we describe a monoclonal antibody that recognizes a conserved epitope in the homeodomain of engrailed proteins of a number of different arthropods, annelids, and chordates; we use this antibody to isolate the grasshopper engrailed gene. In Drosophila embryos, the antibody reveals engrailed protein in the posterior portion of each segment during segmentation, and in a segmentally reiterated subset of neuronal cells during neurogenesis. Other arthropods, including grasshopper and two crustaceans, have similar patterns of engrailed expression. However, these patterns of expression are not shared by the annelids or chordates we examined. Our results provide the most comprehensive view that has been obtained of how expression patterns of a regulatory gene vary during evolution. On the basis of these patterns, we suggest that engrailed is a gene whose ancestral function was in neurogenesis and whose function was co-opted during the evolution of segmentation in the arthropods, but not in the annelids and chordates.

Most models of optimal progeny size assume that there is a trade-off between progeny size and number, and that progeny fitness increases with increasing investment per young. We find that both assumptions are supported by empirical studies but that the trade-off is less apparent when organisms are iteroparous, use adult-acquired resources for reproduction, or provide parental care. We then review patterns of variation in progeny size among species, among populations within species, among individuals within populations, and among progeny produced by a single female. We argue that much of the variation in progeny size among species, and among populations within species, is likely due to variation in natural selection. However, few studies have manipulated progeny environments and demonstrated that the relationship between progeny size and fitness actually differs among environments, and fewer still have demonstrated why selection favors different sized progeny in different environments. We argue that much of the variation in progeny size among females within populations, and among progeny produced by a single female, is probably nonadaptive. However, some species of arthropods exhibit plasticity in progeny size in response to several environmental factors, and much of this plasticity is likely adaptive. We conclude that advances in theory have substantially outpaced empirical data. We hope that this review will stimulate researchers to examine the specific factors that result in variation in selection on progeny size within and among populations, and how this variation in selection influences the evolution of the patterns we observe.

Wolbachia are intracellular bacteria that manipulate the reproduction of their arthropod hosts in remarkable ways. They are predominantly transmitted vertically from mother to offspring but also occasionally horizontally between species. In doing so, they infect a huge range of arthropod species worldwide. Recently, a statistical analysis estimated the infection frequency of Wolbachia among arthropod hosts to be 66%. At the same time, the authors of this analysis highlighted some weaknesses of the underlying data and concluded that in order to improve the estimate, a larger number of individuals per species should be assayed and species be chosen more randomly. Here we apply the statistical approach to a more appropriate data set from a recent survey that tested both a broad range of species and a sufficient number of individuals per species. Indeed, we find a substantially different infection frequency: We now estimate the proportion of Wolbachiainfected species to be around 40% which is lower than the previous estimate but still points to a surprisingly high number of arthropods harboring the bacteria. Notwithstanding this difference, we confirm the previous result that, within a given species, typically most or only a few individuals are infected. Moreover, we extend our analysis to include several reproductive parasites other than Wolbachia that were also screened for in the aforementioned empirical survey. For these symbionts we find a large variation in estimated infection frequencies and corroborate the finding that Wolbachia are the most abundant endosymbionts among arthropod species.

Soil health is presented as an integrative property that reflects the capacity of soil to respond to agricultural intervention, so that it continues to support both the agricultural production and the provision of other ecosystem services. The major challenge within sustainable soil management is to conserve ecosystem service delivery while optimizing agricultural yields. It is proposed that soil health is dependent on the maintenance of four major functions: carbon transformations; nutrient cycles; soil structure maintenance; and the regulation of pests and diseases. Each of these functions is manifested as an aggregate of a variety of biological processes provided by a diversity of interacting soil organisms under the influence of the abiotic soil environment. Analysis of current models of the soil community under the impact of agricultural interventions (particularly those entailing substitution of biological processes with fossil fuel-derived energy or inputs) confirms the highly integrative pattern of interactions within each of these functions and leads to the conclusion that measurement of individual groups of organisms, processes or soil properties does not suffice to indicate the state of the soil health. A further conclusion is that quantifying the flow of energy and carbon between functions is an essential but non-trivial task for the assessment and management of soil health.

Metagenome studies have retrieved vast amounts of sequence data from a variety of environments leading to new discoveries and insights into the uncultured microbial world. Except for very simple communities, the encountered diversity has made fragment assembly and the subsequent analysis a challenging problem. A taxonomic characterization of metagenomic fragments is required for a deeper understanding of shotgunsequenced microbial communities, but success has mostly been limited to sequences containing phylogenetic marker genes. Here we present PhyloPythia, a composition-based classifier that combines higher-level generic clades from a set of 340 completed genomes with sample-derived population models. Extensive analyses on synthetic and real metagenome data sets showed that PhyloPythia allows the accurate classification of most sequence fragments across all considered taxonomic ranks, even for unknown organisms. The method requires no more than 100 kb of training sequence for the creation of accurate models of sample-specific populations and can assign fragments Z1 kb with high specificity.

Viral vectors are the most efficient tools for gene delivery, and the search for tissue-specific infecting viruses is important for the development of in vivo gene therapy strategies. The baculovirus Autographa californica nuclear polyhedrosis virus is widely used as a vector for expression of foreign genes in insect cells, and its host specificity is supposed to be restricted to arthropods. Here we demonstrate that recombinant A. californica nuclear polyhedrosis virus is efficiently taken up by human hepatocytes via an endosomal pathway. High-level reporter gene expression from heterologous promoters was observed in human and rabbit hepatocytes in vitro. Mouse hepatocytes and some other epithelial cell types are targeted at a considerably lower rate. The efficiency of gene transfer by baculovirus considerably exceeds that obtained by calcium phosphate or lipid transfection. These properties of baculovirus suggest a use for it as a vector for liver-directed gene transfer but highlight a potential risk in handling certain recombinant baculoviruses.

Ecological niche models (ENMs) provide a means of characterizing the spatial distribution of suitable conditions for species, and have recently been applied to the challenge of locating potential distributional areas at the Last Glacial Maximum (LGM) when unfavorable climate conditions led to range contractions and fragmentation. Here, we compare and contrast ENM-based reconstructions of LGM refugial locations with those resulting from the more traditional molecular genetic and phylogeographic predictions. We examined 20 North American terrestrial vertebrate species from different regions and with different range sizes for which refugia have been identified based on phylogeographic analyses, using ENM tools to make parallel predictions. We then assessed the correspondence between the two approaches based on spatial overlap and areal extent of the predicted refugia. In 14 of the 20 species, the predictions from ENM and predictions based on phylogeographic studies were significantly spatially correlated, suggesting that the two approaches to development of refugial maps are converging on a similar result. Our results confirm that ENM scenario exploration can provide a useful complement to molecular studies, offering a less subjective, spatially explicit hypothesis of past geographic patterns of distribution.

Fifteen species richness estimators (three asymptotic based on species accumulation curves, 11 nonparametric, and one based in the species-area relationship) were compared by examining their performance in estimating the total species richness of epigean arthropods in the Azorean Laurisilva forests. Data obtained with standardized sampling of 78 transects in natural forest remnants of five islands were aggregated in seven different grains (i.e. ways of defining a single sample): islands, natural areas, transects, pairs of traps, traps, database records and individuals to assess the effect of using different sampling units on species richness estimations. 2. Estimated species richness scores depended both on the estimator considered and on the grain size used to aggregate data. However, several estimators (ACE, Chao1, Jackknife1 and 2 and Bootstrap) were precise in spite of grain variations. Weibull and several recent estimators [proposed by Rosenzweig et al . ( Conservation Biology , 2003, 17 , 864-874), and Ugland et al . ( Journal of Animal Ecology , 2003, 72 , 888-897)] performed poorly. 3. Estimations developed using the smaller grain sizes (pair of traps, traps, records and individuals) presented similar scores in a number of estimators (the above-mentioned plus ICE, Chao2, Michaelis-Menten, Negative Exponential and Clench). The estimations from those four sample sizes were also highly correlated. 4. Contrary to other studies, we conclude that most species richness estimators may be useful in biodiversity studies. Owing to their inherent formulas, several nonparametric and asymptotic estimators present insensitivity to differences in the way the samples are aggregated. Thus, they could be used to compare species richness scores obtained from different sampling strategies. Our results also point out that species richness estimations coming from small grain sizes can be directly compared and other estimators could give more precise results in those cases. We propose a decision framework based on our results and on the literature to assess which estimator should be used to compare species richness scores of different sites, depending on the grain size of the original data, and of the kind of data available (species occurrence or abundance data).

A clone which conferred lipolytic activity at low temperature was identified from a fosmid library constructed from a South China Sea marine sediment sample. The gene responsible, estF, consisted of 1,080 bp that encoded 359 amino acid residues, with a typical N-terminal signal peptide of 28 amino acid residues. A phylogenetic analysis of amino acid sequence with other lipolytic enzymes revealed that EstF and seven closely related putative lipolytic enzymes comprised a unique clade in the phylogenetic tree. Moreover, these hypothetic esterases showed unique conservative sites in the amino acid sequence. The recombinant EstF was overexpressed and purified, and its biochemical properties were partially characterized. The optimal substrate for EstF to hydrolyze among a panel of p-nitrophenyl esters (C2 to C16) was pnitrophenyl butyrate (C4), with a K m of 0.46 mM. Activity quickly decreased with substrates containing an acyl chain length longer than 10 carbons. We found that EstF was active in the temperature range of 0-60°C, showed the best activity at 50°C, but was unstable at 60°C. It exhibited a high level of activity in the pH range of 7.0-10.0 showing the highest activity at pH 9.0.

The complete 18s rDNA gene sequence of Mucrobiotus group hufelundi (Tardigrada) was obtained and aligned with 18s rDNA and rRNA gene sequences of 24 metazoans (mainly protostomes). Discrete character (maximumparsimony) and distance (neighbor-joining) methods were used to infer their phylogeny. The evolution of bootstrap proportions with sequence length (pattern of resolved nodes, PRN) was studied to test the resolution of the nodes in neighbor-joining trees. The results show that arthropods are monophyletic. Tardigrades represent the sister group of arthropods (in parsimony analyses) or they are related with crustaceans (distance analysis and PRN). Arthropoda are divided into two main evolutionary lines, the Hexapoda + Crustacea line (weakly supported), and the Myriapoda + Chelicerata line. The Hexapoda + Crustacea line includes Pentastomida, but the internal resolution is far from clear. The Insecta (Ectognatha) are monophyletic, but no evidence for the monophyly of Hexapoda is found. The Chelicerata are a monophyletic group and the Myriapoda cluster close to Arachnida. Overall, the results obtained represent the first molecular evidence for a Tardigrada + Arthropoda clade. In addition, the congruence between molecular phylogenies of the Arthropoda from other authors and this obtained here indicates the need to review those obtained solely on morphological characters.

Insight into the origin and early evolution of the animal phyla requires an understanding of how animal groups are related to one another. Thus, we set out to explore animal phylogeny by analyzing with maximum parsimony 138 morphological characters from 40 metazoan groups, and 304 18S rDNA sequences, both separately and together. Both types of data agree that arthropods are not closely related to annelids: the former group with nematodes and other molting animals (Ecdysozoa), and the latter group with molluscs and other taxa with spiral cleavage. Furthermore, neither brachiopods nor chaetognaths group with deuterostomes; brachiopods are allied with the molluscs and annelids (Lophotrochozoa), whereas chaetognaths are allied with the ecdysozoans. The major discordance between the two types of data concerns the rooting of the bilaterians, and the bilaterian sister-taxon. Morphology suggests that the root is between deuterostomes and protostomes, with ctenophores the bilaterian sister-group, whereas 18S rDNA suggests that the root is within the Lophotrochozoa with acoel flatworms and gnathostomulids as basal bilaterians, and with cnidarians the bilaterian sister-group. We suggest that this basal position of acoels and gnathostomulids is artifactal because for 1,000 replicate phylogenetic analyses with one random sequence as outgroup, the majority root with an acoel flatworm or gnathostomulid as the basal ingroup lineage. When these problematic taxa are eliminated from the matrix, the combined analysis suggests that the root lies between the deuterostomes and protostomes, and Ctenophora is the bilaterian sister-group. We suggest that because chaetognaths and lophophorates, taxa traditionally allied with deuterostomes, occupy basal positions within their respective protostomian clades, deuterostomy most likely represents a suite of characters plesiomorphic for bilaterians.

An international initiative is developing a scientifically rigorous approach to evaluate the potential risks to nontarget arthropods (NTAs) posed by insect-resistant, genetically modified (IRGM) crops. It adapts the tiered approach to risk assessment that is used internationally within regulatory toxicology and environmental sciences. The approach focuses on the formulation and testing of clearly stated risk hypotheses, making maximum use of available data and using formal decision guidelines to progress between testing stages (or tiers). It is intended to provide guidance to regulatory agencies that are currently developing their own NTA risk assessment guidelines for IRGM crops and to help harmonize regulatory requirements between different countries and different regions of the world.

Islands have traditionally been considered to be any relatively small body of land completely surrounded by water. However, their primary biological characteristic, an extended period of isolation from a source of colonists, is common also to many situations on continents. Accordingly, theories and predictions developed for true islands have been applied to a huge array of systems, from rock pools, to single tree species in forests, to oceanic islands. Here, we examine the literature on islands in the broadest sense (i.e., whether surrounded by water or any other uninhabitable matrix) as it pertains to terrestrial arthropods. We categorize islands according to the features they share. The primary distinction between different island systems is "darwinian" islands (formed de novo) and "fragment" islands. In the former, the islands have never been in contact with the source of colonists and have abundant "empty" ecological niche space. On these islands, species numbers will initially increase through immigration, the rate depending on the degree of isolation. If isolation persists, over time species formation will result in "neo-endemics." When isolation is extreme, the ecological space will gradually be filled through speciation (rather than immigration) and adaptive radiation of neo-endemics. Fragment islands are fundamentally different. In these islands, the ecological space will initially be filled as a consequence of connection to the source of colonists prior to insularization. Species numbers will decrease following fragmentation through the process of relaxation. If these islands become more isolated, species will eventually arise through relictualization with the formation of "paleo-endemics." Given sufficient time, this process can result in generic level endemism on ancient fragment islands, a phenomenon well illustrated in Madagascar and New Zealand. Recognizing the distinction between the different kinds of islands is fundamental for understanding emerging patterns on each, in particular speciation, biodiversity (e.g., neo-endemics versus paleo-endemics), and conservation (e.g., naiveté in interactions with alien species).

Plant diversity is predicted to be positively linked to the diversity of herbivores and predators in a foodweb. Yet, the relationship between plant and animal diversity is explained by a variety of competing hypotheses, with mixed empirical results for each hypothesis. We sampled arthropods for over a decade in an experiment that manipulated the number of grassland plant species. We found that herbivore and predator species richness were strongly, positively related to plant species richness, and that these relationships were caused by different mechanisms at herbivore and predator trophic levels. Even more dramatic was the threefold increase, from low- to high-plant species richness, in abundances of predatory and parasitoid arthropods relative to their herbivorous prey. Our results demonstrate that, over the long term, the loss of plant species propagates through food webs, greatly decreasing arthropod species richness, shifting a predator-dominated trophic structure to being herbivore dominated, and likely impacting ecosystem functioning and services.

In the past, molecular clocks have been used to estimate divergence times among animal phyla, but those time estimates have varied widely (1200^670 million years ago, Ma). In order to obtain time estimates that are more robust, we have analysed a larger number of genes for divergences among three well-represented animal phyla, and among plants, animals and fungi. The time estimate for the chordate^arthropod divergence, using 50 genes, is 993 AE46 Ma. Nematodes were found to have diverged from the lineage leading to arthropods and chordates at 1177 AE79 Ma. Phylogenetic analyses also show that a basal position of nematodes has strong support (p499%) and is not the result of rate biases. The three-way split (relationships unresolved) of plants, animals and fungi was estimated at 1576 AE 88 Ma. By inference, the basal animal phyla (Porifera, Cnidaria, Ctenophora) diverged between about 1200^1500 Ma. This suggests that at least six animal phyla originated deep in the Precambrian, more than 400 million years earlier than their ¢rst appearance in the fossil record.

The attack of a plant by herbivorous arthropods can result in considerable changes in the plant's chemical phenotype. The emission of so-called herbivore-induced plant volatiles (HIPV) results in the attraction of carnivorous enemies of the herbivores that induced these changes. HIPV induction has predominantly been investigated for interactions between one plant and one attacker. However, in nature plants are exposed to a variety of attackers, either simultaneously or sequentially, in shoots and roots, causing much more complex interactions than have usually been investigated in the context of HIPV. To develop an integrated view of how plants respond to their environment, we need to know more about the ways in which multiple attackers can enhance, attenuate, or otherwise alter HIPV responses. A multidisciplinary approach will allow us to investigate the underlying mechanisms of HIPV emission in terms of phytohormones, transcriptional responses and biosynthesis of metabolites in an effort to understand these complex plant-arthropod interactions.

More than 5000 introductions of about 2000 species of exotic arthropod agents for control of arthropod pests in 196 countries or islands during the past 120 years rarely have resulted in negative environmental effects. Yet, risks of environmental effects caused by releases of exotics are of growing concern. Twenty countries have implemented regulations for release of biological control agents. Soon, the International Standard for Phytosanitary Measures (ISPM3) will become the standard for all biological control introductions worldwide, but this standard does not provide methods by which to assess environmental risks. This review summarizes documented nontarget effects and discusses the development and application of comprehensive and quick-scan environmental risk assessment methods.

The application of DNA barcoding to dietary studies allows prey taxa to be identified in the absence of morphological evidence and permits a greater resolution of prey identity than is possible through direct examination of faecal material. For insectivorous bats, which typically eat a great diversity of prey and which chew and digest their prey thoroughly, DNAbased approaches to diet analysis may provide the only means of assessing the range and diversity of prey within faeces. Here, we investigated the effectiveness of DNA barcoding in determining the diets of bat species that specialize in eating different taxa of arthropod prey. We designed and tested a novel taxon-specific primer set and examined the performance of short barcode sequences in resolving prey species. We recovered prey DNA from all faecal samples and subsequent cloning and sequencing of PCR products, followed by a comparison of sequences to a reference database, provided species-level identifications for 149 ⁄ 207 (72%) clones. We detected a phylogenetically broad range of prey while completely avoiding detection of nontarget groups. In total, 37 unique prey taxa were identified from 15 faecal samples. A comparison of DNA data with parallel morphological analyses revealed a close correlation between the two methods. However, the sensitivity and taxonomic resolution of the DNA method were far superior. The methodology developed here provides new opportunities for the study of bat diets and will be of great benefit to the conservation of these ecologically important predators.

In agriculture, sustainability can be linked to ecological resilience. In view of present or imminent environmental changes in agricultural landscapes, the diversity of species and genotypes, particularly of potential beneficials and alternative prey, may become of increasing importance. However, the available methods and empirical data concerning species diversity of above-ground insects in agricultural landscapes do not yet allow comprehensive evaluation. Standardized inventory methods must be used more rigorously and over longer time periods to detect significant differences in space and in time. Indicator groups for biodiversity estimates must be defined.

Chemoreception is a biological process essential for the survival of animals, as it allows the recognition of important volatile cues for the detection of food, egg-laying substrates, mates, or predators, among other purposes. Furthermore, its role in pheromone detection may contribute to evolutionary processes, such as reproductive isolation and speciation. This key role in several vital biological processes makes chemoreception a particularly interesting system for studying the role of natural selection in molecular adaptation. Two major gene families are involved in the perireceptor events of the chemosensory system: the odorantbinding protein (OBP) and chemosensory protein (CSP) families. Here, we have conducted an exhaustive comparative genomic analysis of these gene families in 20 Arthropoda species. We show that the evolution of the OBP and CSP gene families is highly dynamic, with a high number of gains and losses of genes, pseudogenes, and independent origins of subfamilies. Taken together, our data clearly support the birth-and-death model for the evolution of these gene families with an overall high gene turnover rate. Moreover, we show that the genome organization of the two families is significantly more clustered than expected by chance and, more important, that this pattern appears to be actively maintained across the Drosophila phylogeny. Finally, we suggest the homologous nature of the OBP and CSP gene families, dating back their most recent common ancestor after the terrestrialization of Arthropoda (380-450 Ma) and we propose a scenario for the origin and diversification of these families.

Arthropods were the first animals to conquer land and air. They encompass more than three quarters of all described living species. This extraordinary evolutionary success is based on an astoundingly wide array of highly adaptive body organizations. A lack of robustly resolved phylogenetic relationships, however, currently impedes the reliable reconstruction of the underlying evolutionary processes. Here, we show that phylogenomic data can substantially advance our understanding of arthropod evolution and resolve several conflicts among existing hypotheses. We assembled a data set of 233 taxa and 775 genes from which an optimally informative data set of 117 taxa and 129 genes was finally selected using new heuristics and compared with the unreduced data set. We included novel expressed sequence tag (EST) data for 11 species and all published phylogenomic data augmented by recently published EST data on taxonomically important arthropod taxa. This thorough sampling reduces the chance of obtaining spurious results due to stochastic effects of undersampling taxa and genes. Orthology prediction of genes, alignment masking tools, and selection of most informative genes due to a balanced taxa-gene ratio using new heuristics were established. Our optimized data set robustly resolves major arthropod relationships. We received strong support for a sister group relationship of onychophorans and euarthropods and strong support for a close association of tardigrades and cycloneuralia. Within pancrustaceans, our analyses yielded paraphyletic crustaceans and monophyletic hexapods and robustly resolved monophyletic endopterygote insects. However, our analyses also showed for few deep splits that were recently thought to be resolved, for example, the position of myriapods, a remarkable sensitivity to methods of analyses.

Background Rickettsia are intracellular symbionts of eukaryotes that are best known for infecting and causing serious diseases in humans and other mammals. All known vertebrate-associated Rickettsia are vectored by arthropods as part of their life-cycle, and many other Rickettsia are found exclusively in arthropods with no known secondary host. However, little is known about the biology of these latter strains. Here, we have identified 20 new strains of Rickettsia from arthropods, and constructed a multi-gene phylogeny of the entire genus which includes these new strains. Results We show that Rickettsia are primarily arthropod-associated bacteria, and identify several novel groups within the genus. Rickettsia do not co-speciate with their hosts but host shifts most often occur between related arthropods. Rickettsia have evolved adaptations including transmission through vertebrates and killing males in some arthropod hosts. We uncovered one case of horizontal gene transfer among Rickettsia, where a strain is a chimera from two distantly related groups, but multi-gene analysis indicates that different parts of the genome tend to share the same phylogeny. Conclusion Approximately 150 million years ago, Rickettsia split into two main clades, one of which primarily infects arthropods, and the other infects a diverse range of protists, other eukaryotes and arthropods. There was then a rapid radiation about 50 million years ago, which coincided with the evolution of life history adaptations in a few branches of the phylogeny. Even though Rickettsia are thought to be primarily transmitted vertically, host associations are short lived with frequent switching to new host lineages. Recombination throughout the genus is generally uncommon, although there is evidence of horizontal gene transfer. A better understanding of the evolution of Rickettsia will help in the future to elucidate the mechanisms of pathogenicity, transmission and virulence.

Background: Uncertainty persists over the environmental effects of genetically-engineered crops that produce the insecticidal Cry proteins of Bacillus thuringiensis (Bt). We performed meta-analyses on a modified public database to synthesize current knowledge about the effects of Bt cotton, maize and potato on the abundance and interactions of arthropod non-target functional guilds. Methodology/Principal Findings: We compared the abundance of predators, parasitoids, omnivores, detritivores and herbivores under scenarios in which neither, only the non-Bt crops, or both Bt and non-Bt crops received insecticide treatments. Predators were less abundant in Bt cotton compared to unsprayed non-Bt controls. As expected, fewer specialist parasitoids of the target pest occurred in Bt maize fields compared to unsprayed non-Bt controls, but no significant reduction was detected for other parasitoids. Numbers of predators and herbivores were higher in Bt crops compared to sprayed non-Bt controls, and type of insecticide influenced the magnitude of the difference. Omnivores and detritivores were more abundant in insecticide-treated controls and for the latter guild this was associated with reductions of their predators in sprayed non-Bt maize. No differences in abundance were found when both Bt and non-Bt crops were sprayed. Predator-to-prey ratios were unchanged by either Bt crops or the use of insecticides; ratios were higher in Bt maize relative to the sprayed non-Bt control. Conclusions/Significance: Overall, we find no uniform effects of Bt cotton, maize and potato on the functional guilds of non-target arthropods. Use of and type of insecticides influenced the magnitude and direction of effects; insecticde effects were much larger than those of Bt crops. These meta-analyses underscore the importance of using controls not only to isolate the effects of a Bt crop per se but also to reflect the replacement of existing agricultural practices. Results will provide researchers with information to design more robust experiments and will inform the decisions of diverse stakeholders regarding the safety of transgenic insecticidal crops.

Many intracellular micro-organisms are now known to cause reproductive abnormalities and other phenomena in their hosts. The endosymbiont Wolbachia is the best known of these reproductive manipulators owing to its extremely high incidence among arthropods and the diverse host effects it has been implicated as causing. However, recent evidence suggests that another intracellular bacterium, a Cytophaga-like organism (CLO), may also induce several reproductive effects in its hosts. Here, we present the first survey of arthropod hosts for infection by the CLO. We use a sensitive hemi-nested polymerase chain reaction method to screen 223 species from 20 arthropod orders for infection by the CLO and Wolbachia. The results indicate that, although not as prevalent as Wolbachia, the CLO infects a significant number of arthropod hosts (ca. 7.2%). In addition, double infections of the CLO and Wolbachia were found in individuals of seven arthropod species. Sequencing analysis of the 16S rDNA region of the CLO indicates evidence for horizontal transmission of the CLO strains. We discuss these results with reference to future studies on host effects induced by intracellular micro-organisms.

The Arrhenius equation has emerged as the favoured model for describing the temperature dependence of consumption in predator-prey models. To examine the relevance of this equation, we undertook a metaanalysis of published relationships between functional response parameters and temperature. We show that, when plotted in lin-log space, temperature dependence of both attack rate and maximal ingestion rate exhibits a hump-shaped relationship and not a linear one as predicted by the Arrhenius equation. The relationship remains significantly downward concave even when data from temperatures above the peak of the hump are discarded. Temperature dependence is stronger for attack rate than for maximal ingestion rate, but the thermal optima are not different. We conclude that the use of the Arrhenius equation to describe consumption in predator-prey models requires the assumption that temperatures above thermal optima are unimportant for population and community dynamics, an assumption that is untenable given the available data.

Arguably the majority of species on Earth utilise tropical rainforest canopies, and much progress has been made in describing arboreal assemblages, especially for arthropods. The most commonly described patterns for tropical rainforest insect communities are host specificity, spatial specialisation (predominantly vertical stratification), and temporal changes in abundance (seasonality and circadian rhythms). Here I review the recurrent results with respect to each of these patterns and discuss the evolutionary selective forces that have generated them in an attempt to unite these patterns in a holistic evolutionary framework. I propose that species can be quantified along a generalist-specialist scale not only with respect to host specificity, but also other spatial and temporal distribution patterns, where specialisation is a function of the extent of activity across space and time for particular species. When all of these distribution patterns are viewed through the paradigm of specialisation, hypotheses that have been proposed to explain the evolution of host specificity can also be applied to explain the generation and maintenance of other spatial and temporal distribution patterns. The main driver for most spatial and temporal distribution patterns is resource availability. Generally, the distribution of insects follows that of the resources they exploit, which are spatially stratified and vary temporally in availability. Physiological adaptations are primarily important for host specificity, where nutritional and chemical variation among host plants in particular, but also certain prey species and fungi, influence host range. Physiological tolerances of abiotic conditions are also important for explaining the spatial and temporal distributions of some insect species, especially in drier forest environments where desiccation is an ever-present threat. However, it is likely that for most species in moist tropical rainforests, abiotic conditions are valuable indicators of resource availability, rather than physiologically limiting factors. Overall, each distribution pattern is influenced by the same evolutionary forces, but at differing intensities. Consequently, each pattern is linked and not mutually exclusive of the other distribution patterns. Most studies have examined each of these patterns in isolation. Future work should focus on examining the evolutionary drivers of these patterns in concert. Only then can the relative strength of resource availability and distribution, host defensive phenotypes, and biotic and abiotic interactions on insect distribution patterns be determined.

A blood-sucking habit appeared independently several times in the course of arthropod evolution. However, from more than a million species of insects and arachnids presently living on earth, only about 14,000 species developed the capacity to feed on vertebrate blood. This figure suggests the existence of severe physiological constraints for the evolution of hematophagy, implying the selective advantage of special adaptations related to the use of blood as a food source. Digestion of vertebrate hemoglobin in the midgut of blood-feeding arthropods results in the production of large amounts of heme, a potentially cytotoxic molecule. Here we will review mechanisms by which heme can exert biological damage, together with a wide spectrum of adaptations developed by blood-feeding insects and ticks to counteract its deleterious effects. In spite of the existence of a great molecular diversity of protective mechanisms, different hematophagous organisms developed convergent solutions that may be physiologically equivalent.

The earliest fossil evidence of terrestrial animal activity is from the Ordovician, 450 million years ago (Ma). However, there are earlier animal fossils, and most molecular clocks suggest a deep origin of animal phyla in the Precambrian, leaving open the possibility that animals colonized land much earlier than the Ordovician. To further investigate the time of colonization of land by animals, we sequenced two nuclear genes, glyceraldehyde-3-phosphate dehydrogenase and enolase, in representative arthropods and conducted phylogenetic and molecular clock analyses of those and other available DNA and protein sequence data. To assess the robustness of animal molecular clocks, we estimated the deuterostome-arthropod divergence using the arthropod fossil record for calibration and tunicate instead of vertebrate sequences to represent Deuterostomia. Nine nuclear and 15 mitochondrial genes were used in phylogenetic analyses and 61 genes were used in molecular clock analyses.

Limb morphology across the arthropods is reviewed using external morphological and internal anatomical data from both recent and fossil arthropods. Evolutionary trends in limb structure are identified primarily by reference to the more rigorous of the many existing phylogenetic schemes, but no major new phylogenetic inferences are presented. Tagmosis patterns are not considered, although the origins and patterns of heteronomy within the postantennulary limb series are analysed. The phenomenon of annulation is examined and two basic types of annuli are recognised : terminal and intercalary. The annulation of the apical segment of a limb results in the formation of terminal flagella, and is typical of primarily sensory appendages such as insect and malacostracan antennules and maxillary palps of some hexapods. Intercalary annulation, arising by subdivision of existing subterminal segments, is common, particularly in the tarsal region of arthropodan walking limbs. Differentiating between segments and annuli is discussed and is recognised as a limiting factor in the interpretation of fossils, which usually lack information on intrinsic musculature, and in the construction of groundplans. Rare examples of secondary segmentation, where the criteria for distinguishing between segments and annuli fail, are also highlighted. The basic crown-group arthropodan limb is identified as tripartite, comprising protopodite, telopodite and exopodite, and the basic segmentation patterns of each of these parts are hypothesised. Possible criteria are discussed that can be used for establishing the boundary between protopodite and telopodite in limbs that are uniramous through loss of the exopodite. The subdivision of the protopodite, which is typical of the postantennulary limbs of mandibulates, is examined. The difficulties resulting from the partial or complete failure of expression of articulations within the mandibulate protopodite and subsequent incorporation of partial protopodal segments into the body wall, are also discussed. The development and homology between the various exites, including gills, on the postantennulary limbs of arthropods are considered in some detail, and the question of the possible homology between crustacean gills and insect wings is critically addressed. The hypothesis that there are only two basic limb types in arthropods, antennules and postantennulary limbs, is proposed and its apparent contradiction by the transformation of antennules into walking limbs by homeotic mutation is discussed with respect to the appropriate level of serial homology between these limbs.