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How distantly related are eusocial insects? Aren't members of a species much more related than 1/4, 1/2, or 3/4?

How distantly related are eusocial insects? Aren't members of a species much more related than 1/4, 1/2, or 3/4?


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How many genes do we share with our mother?

I went to a lecture that talked about the behavior of social insects in terms of their relatedness of genes. For instance, workers were 3/4ths related to each other, so it was in their gene's interest to care for each other instead of having their own, half-related offspring.

However isn't it the case that members of the same species are much more related than 1/2 or 3/4s? I've read that we share anywhere from 95-99% of our genes with chimpanzees. Also I believe I've read (perhaps in Pinker) that we share some 60% of our genes with daffodils.

So when they talk about the relatedness of social insects, and they say it's 1/2 or 2/3 or 1/4, aren't they really talking about the 1% to 5% of genes that are in play in sexual reproduction within the species? In other words, aren't I 97% + 1/2 * 3% related to my sister, assuming human beings share 97% of our genes?


In evolutionary genetic comparison, you are talking about members within species. They will share almost all genes, because if they didn't they would belong to a different species.

However, within species there exist different versions of the same genes, called 'alleles'. When we say that you are 0.5 related to each of your parents, we mean that statistically, 50% of your alleles should be those which your father has, and 50% of your alleles are those passed down from your mother.

Eusocial insects have different mechanisms. Bee males are produced without fertilisation, meaning that they only have one copy of each bee gene. When the male produces sperm, it only has this one set, so all sperms end up carrying the same set of alleles.

Females on the other hand have the normal double set, with two different versions of each gene. So if you look at one gene, half of the female's egg should have one version and the other half should have the other version. All females of one hive are produced by the same queen and the one male that she mated with. Remember, the male only has one set, so the versions coming from the male are the same in all female offspring.

This means a female's genes are made up of: 50% from the father (these are the same across all females) and 50% from the mother (where half the females have one version and half have the other version). Statistically, this means that looking at one gene, there is a 75% chance that two bees will have the same version of that gene.


The part that is probably specific to this question is that some species of insects divide their population in the reproducing individuals, that are usually diploid and the working individuals, that are sometimes haploids, meaning they only have one copy of their set of chromosomes. This means that the workers have about half the amount of DNA material compared to reproducing individuals, although they share that copy in full with the diploid individuals.

There are variations of this mechanism in different species. Some species keep the copies but epigenetically "shut down" one of the copies. Others shed specific bits of the genome in the somatic cells. Sometimes this loss of DNA is not perfect, so different parts of the genome are lost just by chance.

We humans don't lose any material at all, we just epigenetically "shut down" one of the X chromosomes in XX individuals -- females.


Tool use by animals

Tool use by animals is a phenomenon in which an animal uses any kind of tool in order to achieve a goal such as acquiring food and water, grooming, defense, communication, recreation or construction. Originally thought to be a skill possessed only by humans, some tool use requires a sophisticated level of cognition. There is considerable discussion about the definition of what constitutes a tool and therefore which behaviours can be considered true examples of tool use. A wide range of animals, including mammals, birds, fish, cephalopods, and insects, are considered to use tools.

Primates are well known for using tools for hunting or gathering food and water, cover for rain, and self-defence. Chimpanzees have often been the object of study in regard to their usage of tools, most famously by Jane Goodall, since these animals are frequently kept in captivity and are closely related to humans. Wild tool-use in other primates, especially among apes and monkeys, is considered relatively common, though its full extent remains poorly documented, as many primates in the wild are mainly only observed distantly or briefly when in their natural environments and living without human influence. Some novel tool-use by primates may arise in a localized or isolated manner within certain unique primate cultures, being transmitted and practiced among socially connected primates through cultural learning. Many famous researchers, such as Charles Darwin in his book The Descent of Man, mentioned tool-use in monkeys (such as baboons).

Among other mammals, both wild and captive elephants are known to create tools using their trunks and feet, mainly for swatting flies, scratching, plugging up waterholes that they have dug (to close them up again so the water doesn't evaporate), and reaching food that is out of reach. In addition to primates and elephants, many other social mammals particularly have been observed engaging in tool-use. A group of dolphins in Shark Bay uses sea sponges to protect their beaks while foraging. Sea otters will use rocks or other hard objects to dislodge food (such as abalone) and break open shellfish. Many or most mammals of the order Carnivora have been observed using tools, often to trap or break open the shells of prey, as well as for scratching.

Corvids (such as crows, ravens and rooks) are well known for their large brains (among birds) and tool use. New Caledonian crows are among the only animals that create their own tools. They mainly manufacture probes out of twigs and wood (and sometimes metal wire) to catch or impale larvae. Tool use in some birds may be best exemplified in nest intricacy. Tailorbirds manufacture 'pouches' to make their nests in. Some birds, such as weaver birds, build complex nests utilizing a diverse array of objects and materials, many of which are specifically chosen by certain birds for their unique qualities. Woodpecker finches insert twigs into trees in order to catch or impale larvae. Parrots may use tools to wedge nuts so that they can crack open the outer shell of nuts without launching away the inner contents. Some birds take advantage of human activity, such as carrion crows in Japan, which drop nuts in front of cars to crack them open.

Several species of fish use tools to hunt and crack open shellfish, extract food that is out of reach, or clear an area for nesting. Among cephalopods (and perhaps uniquely or to an extent unobserved among invertebrates), octopuses are known to utilize tools relatively frequently, such as gathering coconut shells to create a shelter or using rocks to create barriers.


Abstract

Spectacular progress in molecular biology, genome-sequencing projects and genomics makes this an appropriate time to attempt a comprehensive understanding of the molecular basis of social life. Promising results have already been obtained in identifying genes that influence animal social behaviour and genes that are implicated in social evolution. These findings — derived from an eclectic mix of species that show varying levels of sociality — provide the foundation for the integration of molecular biology, genomics, neuroscience, behavioural biology and evolutionary biology that is necessary for this endeavour.


Comparative transcriptomic analysis of the mechanisms underpinning ageing and fecundity in social insects

The exceptional longevity of social insect queens despite their lifelong high fecundity remains poorly understood in ageing biology. To gain insights into the mechanisms that might underlie ageing in social insects, we compared gene expression patterns between young and old castes (both queens and workers) across different lineages of social insects (two termite, two bee and two ant species). After global analyses, we paid particular attention to genes of the insulin/insulin-like growth factor 1 signalling (IIS)/target of rapamycin (TOR)/juvenile hormone (JH) network, which is well known to regulate lifespan and the trade-off between reproduction and somatic maintenance in solitary insects. Our results reveal a major role of the downstream components and target genes of this network (e.g. JH signalling, vitellogenins, major royal jelly proteins and immune genes) in affecting ageing and the caste-specific physiology of social insects, but an apparently lesser role of the upstream IIS/TOR signalling components. Together with a growing appreciation of the importance of such downstream targets, this leads us to propose the TI–J–LiFe (TOR/IIS–JH–Lifespan and Fecundity) network as a conceptual framework for understanding the mechanisms of ageing and fecundity in social insects and beyond.

This article is part of the theme issue ‘Ageing and sociality: why, when and how does sociality change ageing patterns?’

1. Introduction

Why do organisms age? This is a major question in evolutionary biology, given that an unlimited lifespan associated with continuous reproduction would increase fitness and hence should be favoured. The classical evolutionary theory of ageing, developed by Medawar, Williams and Hamilton [1–3], has, in principle, explained why ageing evolves. However, we still understand very little about the tremendous diversity of ageing rates among organisms and the mechanisms that might underlie this diversity [4] (reviewed in [5,6]).

During the last decades, results from model organisms have revealed the existence of a conserved set of gene networks and pathways involved in ageing in animals ranging from nematodes and flies to mice and humans (see [6–20], and references therein). In many insects, for example, the insulin/insulin-like growth factor 1 signalling (IIS)/target of rapamycin (TOR)/juvenile hormone (JH) network has emerged as a key regulator of lifespan and somatic maintenance, growth and fecundity, and explains trade-offs between these processes (figure 1). The IIS and TOR pathways sense the availability of nutrients, such as carbohydrates and amino acids. Through a cascade of signalling activities, they positively affect the production of the lipophilic sesquiterpenoid hormone JH (as well as the steroid hormone 20-hydroxy-ecdysone) and regulate various physiological processes including reproductive physiology (e.g. egg maturation, by affecting the expression of yolk proteins or the yolk precursor protein vitellogenin see [13–19]), somatic maintenance (e.g. humoral innate immunity and oxidative stress resistance) and lifespan (see reviews in [6–20] and references therein). In particular, results from the fruit fly Drosophila melanogaster as well as from other relatively short-lived insects (e.g. grasshoppers, butterflies, bugs and planthoppers) suggest that downregulation of this signalling network (e.g. via experimental ablation of insulin-producing cells or of the gland that produces JH) promotes somatic maintenance and longevity at the expense of fecundity (e.g. [7,13,15–17,20] and references therein). Because of its central role in modulating insect life history and ageing, we herein refer to this integrated network and the downstream processes that it affects as the TI–J–LiFe network (TOR/IIS–JH–Lifespan and Fecundity) (figure 1).

Figure 1. The ‘TI–J–LiFe’ network. The TI–J–LiFe network represents a set of interacting pathways that comprise the nutrient sensing TOR (target of rapamycin) and IIS (insulin/insulin-like growth factor 1 signalling) pathways, the Juvenile Hormone (JH, a major lipophilic hormone whose production is regulated by IIS and TOR), as well as downstream processes targeted by this network, including somatic maintenance functions (e.g. immunity and oxidative stress resistance) and reproductive physiology (including vitellogenins and yolk proteins), that have profound effects upon insect life history, especially on Lifespan and Fecundity. This network is thought to be one of the major regulatory circuits underpinning variation of insect lifespan and the trade-off between fecundity and longevity. The core components and feedback loops depicted here are mainly based on experimental findings in Drosophila melanogaster (for detailed information, see https://flybase.org e.g. IIS gene lists at: https://flybase.org/reports/FBgg0000904.html https://flybase.org/reports/FBgg0000900.html https://flybase.org/reports/FBgg0000898.html). Previous work suggests that this network and its effects are evolutionarily highly conserved among insects beyond Drosophila. In some social insects (e.g. Apis mellifera), some parts of this network might be ‘wired’ differently, but whether such a ‘rewiring’ is common among social insects remains largely unknown (for further discussion, see [18]). (Online version in colour.)

Considerably less is known, however, about the role of this signalling system in affecting ageing of social insects in which queens have extraordinarily long lifespans of up to several decades and that seemingly defy the commonly observed trade-off between fecundity and longevity [21–24]. Social insects (termites and ants as well as some bees and wasps) are further characterized by a reproductive division of labour: within a colony, the typically long-lived queens (and in termites, also kings) are the only reproducing individuals, while the other colony members (workers and sometimes soldiers) perform all non-reproductive tasks, such as foraging, brood care and defence, and are comparatively short-lived. Thus, as is the case in long-lived social mole-rats [25,26], reproductive individuals with exceptionally long lifespans (queens) have evolved in social insects. The convergent evolution of sociality and reproductive division of labour (‘castes’, comprising reproductives, workers and sometimes soldiers) appear to be associated with selection for long lifespans in reproductives (see also [21,24,27]). This calls for investigation of the convergent, or possibly parallel, evolution of the mechanisms underlying a long lifespan in reproductives.

Social animals are especially suited for ageing studies because both short- and long-lived phenotypes are encoded by the same genome within a colony (e.g. [17,24,28] and references therein). Indeed, outside social insects and mole-rats, such extreme (and in this case phenotypically plastic) differences in lifespan are only found in a few, distantly related taxa (e.g. [4]), which makes controlled comparisons difficult. The shared genetic background among castes within a colony furthermore means that caste-associated differences in longevity are generally not the result of genetic variation among individuals but are due to differences in gene expression. Transcriptomic studies of social insects therefore hold great promise for uncovering the physiological mechanisms underlying large differences in lifespan (e.g. [22,28,29]). To date, however, most such studies have focused on single species and not leveraged the potential power of comparative transcriptomics across taxa.

Here, we have examined the mechanisms underlying ageing in social insects by comparing gene expression patterns between young and old queens (and for termites, also kings) and workers across different social insect lineages: two termite (Blattodea, Isoptera), two bee (Hymenoptera, Apoidea) and two ant species (Hymenoptera, Formicidae) (for species and lifespan characteristics, see table 1). We studied patterns of life history and ageing of these species comparatively within a collaborative framework, the ‘So-Long’ consortium (www.so-long.org). This consortium tackles major questions about the apparent ‘reversal’ of the fecundity–longevity trade-off in the context of insect sociality by using species of different social complexity for each lineage and applying standardized methods when technically feasible. However, major biological differences among the species studied by our consortium sometimes necessitated the use of, for example, different tissues for transcriptomic analysis since the amount and quality of tissue that could be obtained constrained our use of specific tissues. In brief, we employed gene expression data derived from transcriptomes of target species to identify putative differences and commonalities in ageing-related expression patterns across three social insect lineages, with a special focus on the TI–J–LiFe network (figure 1 electronic supplementary material, §S1.0 and table S1). By comparing our results with published work from the well-established ageing model D. melanogaster, we begin to uncover how long-lived social insects might differ in their molecular underpinning of ageing and life-history traits when compared with short-lived solitary insects.

Table 1. Overview of samples included in this study.

a In [30], the same samples were referred to as ‘head’, yet the prothorax was attached to the head.


Contents

Reproductives [ edit | edit source ]

Fertile termite queen (Coptotermes formosanus), showing ovary-filled, distended abdomen. The rest of its body is the same size as that of a worker.

A female that has flown, mated, and is producing eggs is called a "queen." Similarly, a male that has flown, mated, and is in proximity to a queen is termed a "king." Research using genetic techniques to determine relatedness of colony members has shown that the original idea that colonies are only ever headed by a monogamous royal pair is wrong. Multiple pairs of reproductives within a colony are commonly encountered. In the families Rhinotermitidae and Termitidae, and possibly others, sperm competition does not seem to occur (male genitalia are very simple and the sperm are anucleate), suggesting that only one male (king) generally mates within the colony.

At maturity, a primary queen has a great capacity to lay eggs. In physogastric species, the queen adds an extra set of ovaries with each molt, resulting in a greatly distended abdomen and increased fecundity, often reported to reach a production of more than 2,000 eggs a day. The distended abdomen increases the queen's body length to several times more than before mating and reduces her ability to move freely, though attendant workers provide assistance. The queen is widely believed to be a primary source of pheromones useful in colony integration, and these are thought to be spread through shared feeding (trophallaxis).

The king grows only slightly larger after initial mating and continues to mate with the queen for life (a termite queen can live for forty-five years). This is very different from ant colonies, in which a queen mates once with the male(s) and stores the gametes for life, as the male ants die shortly after mating.

Two termites in the process of shedding their wings after mating. Maun, Botswana.

The winged (or "alate'") caste, also referred to as the reproductive caste, are generally the only termites with well-developed eyes, although workers of some harvesting species do have well-developed compound eyes, and, in other species, soldiers with eyes occasionally appear. Termites on the path to becoming alates (going through incomplete metamorphosis) form a subcaste in certain species of termites, functioning as workers ("pseudergates") and also as potential supplementary reproductives. Supplementaries have the ability to replace a dead primary reproductive and, at least in some species, several are recruited once a primary queen is lost.

In areas with a distinct dry season, the alates leave the nest in large swarms after the first good soaking rain of the rainy season. In other regions, flights may occur throughout the year, or more commonly, in the spring and autumn. Termites are relatively poor fliers and are readily blown downwind in wind speeds of less than 2 km/h, shedding their wings soon after landing at an acceptable site, where they mate and attempt to form a nest in damp timber or earth.

Workers [ edit | edit source ]

Worker termites undertake the labors of foraging, food storage, brood and nest maintenance, and some defense duties in certain species. Workers are the main caste in the colony for the digestion of cellulose in food and are the most likely to be found in infested wood. This is achieved in one of two ways. In all termite families except the Termitidae, there are flagellate protists in the gut that assist in cellulose digestion. [citation needed] However, in the Termitidae, which account for approximately 60 percent of all termite species, the flagellates have been lost and this digestive role is taken up, in part, by a consortium of prokaryotic organisms. This simple story, which has been in entomology textbooks for decades, is complicated by the finding that all studied termites can produce their own cellulase enzymes, and therefore might digest wood in the absence of their symbiotic microbes although there is now evidence suggesting that these gut microbes make use of termite-produced cellulase enzymes. [citation needed] Ώ] Our knowledge of the relationships between the microbial and termite parts of their digestion is still rudimentary. What is true in all termite species, however, is that the workers feed the other members of the colony with substances derived from the digestion of plant material, either from the mouth or anus. This process of feeding of one colony member by another is known as trophallaxis and is one of the keys to the success of the group. It frees the parents from feeding all but the first generation of offspring, allowing for the group to grow much larger and ensuring that the necessary gut symbionts are transferred from one generation to another. Some termite species do not have a true worker caste, instead relying on nymphs that perform the same work without differentiating as a separate caste. [citation needed]

Soldiers [ edit | edit source ]

A picture of a soldier termite (Macrotermitinae) with an enlarged jaw in the Okavango Delta.

The soldier caste has anatomical and behavioural specializations, providing strength and armour which are primarily useful against ant attack. The proportion of soldiers within a colony varies both within and among species. Many soldiers have jaws so enlarged that they cannot feed themselves, but instead, like juveniles, are fed by workers. The pantropical subfamily Nasutitermitinae have soldiers with the ability to exude noxious liquids through either a horn-like nozzle (nasus). Simple holes in the forehead called "fontanelles" and which exude defensive secretions are a feature of the family Rhinotermitidae. Many species are readily identified using the characteristics of the soldiers' heads, mandibles, or nasus. Among the drywood termites, a soldier's globular ("phragmotic") head can be used to block their narrow tunnels. Termite soldiers are usually blind, but in some families, particularly among the dampwood termites, soldiers developing from the reproductive line may have at least partly functional eyes.

The specialization of the soldier caste is principally a defence against predation by ants. The wide range of jaw types and phragmotic heads provides methods that effectively block narrow termite tunnels against ant entry. A tunnel-blocking soldier can rebuff attacks from many ants. Usually more soldiers stand by behind the initial soldier so once the first one falls another soldier will take the place. In cases where the intrusion is coming from a breach that is larger than the soldier's head, defense requires special formations where soldiers form a phalanx-like formation around the breach and bite at intruders or exude toxins from the nasus or fontanelle. This formation involves self-sacrifice because once the workers have repaired the breach during fighting, no return is provided, thus leading to the death of all defenders. Another form of self-sacrifice is performed by Southeast Asian tar-baby termites (Globitermes sulphureus). The soldiers of this species commit suicide by autothysis—rupturing a large gland just beneath the surface of their cuticle. The thick yellow fluid in the gland becomes very sticky on contact with the air, entangling ants or other insects who are trying to invade the nest. ΐ] Α]

Termites undergo incomplete metamorphosis. Freshly hatched young appear as tiny termites that grow without significant morphological changes (other than wings and soldier specializations). Some species of termite have dimorphic soldiers (up to three times the size of smaller soldiers). Though their value is unknown, speculation is that they may function as an elite class that defends only the inner tunnels of the mound. Evidence for this is that, even when provoked, these large soldiers do not defend themselves but retreat deeper into the mound. On the other hand, dimorphic soldiers are common in some Australian species of Schedorhinotermes that neither build mounds nor appear to maintain complex nest structures. Some termite taxa are without soldiers perhaps the best known of these are in the Apicotermitinae.

Diet [ edit | edit source ]

Termites are generally grouped according to their feeding behaviour. Thus, the commonly used general groupings are subterranean, soil-feeding, drywood, dampwood, and grass-eating. Of these, subterraneans and drywoods are primarily responsible for damage to human-made structures.

All termites eat cellulose in its various forms as plant fibre. Cellulose is a rich energy source (as demonstrated by the amount of energy released when wood is burned), but remains difficult to digest. Termites rely primarily upon symbiotic protozoa (metamonads) such as Trichonympha, and other microbes in their gut to digest the cellulose for them and absorb the end products for their own use. Gut protozoa, such as Trichonympha, in turn rely on symbiotic bacteria embedded on their surfaces to produce some of the necessary digestive enzymes. This relationship is one of the finest examples of mutualism among animals. Most so-called higher termites, especially in the Family Termitidae, can produce their own cellulase enzymes. However, they still retain a rich gut fauna and primarily rely upon the bacteria. Owing to closely related bacterial species, it is strongly presumed that the termites' gut flora are descended from the gut flora of the ancestral wood-eating cockroaches, like those of the genus Cryptocercus.

Some species of termite practice fungiculture. They maintain a “garden” of specialized fungi of genus Termitomyces, which are nourished by the excrement of the insects. When the fungi are eaten, their spores pass undamaged through the intestines of the termites to complete the cycle by germinating in the fresh faecal pellets. Β] Γ] They are also well known for eating smaller insects in a last resort environment.

In captivity [ edit | edit source ]

Few zoos hold termites, due to the difficulty in keeping them captive and the reluctance of authorities to permit potential pests. One of them is Zoo Basel in Switzerland. At Zoo Basel, two African termite (Macrotermes bellicosus) populations exist and thrive - resulting in very rare (in captivity) mass migrations of young flying termites. This happened last in September 2008, when thousands of male termites left their mound each night, died, and covered the floors and water pits of the house their exhibit is in. Δ]


Materials and methods

Materials

We chose three trees of Ficus hispida in Danzhou (19° 30’ N, 109° 31’ E), Hainan province, China for inoculation of the fig pollinator species (Ceratosolen solmsi). Samples for genomic DNA extraction were collected from June to August in 2010. In each inoculation experiment, naturally growing figs were covered with mesh bags from their very early developmental stages to exclude all insects, including fig wasps. One mated female (‘mother’) fig wasp was introduced into each bagged fig to lay eggs. About 1 month later, when the offspring were mature, we transferred several mated female daughter fig wasps into other bagged figs at their receptive stages. Following this second generation of development, we collected all male grandson fig wasps for genome sequencing. In fig wasps, like other hymenopterans, males are haploid and provide better targets than diploid females for genome sequencing projects. These processes reduced genomic heterozygosity and, thus, improved the quality of assembly. After thoroughly washing with double-distilled water, we immediately froze the samples in liquid nitrogen and transported them to the laboratory on dry ice. DNA extraction occurred immediately on arrival.

The following groups of samples were selected for transcriptome analyses: (1) larval female-16th day (larva), (2) larval male-16th day (larva), (3) pupal female-21th day (early pupa), (4) pupal male-21th day (early pupa), (5) pupal female-25th day (late pupa), (6) pupal male-25th day (late pupa), (7) adult female-29th day, and (8) adult male-29th day. The days indicate time since the eggs were laid. For sample groups 1 to 6, we used only one fig wasp for each RNA extraction. For each adult group, we used 50 individuals. All RNA extractions occurred immediately after collection.

DNA extraction

About 500 male fig wasps were divided into 10 samples of 50 individuals each and used for DNA extraction using a method modified from the protocol developed by J. Rehm, in the Berkeley Drosophila Genome Project [48]. Briefly, 50 fig wasps were completely homogenized in 400 μL Buffer A (100 mM Tris–HCl, pH 7.5 500 mM EDTA 100 mM NaCl 0.5% SDS). A total of 3 μL RNaseA was added to the homogenate, followed by 2 h incubation at 37°C. Then 3 μL Proteinase K was added to the mixture, followed by 2 h incubation at 58°C. Next, 800 μL of LiCL/KAc solution (5 M KAc and 6 M LiCl) was added before the tube was incubated on ice for 10 min. Then the mixture was centrifuged at 14,000 g for 15 min at 4°C, and 1 mL of supernatant was transferred into a new 2 mL tube. To precipitate the genomic DNA from the supernatant, 0.8 mL ice isopropanol was added, and centrifuged at 14,000 g for 15 min at 4°C. The supernatant was then aspirated. The DNA pellet was washed with 70% ethanol, followed by drying for 5 min. The DNA was dissolved in 50 μL of TE buffer and stored at −80°C.

RNA extraction

Total RNA was isolated using the RNeasy® Micro Kit (Qiagen, Shanghai, China) and treated with DNase (Qiagen, Shanghai, China). A NanoDrop ND-1000 Spectrophotometer (Nano-Drop Technologies, Wilmington, DE, USA) was used to confirm adequate RNA concentration and A260/A280 ratio. RNA was dissolved in 20 μL RNase-free water and kept at −80°C. Larval females and males that had no distinct morphological divergence were discriminated by the variable splicing pattern of the sex determination gene doublesex[49]. The procedure used 50 ng dissolved RNA of larva fig wasp to synthesize first-strand cDNA by priming with oligo(dT) with TransScript® II First-Strand cDNA Synthesis SuperMix (TransGen Biotech, Beijing, China). The sex of larva individual was then confirmed by PCR of the male-specific splice isoform of doublesex.

Shotgun libraries construction and sequencing

Genomic DNA was sheared into fragments and seven libraries were constructed with inserted fragment sizes ranging from 200 bp, 500 bp, 800 bp, 2 kb, 5 kb, and 10 kb to 20 kb by the manufacturer’s library kit (Illumina) [50]. A PCR-free library was also constructed. The libraries were sequenced using the Illumina-HiSeq™ 2000 platform with paired-end sequencing approaches.

For RNA-seq, beads with oligo(dT) were used to isolate poly(A) mRNA. Fragmentation buffer was then added for cutting mRNA into short fragments, which were used as templates. Random hexamer primers were used to synthesize first-strand cDNA. Second-strand cDNA was synthesized using a mixture of buffer, dNTPs, RNase H, and DNA polymerase I. Short fragments were purified with QiaQuick PCR extraction kits and resolved with EB buffer for end repair and addition of poly(A). Next, the short fragments were connected with sequencing adaptors. For amplification with PCR, we selected suitable fragments as templates based on agarose gel electrophoresis. Finally, the libraries were sequenced using an Illumina HiSeq™ 2000. RNA-seq for abdomen of adult Drosophila willistoni was downloaded and compared between female and male [51].

Genome assembly

We used SOAPdenovo (version 2.01) to assemble the genome with the following procedures (basic information in Table 1 details are referred to the giant panda [50]):

construct contig: split the short-insert size library data into 43-mers and construct a de Bruijn graph. Next, obtain parameters from a simplified graph. Finally, connect the 43-mer path to get contigs

construct scaffold: realign all usable reads onto contig sequences, then calculate the amount of shared paired-end relationships between each pair of contigs, weight the rate of consistent and conflicting paired-ends, and then construct scaffolds

fill gaps: use the paired-end information from the short insert size library and the PCR-free library to retrieve read-pairs that had one end mapped to a unique contig and the other end located in the gap region. We then carried out a local assembly of the collected reads to fill the gaps using Gapcloser.

To evaluate the assembly, we employed CEGMA and EST evaluations. For CEGMA (version 2.4) [19] evaluation, we used 248 ultra-conserved core eukaryotic genes (CEGs) that were widely distributed and conserved in species to assess the completeness of genome assembly and gene-set. The CEGMA evaluation utilized several software packages, including tblastn (blast-2.2.25), genewise (wise2.2.3), hmmer (hmmer-3.0), and geneid (geneid v1.4). Four insect genomes including C. solmsi were compared. For the EST evaluation, we used BLAT[52] to map Sanger-sequenced ESTs or Trinity [53] assembled tgicl [54] clustered unigenes to the C. solmsi genome assembly. Then we calculated genome coverage using both the percentage of bases covered by ESTs and the percentage of scaffold numbers with >90% or 50% covered by ESTs. Eight transcriptome datasets were assembled separately by Trinity and then clustered to remove redundancy by tgicl to get unigene sequences before evaluation.

Gene prediction

Multiple approaches were used to predict gene structures in this genome including de novo, homology-based, EST and RNA-seq based predictions. De novo prediction was performed based on the repeat-masked genome and with the help of the HMM model using AUGUSTUS [55], GENSCAN, and SNAP. Homologous proteins of the following species were mapped to the genome using tblastn with an E-value cutoff 1e-5: Homo sapiens (H. sap): data downloaded from [56] Apis mellifera (A. mel): data downloaded from [57]: Bombyx mori (B. mor): data downloaded from [58] Drosophila melanogaster (D. mel): data downloaded from [59] and Nasonia vitripennis (N. vit): data downloaded from [60]. The aligned sequences, as well as their corresponding query proteins, were then filtered and passed to GeneWise[61] to search for accurately spliced alignments. ESTs (unpublished data in our lab) were aligned to the genome using BLAT[52] to generate spliced alignments. The alignments were then linked according to overlap using PASA. Source evidence generated from the above three approaches was integrated by GLEAN to produce a consensus gene set.

To improve the integrity and correctness of the genome, transcriptome reads were aligned against the genome using TopHat to identify candidate exon regions and the donor and acceptor sites. Cufflinks[62] was employed to assemble the alignments into transcripts. Then, based on assembled candidate transcript sequences, ORFs were predicted to get reliable transcripts by using HMM-based training parameters. Finally, we combined the GLEAN set with the transcripts from RNA-seq to generate a confident gene set.

Gene function annotation

Gene functions were assigned according to the best match of the alignments based on blastp to the databases SwissProt (release2011 01) [63] and TrEMBL (release2011 01). The motifs and domains of genes were determined by InterProScan (iprscan 4.7) [64] against protein databases such as ProDom, PRINTS, Pfam, SMART, PANTHER, and PROSITE. Gene Ontology (GO) [65] IDs for each gene were obtained from the corresponding InterPro entries. All genes were aligned against KEGG (release54) [66] proteins, and the pathway in which the gene might be involved was inferred from matched genes.

Annotation of repeats and non-coding RNA

Initially, non-interspersed repetitive regions (including simple repeats, satellites, and low complexity repeats) were predicted by RepeatMasker [67] with the ‘-noint’ option. These tandem repeats were also annotated using Tandem Repeats Finder (v.4.04) with parameters of ‘Match = 2, Mismatch = 7, Delta = 7, PM = 80, PI = 10, Minscore = 50, and MaxPeriod = 2000’ [68].

Implementing a homology strategy, we identified known transposable elements (TEs) against the Repbase database (v.20120418) in the genome of C. solmsi using RepeatMasker v.open-3.3.0 (ab-blast engine [69], with parameters ‘-nolow, -no_is -norna, -parallel 1 -s’) [67] and RepeatProteinMask (with parameters ‘-noLowSimple, -pvalue 0.0001’) at the DNA and protein level, respectively [70].

A de novo repeat library was also generated using RepeatModeler (v1.0.5) [71] and PILER-DF [72], and a RepeatMasker analysis against the final non-redundant library was performed again to find homologs in the genome and to classify the found repeats.

We searched the whole genome sequence to detect four types of non-coding RNAs. Employment of tRNAscan-SE[73] identified reliable tRNA positions. We searched for small nuclear RNAs and microRNAs using a two-step method: sequences were aligned with blast and then searched with INFERNAL against the Rfam database (release 9.1) [74]. The rRNAs were found by aligning with blastn against a ref rRNA sequence from the closest related species.

Orthologous gene clusters and the phylogeny of arthropods

We identified gene families using TreeFam [75] and the following steps: first, blastp was used to compare all the protein sequences of 10 species including C. solmsi. The E-value threshold was set to 1e-7 second, HSP segments of each protein pair were concatenated by solar[50], H-scores were computed based on Bit-scores, and these were taken to evaluate the similarity between genes finally, gene families were identified by clustering of homologous gene sequences using hcluster_sg. Genes specific to C. solmsi were those that did not cluster with other arthropods chosen for gene family construction, and those that did not have homologs in the predicted gene repertoire of the compared genomes (Figure 3). However, these genes could have GO annotation if they had the functional motifs. The motifs and domains of these genes were determined by InterProScan (iprscan 4.7) [64] against protein databases such as ProDom, PRINTS, Pfam, SMART, PANTHER, and PROSITE. GO IDs of each gene were obtained from the corresponding InterPro entries, from which we also obtained gene functional enrichment.

Single-copy orthologous genes were used to reconstruct the phylogeny. CDS sequences from each gene were aligned using MUSCLE and protein sequences were concatenated to form one super gene for each species. Codon position 2 of aligned CDS sequences was extracted for subsequent analysis. PhyML [76] was used to construct the phylogeny using the GTR substitution model and gamma distribution rates model across sites. Branch reliability was assessed via aLRT values.

Divergence times were estimated using PAML mcmctree[77] while implementing the approximate likelihood calculation method. Gamma prior and alpha parameters were computed based on the substitution rate per time unit estimated by PAML baseml. We ran mcmctree to sample 10,000 times, with sampling frequency set to 5,000, and burnin parameter set to 5,000,000 using correlated molecular clock and REV substitution model. Finetune parameters were set to make acceptance proportions fall in range (0.2, 0.4). Other parameters were the defaults. Convergence of results was checked by Tracer[78] and two independent runs were performed to confirm convergence.

Gene family expansion and contraction

We identified gene families using CAFE [79], which employed a random birth and death model to study gene gains and losses in gene families across a user-specified phylogeny. The global parameter λ, which described both the gene birth (λ) and death (μ = −λ) rate across all branches in the tree for all gene families, was estimated using maximum likelihood. A conditional P value was calculated for each gene family, and families with conditional P values less than threshold (0.0001) were considered as having an accelerated rate of gain or loss. We identified branches responsible for low overall P values of significant families.

Evolutionary rates of genes

We calculated ka/ks ratios for all single copy orthologs of C. solmsi, Nasonia vitripennis, Apis mellifera, Camponotus floridanus, and Tribolium castaneum. Alignment quality was essential for estimating positive selection. Thus, orthologous genes were first aligned by PRANK [80], a good alignment tool for studies of molecular evolution. We used Gblocks [81] to remove ambiguously aligned blocks within PRANK alignments. We employed ‘codeml’ in the PAML package [23] with the free-ratio model to estimate Ka, Ks, and Ka/Ks ratios on different branches. The difference in mean Ka/Ks ratios for single-copy orthologous genes between C. solmsi and each of the other species were compared with paired Wilcoxon rank sum tests.

Genes that showed values of Ka/Ks higher than 1 along the branch leading to C. solmsi were reanalyzed using the codon based branch-site tests implemented in PAML [23, 82]. The branch-site model, which allowed ω to vary both among sites in the protein and across branches, was used to detect episodic positive selection. Test 1 (M1a vs. branch site model) and test 2 (branch site null model vs. branch site model), which differentiated positive selection from the relaxation of selective constrains, were used. The pairwise comparisons M1a vs. branch-site model, and branch-site model (model = 2, NSsites = 2) vs. branch-site null model (fixed ω = 1 and ω = 1) were used to perform likelihood ratio tests (LRTs). Their significance was evaluated using a χ2 distribution. When the LRT was significant, a Bayes Empirical Bayes (BEB) analysis was conducted to identify putatively positively selected sites, which may also be relaxed selected sites though.

Manual annotation and evolutionary analyses of interested genes

For genes requiring greater annotation, protein homologs of N. vitripennis, A. mellifera, and sometimes D. melanogaster were collected from NCBI, Hymenoptera Genome Database [83], and FlyBase [84]. Both tblastn and blastp searches were performed for candidate genes in the assembled genome of C. solmsi. The annotated protein set used an E-threshold 0.005. The threshold was raised when protein sequences were short and few blast hits were found. A blast of candidate genes to the NCBI non-redundant (nr) protein database confirmed their orthology. IGV browser was used to view gene annotations, EST, and RNA-seq BAM alignments in the genome of C. solmsi. Gene models were refined manually according to RNA evidence and tblastn results conducted with the assistance of custom perl scripts. Pseudogenes and irregular features such as missing start codons, stop codons, and other anomalies were noted. For annotation of cuticular proteins with an R&R consensus, the ‘chitin_bind_4’ domain was required. Proteins containing cysteines were removed unless the cysteines lay in signal peptide regions, which were identified by SignalP [85]. For P450s, gene models for C. solmsi were searched by tblastn and blastp against D. melanogaster, A. mellifera, and N. vitripennis CYP sequences representing the CYP2, 3, 4 and mitochondrial P450 clans (E-value cutoff = 10 -4 ). All models with predicted proteins that included the canonical heme binding sequence were manually verified for the presence of the other key features of P450 enzymes the gene model was corrected whenever necessary (incorrect predictions such as fusions with adjacent genes or fragmentation) or possible (when RNA-seq sequences were available). Final gene models were confirmed by blasting back to the reference gene set to confirm reciprocal best hits. The obtained gene models were inspected and, if necessary, edited. Care was taken to ensure that the predicted gene structures matched corresponding transcriptomic data. Pseudogenes and gene fragments (detritus exons) were separated from putative full-length CYP coding sequences. To annotate genes involved in the development of eyes and wings, proteins putatively participating in development of eyes and wings described for D. melanogaster were used as query sequences. These queries were used in blastp and tblastn searches (E-value cutoff = 10 -4 ) against the protein predictions and scaffolds of the C. solmsi genome. Iterative searches were also conducted with each new protein of C. solmsi as a query until no new genes were identified in each major subfamily or lineage.

To further understand the evolutionary history and homologies between gene families of C. solmsi and other insects, we performed a phylogenetic analysis using the genes found in C. solmsi and some other insect taxa with completed genomes: A. mellifera, N. vitripennis, and D. melanogaster. The amino acid sequences of homologous genes were aligned with ClustalX v2.0 [86]. ProtTest [87] identified evolutionary models that best fit this dataset according to the Akaike Information Criterion. A maximum likelihood tree was then reconstructed with PhyML 3.0 using the best-fit model with a gamma correction using four discrete classes, an estimated alpha parameter and proportion of invariable sites [76]. Node support values were obtained by the rapid bootstrap algorithm as implemented in PhyML 3.0 (100 replicates). Some tree images were created using the iTOL web server [88]. Gray circles on branches were used to indicate bootstrap values >80% from 100 bootstrap replicates.

We tested for selection on Gustatory receptor (Gr) and Olfactory receptor (Or) genes. The calculation of each Ka/Ks value of Gr or Or gene were based on each orthologous group of Gr or Or gene members among A. mellifera, N. vitripennis, and C. solmsi. The difference between mean Ka/Ks of Or plus Gr genes and all single-copy genes was compared using paired Wilcoxon rank sum tests. This determined if Or and Gr genes underwent different selective pressures than single-copy genes.

Validation of extremely contracted CCE, OBP, and Gr gene families

Since the fig wasp had many fewer genes than other insect species, we performed additional analyses to confirm that the absence of genes is not due to a poor or incomplete assembly or inadequate annotation. Validation of assembly quality was fully analyzed and described above (see genome assembly). We also tried to confirm the absence of genes by focusing on the annotations of the gene families CCE, OBP, and Gr, in which the fig wasp had the fewest gene members among the studied insects, by delving into the raw genomic reads.

We chose the library with the inserted fragment size of 500 bp (altogether 68,549,132 pair end reads after correction), which was 25× coverage of the genome, and aligned it to the assembled genome of C. solmsi by SOAP [89] (default except: -v, 5 -g, 3). Altogether, 91.2% of reads mapped to the genome. We then compared the unmapped reads (= 6,042,453) to the protein sequences of all CCE, OBP, and Gr members of the fig wasp, jewel wasp, and honeybee by blastx[69] (references given in Table 2 and Additional file 1: Table S10) with e-value of 1e-5. Only one read mapped to gene AmGr9 (a Gr gene member in the honeybee), which indicated that the unmapped reads scarcely had any similar sequences in the three gene families. Thus, the absence of genes in these gene families was not due to incomplete annotation.

Detection of horizontal gene transfer

Two independent approaches were used to identify possible HGT events. The first used gene models. We used blastp (E-value cutoff 10 -10 and a continuous overlap threshold of 33%) to compare the predicted protein sequences of C. solmsi against sequences in the RefSeq and nr databases to exclude unique genes and those with high similarity to other insects. Next, we constructed phylogenies for the retained proteins with highest similarity to non-insects. Multiple alignments were performed by using ClustalW2 [86], followed by manual refinement. Phylogenetic analyses were conducted using maximum likelihood (ML) and Bayesian inference (BI). A distance-based phenogram was also constructed using neighbor joining (NJ). ML trees were estimated by PhyML [76] using best-fit substitution model estimated by Prottest 3.0 [87]. In all cases, we used a discrete gamma-distribution model with four rate-categories plus invariant positions. The gamma parameter and proportion of invariant sites were estimated from the data. Bootstrap branch-support values involved 1,000 pseudoreplicates. BI used MrBayes 3.1.2 [93]. For each HGT, we ran two independent analyses using four MCMC chains (one cold and three hot) for one million generations and stopped them when the average deviation of split frequencies fell well below 0.01. We sampled trees every 100 generations and discarded the initial 25% of the total trees as burn-in. Compatible groups were shown in the majority rule consensus tree. NJ trees were constructed by using Neighbor in Mega5 [94]. Bootstrap values were obtained by generating 1,000 pseudoreplicates. HGTs were detected by clustering non-related species on a well-supported node.

Using the scaffold sets of C. solmsi, we identified regions involved in recent HGT events between bacteria, fungi, plants, and other non-insects to C. solmsi. The pipeline involved blastn of C. solmsi against other non-insects. Usually, this approach has detected two categories of genes: candidate HGT events and highly conserved genes shared by non-insects and C. solmsi (not HGTs). Thus, the pipeline formed two categories based on the following certain criteria:

HGT_scaf: non-insects score > animal score AND scaffold length > 5 K AND range_per < 90%

highly_cons: best non-insects score < insect score.

The scaffolds with both non-insect and insect sequences served as evidence of non-contamination.

Data availability

Data for this Whole Genome Shotgun project were deposited at DDBJ/EMBL/GenBank under the accession no. ATAC00000000. The version described in this paper is version ATAC01000000. The transcriptome data reported in this paper were deposited in the National Center for Biotechnology Information Short Read Archive [95] under the accession no. SRP029703.


Published by the Royal Society. All rights reserved.

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Bug Reference

Want to find out more about common pests? Simply click on any of the bugs below, and read more information. If you have problems with any of these pests, The Bug Man can help you get rid of them for good!

Harmonia axyridis is a "typical" coccinellid beetle in shape and structure, being domed and having a "smooth" transition between its elytra (wing coverings), pronotum and head. It occurs in three main color forms: red or orange with black spots (known as form succinea) black with four red spots (form spectabilis) and black with two red spots (form conspicua). However, numerous intermediate and divergent forms have also been recorded. The species is typically large (7–8 mm long) and even more dome-shaped than native European species (these characteristics distinguish Harmonia axyridis from native species in the UK). It often has white markings (typically "M" or "W" shaped) on its pronotum, and usually brown or reddish legs.

This species was possibly established in North America as the result of introductions into the United States in an attempt to control the spread of aphids. Whatever the source, in the last two decades, this insect has spread throughout the United States and Canada and has been a prominent factor in controlling aphid populations. However, many people now view this species as a nuisance, partly due to their tendency to overwinter indoors and the unpleasant odor and stain left by their bodily fluid when frightened or squashed. (It is also currently increasing in Europe to the detriment of indigenous species, due to its voracious appetite which enables them to out-compete and even eat other lady beetles, as it also does in the United States.)

In the U.S., the first attempts to introduce it took place as far back as 1916. Repeated efforts were not successful. In the early 1980s, aphids were causing significant problems for growers of pecan trees, so the United States Department of Agriculture again attempted to bring the insect into the country—this time in the southeastern United States, using beetles brought from their native region in northeasternAsia. After a period of time, USDA scientists concluded that their attempts had been unsuccessful. However, a population of beetles was observed near New Orleans, Louisiana around 1988, though this may have been an accidental introduction event independent of the original, planned efforts. In the following years it quickly spread to other states, being occasionally observed in the Midwest within 5–7 years, and becoming common in the region by about 2000. The species was also established in the northwest by 1991, and the northeast by 1994, in the former case quite possibly involving additional introductions, rather than reaching there from the southeast. It is reported that it has heavily fed on soybean aphids (which recently appeared in the U.S. after coming from China), supposedly saving farmers vast sums of money in 2001. However, in addition to its household pest status, it has been reported to be a minor agricultural pest (contaminating crops of tender fruits and grapes) in Iowa, Ohio, New York State, and Ontario. The contamination of grapes by this beetle has been found to alter the taste of wine. Native ladybird species have experienced often dramatic declines in abundance in areas invaded by H. axyridis. Despite the troubles the Asian lady beetle causes, many farmers still view it as a beneficial insect.

The common bedbug (Cimex lectularius) is the species best adapted to human environments. It is found in temperate climates throughout the world and feeds on human blood. Adult bedbugs are reddish-brown, flattened, oval, and wingless, with microscopic hairs that give them a banded appearance. A common misconception is that they are not visible to the naked eye. Adults grow to 4–5 mm (1/8th – 3/16th of an inch) in length and do not move quickly enough to escape the notice of an attentive observer. Newly hatched nymphs are translucent, lighter in color and become browner as they moult and reach maturity. In size, they are often compared to lentils or apple seeds.

Feeding habits
Bedbugs are generally active just before dawn, with a peak feeding period about an hour before sunrise. However, they may attempt to feed at other times, given the opportunity, and have been observed to feed at any time of the day. They climb the walls to the ceiling and jump down on feeling a heat wave (in wooden houses). Attracted by warmth and the presence of carbon dioxide, the bug pierces the skin of its host with two hollow tubes. With one tube it injects its saliva, which containsanticoagulants and anesthetics, while with the other it withdraws the blood of its host. After feeding for about five minutes, the bug returns to its hiding place. The bites cannot usually be felt until some minutes or hours later, as a dermatological reaction to the injected agents, and the first indication of a bite usually comes from the desire to scratch the bite site. Because of their dislike for sunlight, bedbugs come out at night.

Although bedbugs can live for a year or as much as eighteen months without feeding, they typically seek blood every five to ten days. Bedbugs that go dormant for lack of food often live longer than a year, well-fed specimens typically live six to nine months. Low infestations may be difficult to detect, and it is not unusual for the victim not to even realize they have bedbugs early on. Patterns of bites in a row or a cluster are typical as they may be disturbed while feeding. Bites may be found in a variety of places on the body.

Bedbugs have been erroneously associated with filth in the mistaken notion that this attracts them. Bedbugs are attracted by exhaled carbon dioxide and body heat, not by dirt, and they feed on blood, not waste. In short, the cleanliness of their environment has an effect on the control of bedbugs but, unlike cockroaches, does not have a direct effect on bedbugs as they feed on their hosts and not on waste. Good housekeeping in association with proper preparation and mechanical removal by vacuuming will certainly assist in control.

Bites
In most observed cases, bites consist of a raised red bump or flat welt, and are often accompanied by intense itching. The red bump or welts are the result of an allergic reaction to the anesthetic contained in the bedbug's saliva, which is inserted into the blood of its victim. Bedbug bites may appear indistinguishable from mosquito bites, though they tend to last for longer periods. Bites may not become immediately visible, and can take up to nine days to appear. Bedbug bites tend to not have a red dot in the center such as is characteristic of flea bites. A trait shared with flea bites, however, is tendency towards arrangements of sequential bites. Bites are often aligned three in a row, giving rise to the colloquialism "breakfast, lunch and dinner." This may be caused by the bedbug being disturbed while eating, and relocating half an inch or so farther along the skin before resuming feeding. Alternatively, the arrangement of bites may be caused by the bedbug repeatedly searching for a blood vein. People react very differently to bedbugs, and individual responses vary with factors including skin type, environment, and the species of bug. In some rare cases, allergic reactions to the bites may cause nausea and illness. In a large number of cases, estimated to 50% of all people, there is no visible sign of bites whatsoever, greatly increasing the difficulty of identifying and eradicating infestations. People commonly respond to bed bug infestations and their bites with anxiety, stress, and insomnia. Individuals may also get skin infections and scars from scratching the bedbug bite locations.

Infestations
here are several means by which dwellings can become infested with bedbugs. People can often acquire bedbugs at hotels, motels, or bed-and-breakfasts, and bring them back to their homes in their luggage. They also can pick them up by inadvertently bringing infested furniture or used clothing to their household. If someone is in a place that is severely infested, bedbugs may actually crawl onto and be carried by people's clothing, although this is atypical behaviour — except in the case of severe infestations, bedbugs are not usually carried from place to place by people on clothing they are currently wearing. Bedbugs may travel between units in multi-unit dwellings, such as condominiums and apartment buildings, after being originally brought into the building by one of the above routes. Bedbugs can also be transmitted via animal vectors including wild birds and household pets.

This spread between sites is dependent in part on the degree of infestation, on the material used to partition units and whether infested items are dragged through common areas while being disposed of, resulting in the shedding of bedbugs and bedbug eggs while being dragged. In some exceptional cases, the detection of bedbug hiding places can be aided by the use of dogs that have been trained to find the insects by their scent much as dogs are trained to find drugs or explosives. A trained dog and handler can detect and pinpoint a bedbug infestation within minutes. This is a fairly costly service that is not used in the majority of cases, but can be very useful in difficult cases.

The numerical size of a bedbug infestation is to some degree variable, as it is a function of the elapsed time from the initial infestation. With regards to the elapsed time from the initial infestation, even a single female bedbug brought into a home has a potential for reproduction, with its resulting offspring then breeding, resulting in a geometric progression of population expansion if control is not undertaken. Sometimes people are not aware of the insects and do not notice the bites. The visible bedbug infestation does not represent the infestation as a whole, as there may be infestations elsewhere in a home. However, the insects do have a tendency to stay close to their hosts, hence the name "bed" bugs.

Locations
Bedbugs travel easily and quickly along pipes and boards, and their bodies are very flat, which allows them to hide in tiny crevices. In the daytime, they tend to stay out of the light, preferring to remain hidden in such places as mattress seams, mattress interiors, bed frames, nearby furniture, carpeting, baseboards, inner walls, tiny wood holes, or bedroom clutter. Bedbugs can be found on their own, but more often congregate in groups. Bedbugs are capable of travelling as far as 100 feet to feed, but usually remain close to the host in bedrooms or on sofas where people may sleep.

Detection
Bedbugs are known for being elusive, transient, and nocturnal, making them difficult to detect. While individuals have the option of contacting a pest control professional to determine if a bedbug infestation exists, there are several do-it-yourself methods that may work equally well. The presence of bedbugs may be confirmed through identification of the insects collected or by a pattern of bites. Though bites can occur singularly, they often follow a distinctive linear pattern marking the paths of blood vessels running close to the surface of the skin. The common bite pattern of three bites often around the ankle or shin close to each other has garnered the macabre colloquialism "breakfast, lunch & dinner."

A technique for catching bedbugs in the act is to have a light source quickly accessible from your bed and to turn it on at about an hour before dawn, which is usually the time when bedbugs are most active. A flashlight/torch is recommended instead of room lights, as the act of getting out of bed will cause any bedbugs present to scatter before you can catch them. If you awaken during the night, leave your lights off but use your flashlight/torch to inspect your mattress. Bedbugs are fairly fast in their movements, about equal to the speed of ants. They may be slowed down if engorged. When the bedroom light is switched on, it may temporarily startle them allowing time for you to get a dust pan and brush kept next to the bed and sweep the bugs into the pan then immediately sweep them into a cup or mug full of water where the bugs drown quickly. Dispose of the water down the sink or toilet. Disinfect the mattress, skirting boards and so on regularly. Glue traps placed in strategic areas around the home, sometimes used in conjunction with heating pads or balloons filled with exhaled breath offering a carbon dioxide source, may be used to trap and thus detect bedbugs. This method has varied reports of success. There are also commercial traps like 'flea' traps whose effectiveness is questionable except perhaps as a means of detection. Perhaps the easiest trapping method is to place double-sided carpet tape in long strips near or around the bed and check the strips after a day or more.

It is found primarily on maple and ash trees. The adults are about 12½ mm (½ in) long with a dark brown or black coloration, relieved by red wing veins and markings on the abdomen. Nymphs and immature bugs are bright red. These insects feed on the softer plant tissues, including leaves, flowers, and new twigs. Unless the population is exceptionally large, the damage to plants is minimal. During years when their population soars, they can damage useful shade trees.

In autumn, they can become household pests. The adult insects seek wintering hibernation locations and find their way into buildings through crevices. They remain inactive inside the walls (and behind siding) while the weather is cool. When the heating systems revive them, they begin to enter inhabited parts of the buildings. In the spring, the bugs leave their winter hibernation locations to lay eggs on maple or ash trees. In late spring, groups of 50-200+ bugs may gather on house siding or brick, usually in a sunny spot. A month or two later you may find pairs of them mating, connected end to end, also in groups of 3 and 4.

The brown recluse spider is uncommon in Ohio. Nonetheless, OSU Extension receives numerous spider specimens that homeowners mistakenly suspect to be the brown recluse. Media attention and public fear contribute to these misdiagnoses. The brown recluse belongs to a group of spiders that is officially known as the "recluse spiders" in the genusLoxosceles (pronounced lox-sos-a-leez). These spiders are also commonly referred to as "fiddleback" spiders or "violin" spiders because of the violin-shaped marking on the top surface of the cephalothorax (fused head and thorax). However, this feature can be very faint depending on the species of recluse spider, particularly those in the southwestern U.S., or how recently the spider has molted. The common name, brown recluse spider, pertains to only one species, Loxosceles reclusa. The name refers to its color and habits. It is a reclusive creature that seeks and prefers seclusion.

Distribution
The brown recluse spider and ten additional species of Loxosceles are native to the United States. In addition, a few non-native species have become established in limited areas of the country. The brown recluse spider is found mainly in the central Midwestern states southward to the Gulf of Mexico (see map). Isolated cases in Ohio are likely attributable to this spider occasionally being transported in materials from other states. Although uncommon, there are more confirmed reports of Loxosceles rufescens (Mediterranean recluse) than the brown recluse in Ohio. It, too, is a human-associated species with similar habits and probably similar venom risks (unverified).

Identification
Recluse spiders have six eyes that are arranged in pairs. In the mature brown recluse spider as well as some other species of recluse spiders, the dark violin marking is well defined, with the neck of the violin pointing toward the bulbous abdomen. The abdomen is uniformly colored, although the coloration can range from light tan to dark brown, and is covered with numerous fine hairs that provide a velvety appearance. The long, thin, brown legs also are covered with fine hairs, but not spines. Adult brown recluse spiders have a leg span about the size of a quarter. Their body is about 3/8 inches long and about 3/16 inches wide. Males are slightly smaller in body length than females, but males have proportionally longer legs. Both sexes are venomous. The immature stages closely resemble the adults except for size and a slightly lighter color. Whereas most spiders have eight eyes, recluse spiders have six eyes that are arranged in pairs in a semicircle on the forepart of the cephalothorax (see close-up view). A 10X hand lens or microscope is needed to see this diagnostic feature. In order to determine the exact species of Loxosceles, the spider's genitalia need to be examined under a high-power microscope. This requires the skills of a spider expert.

Life Cycle and Habits
Egg laying primarily occurs from May through July. The female lays about 50 eggs that are encased in an off-white silken sac that is about 2/3-inch diameter. Each female may produce several egg sacs over a period of several months. Spiderlings emerge from the egg sac in about a month or less. Their development is slow and is influenced by weather conditions and food availability. It takes an average of one year to reach the adult stage from time of egg deposit. Adult brown recluse spiders often live about one to two years. They can survive long periods of time (about 6 months) without food or water.

The brown recluse spider spins a loose, irregular web of very sticky, off-white to grayish threads. This web serves as the spider's daytime retreat, and it often is constructed in an undisturbed corner. This spider roams at night searching for insect prey. Recent research at the University of Kansas indicates that the brown recluse spider is largely a scavenger, preferring dead insects. Mature males also roam in search of females.

Brown recluse spiders generally occupy dark, undisturbed sites, and they can occur indoors or outdoors. In favorable habitats, their populations are usually dense. They thrive in human-altered environments. Indoors, they may be found in attics, basements, crawl spaces, cellars, closets, and ductwork or registers. They may seek shelter in storage boxes, shoes, clothing, folded linens, and behind furniture. They also may be found in outbuildings such as barns, storage sheds, and garages. Outdoors, brown recluse spiders may be found underneath logs, loose stones in rock piles, and stacks of lumber.

The brown recluse spider is not aggressive, and it normally bites only when crushed, handled or disturbed. Some people have been bitten in bed after inadvertently rolling over onto the spider. Others have been bitten after accidentally touching the spider when cleaning storage areas. Some bites occur when people put on seldom used clothing or shoes inhabited by a brown recluse.

They are often mistaken for a bumblebee species, as they can be similar in size and coloration, though most carpenter bees have a shiny abdomen, while in bumblebees the abdomen is completely clothed with dense hair. Males of some species have a white or yellow face, where the females do not males also often have much larger eyes than the females, which relates to their mating behavior. Male bees are often seen hovering near nests, and will approach nearby animals. However, males are harmless since they do not have a stinger. Female bees do have a stinger, but are not aggressive, and will not sting unless directly provoked.

Behavior
Carpenter bees are traditionally considered solitary bees, though some species have simple social nests in which mothers and daughters may cohabit. However, even solitary species tend to be gregarious, and often several will nest near each other. It has been occasionally reported that when females cohabit, there may be a division of labor between them, where one female may spend most of her time as a guard within the nest, motionless and near the entrance, while another female spends most of her time foraging for provisions.

Carpenter bees make nests by tunneling into wood, vibrating their bodies as they rasp their mandibles against the wood, each nest having a single entrance which may have many adjacent tunnels. Carpenter bees do not eat wood. They discard the bits of wood, or re-use particles to build partitions between cells. The tunnel functions as a nursery for brood and the pollen/nectar upon which the brood subsists. The provision masses of some species are among the most complex in shape of any group of bees whereas most bees fill their brood cells with a soupy mass, and others form simple spheroidal pollen masses, Xylocopaform elongate and carefully sculpted masses that have several projections which keep the bulk of the mass from coming into contact with the cell walls, sometimes resembling an irregular caltrop. The eggs are very large relative to the size of the female, and are some of the largest eggs among all insects.

There are two very different mating systems that appear to be common in carpenter bees, and often this can be determined simply by examining specimens of the males of any given species. Species in which the males have large eyes are characterized by a mating system where the males either search for females by patrolling, or by hovering and waiting for passing females, whom they then pursue. In the other type of mating system, the males often have very small heads, but there is a large, hypertrophied glandular reservoir in the mesosoma, which releases pheromones into the airstream behind the male while it flies or hovers. The pheromone advertises the presence of the male to females.

There are about 4,000 species of cockroach, of which 30 species are associated with human habitations and about four species are well known as pests.

Among the best-known pest species are the American cockroach, Periplaneta americana, which is about 30 millimetres (1.2 in) long, the German cockroach, Blattella germanica, about 15 millimetres (½ in) long, the Asian cockroach, Blattella asahinai, also about 15 millimetres (½ in) in length, and the Oriental cockroach, Blatta orientalis, about 25 millimetres (1 in).

Size
Cockroaches are rather large insects. Most species are about the size of a thumbnail, but several species are bigger. The world's largest cockroach is the Australian giant burrowing cockroach, which can reach 9 cm in length and weigh more than 30 grams. Comparable in size is the Central American giant cockroach Blaberus giganteus, which grows to a similar length but is not as heavy.

Eggs and egg capsules
Female cockroaches are sometimes seen carrying egg cases on the end of their abdomen the egg case of the German cockroach holds about 30–40 long, thin eggs, packed like frankfurters in the case called an ootheca. The eggs hatch from the combined pressure of the hatchlings gulping air and are initially bright white nymphs that continue inflating themselves with air and harden and darken within about four hours. Their transient white stage while hatching and later while molting has led to many claims of glimpses of an albino cockroach.

A female German cockroach carries an egg capsule containing around 40 eggs. She drops the capsule prior to hatching, though live births do rarely occur. Development from eggs to adults takes 3-4 months. Cockroaches live up to a year. The female may produce up to eight egg cases in a lifetime in favorable conditions, it can produce 300-400 offspring. Other species of cockroach, however, can produce an extremely high number of eggs in a lifetime, but in some cases a female only needs to be impregnated once to be able to lay eggs for the rest of her life.

Crab spiders make up the Thomisidae family of the Araneae order. They are called crab spiders because they resemble crabs, with two front pairs of legs angled outward and bodies that are flattened and often angular. Also, like crabs, Thomisidae can move sideways or backward.

Crab spiders do not build webs to trap prey, but are hunters and ambushers. Some species sit on or among flowers, bark, fruit or leaves where they grab visiting insects. Individuals of some species, such as Misumena vatia, are able to change color between white and yellow to match the flower on which they're sitting. Other species, with their flattened bodies, hunt in the crevices of tree trunks or under loose bark. Members of the genus Xysticus hunt in the leaf litter on the ground. In each case, crab spiders use their powerful front legs to grab and hold onto prey while paralyzing it with a venomous bite.

The spider family Aphantochilidae was incorporated into the Thomisidae in the late 1980s. Aphantochilus species mimic Cephalotes ants, on which they prey. The spiders of Thomisidae are not known to be harmful to humans. However, spiders of an unrelated genus, Sicarius, which are sometimes referred to as "crab spiders", are close cousins to the recluse spiders, and are highly venomous.

Earwigs is the common name given to the insect order Dermaptera characterized by membranous wings folded underneath short leathery forewings (hence the literal name of the order—"skin wings"). The abdomen extends well beyond the wings, and frequently, though not always, ends in a pair of forceps-like structures termed cerci. The order is relatively small among Insecta, with about 1,800 recorded species in 10 families. Earwigs are, however, quite common globally. There is no evidence that they transmit disease or otherwise harm humans or other animals, despite their nickname pincher bug or "pinch ass".

Appearance and behaviour
Most earwigs are elongated, flattened, and are dark brown. Lengths are mostly in the quarter- to half-inch range (10–14 mm), with the Saint Helena Giant Earwig reaching three inches (80 mm). Cerci range from nonexistent to long arcs up to one-third as long as the rest of the body. Mouthparts are designed for chewing, as in other orthopteroid insects. Flight capability in Dermaptera is varied, as there are species with and without wings. In those earwigs that have wings (are not apterous), the hindwings are folded in a complex fashion, so that they fit under the forewings. Most species of winged earwigs are capable of flight, yet earwigs rarely fly around.

The abdomen of the earwig is flexible and muscular. It is capable of maneuvering as well as opening and closing the forceps. The forceps are used for a variety of purposes. In some species, the forceps have also been observed in use for holding prey, and incopulation. The forceps tend to be more curved in males than in females.

Most earwigs found in Europe and North America are of the species Forficula auricularia, the European or common earwig, which is distributed throughout the cooler parts of the northern hemisphere. This species feeds on other insects, plants, ripe fruit, and garbage. Plants that they feed on typically include clover, dahlias, zinnias, butterfly bush, hollyhock, lettuce, cauliflower,strawberry, sunflowers, celery, peaches, plums, grapes, potatoes, roses, seedling beans andbeets, and tender grass shoots and roots they have also been known to eat corn silk, damaging the corn. Typically they are a nuisance because of their diet, but normally do not present serious hazards to crops. Some tropical species are brightly colored. Occasionally earwigs are confused with cockroaches because of their cerci and their long antennae.

Earwigs are generally nocturnal and can be seen patrolling household walls and ceilings. Interaction with earwigs at this time results in a defensive free fall to the ground below, and the subsequent scramble to a nearby cleft or crevice. Earwigs are also drawn to damp conditions. During the summer, they can be found around sinks and in bathrooms. Earwigs tend to gather in shady cracks or openings or anywhere that they can remain concealed in daylight hours. Picnic tables, compost and waste bins, patios, lawn furniture, window frames, or anything with minute spaces (even artichoke blossoms) can potentially harbor these unwanted residents. Upon gaining entry to the basement and living areas of the home, earwigs can easily find cover in undisturbed magazine and newspaper piles, furniture/wickerwork, base boards, carpeted stairways, pet food dishes, and even inside DVD cases and keyboards. Earwigs are exploratory creatures and are often found trapped in poison baited cups or buckets of soapy water.

Adult Eastern cicada killer wasps are large, 1.5 to 5 cm (2/3 to 2 inches) long, robust wasps with hairy, reddish and black areas on the thorax (middle part), and are black to reddish brown marked with light yellow stripes on the abdominal (rear) segments. The wings are brownish. Coloration may superficially resemble that ofyellowjackets or hornets. The females are somewhat larger than the males, and both are among the largest wasps seen in the Eastern United States, their unusual size giving them a uniquely fearsome appearance. European hornets (Vespa crabro) are often mistaken for Eastern cicada killers. species occurs in the eastern and midwest U.S. and southwards into Mexico and Central America. They are so named because they hunt cicadas and provision their nests with them.

Life cycle and habits
Solitary wasps (such as the Eastern cicada killer) are very different in their behavior from the social wasps such ashornets, yellowjackets, and paper wasps. Cicada killer females use their sting to paralyze their prey (cicadas) rather than to defend their nests. Adults feed on flower nectar and other plantsap exudates. Adults emerge in summer, typically beginning around late June or early July and continuing throughout the summer months. They are present in a given area for 60 to 75 days, until mid-September. The large females are commonly seen in mid-to-late summer skimming around lawns seeking good sites to dig burrows and searching shrubs and trees for cicadas.

The males are more often seen in groups, vigorously challenging one another for position on the breeding aggregation from which they emerged, and generally pursuing anything that moves or flies within close proximity. It is not unusual to see two or three male wasps locked together in midair combat, the aggregate adopting an erratic and uncontrolled flight path until one of the wasps breaks away. The male wasp's aggressive behavior is extremely similar to that of another robust insect of the area, the male carpenter bee. In both cases, while the males' vigorous territorial defense can be extremely frightening and intimidating to human passersby, the males pose no danger whatsoever. They will only grapple with other insects, and cannot sting.While they may be frightfully large, female cicada killer wasps are not aggressive and rarely sting unless they are grasped roughly, stepped upon with bare feet, or caught in clothing, etc. One author who has been stung indicates that, for him, the stings are not much more than a "pinprick". Males aggressively defend their perching areas on nesting sites against rival males but they have no sting. Although they appear to attack anything which moves near their territories, male cicada killers are actually investigating anything which might be a female cicada killer ready to mate. Such close inspection appears to many people to be an attack, but male and female cicada killers don't land on people and attempt to sting. If handled roughly females will sting, and males will jab with a sharp spine on the tip of their abdomen. Both sexes are well equipped to bite, as they have large jaws however, they don't appear to grasp human skin and bite. They are non-aggressive towards humans and usually fly away when swatted at, instead of attacking. Cicada killers exert a natural control on cicada populations and thus may directly benefit the deciduous trees upon which their cicada prey feed.

This ground-burrowing wasp may be found in well-drained, sandy soils to loose clay in bare or grass-covered banks, berms and hills as well as next to raised sidewalks, driveways and patio slabs. Females may share a burrow, digging their own nest cells off the main tunnel. A burrow is 15 to 25 cm (6 - 10 in.) deep and about 3 cm (1.5 in.) wide. The female dislodges the soil with her jaws and pushes loose soil behind her as she backs out of the burrow using her hind legs, which are equipped with special spines that help her push the dirt behind her. The excess soil pushed out of the burrow forms a mound with a trench through it at the burrow entrance. Cicada killers may nest in planters, window boxes, flower beds or under shrubs, ground cover, etc. Nests often are made in the full sun where vegetation is sparse.

After digging a nest chamber in the burrow, female cicada killers capture cicadas, paralyzing them with a sting the cicadas then serve as food to rear their young. After paralyzing a cicada, the female wasp straddles it and takes off toward her burrow this return flight to the burrow is difficult for the wasp because the cicada is often more than twice her weight. After putting the cicada in the nest cell, the female deposits an egg on the cicada and closes the cell with dirt. Male eggs are laid on a single cicada but female eggs are given two or sometimes three cicadas this is because the female wasp is twice as large as the male and must have more food. New nest cells are dug as necessary off the main burrow tunnel and a single burrow may eventually have 10 to 20 cells. The egg hatches in one or two days, and the cicadas serve as food for the grub. The larvae complete their development in about 2 weeks. Overwintering occurs as a mature larva within an earth-coated cocoon. Pupation occurs in the nest cell in the spring and lasts 25 to 30 days. There is only one generation per year and no adults overwinter.

This wasp is frequently attacked by the parasitic "velvet ant" wasp, Dasymutilla occidentalis, also known as the "cow-killer" wasp. It lays an egg in the nest cell of the cicada killer, and when the cicada killer larva pupates, the parasitoid larva consumes the pupa.

Fleas are small (1/16 to 1/8-inch (1.5 to 3.3 mm) long), agile, usually dark colored (for example, the reddish-brown of the cat flea), wingless insects with tube-like mouth-parts adapted to feeding on the blood of their hosts. Their bodies are laterally compressed (human anatomical terms), permitting easy movement through the hairs or feathers on the host's body (or in the case of humans, under clothes). Their legs are long, the hind pair well adapted for jumping (vertically up to seven inches (18 cm) horizontally thirteen inches (33 cm) - around 200 times their own body length, making the flea one of the best jumpers of all known animals (in comparison to body size), second only to thefroghopper. The flea body is hard, polished, and covered with many hairs and short spines directed backward, which also assists its movements on the host. Its tough body is able to withstand great pressure, likely an adaptation to survive attempts to eliminate them such as scratching. Even hard squeezing between the fingers is normally insufficient to kill the flea it may be necessary to capture them with adhesive tape, crush them between the fingernails, roll them between the fingers, or put them in a fire-safe area and burn them with match or lighter. They can also be drowned. Flea larvae emerge from the eggs to feed on any available organic material such as dead insects, feces, and vegetable matter. They are blind and avoid sunlight, keeping to dark places like sand, cracks and crevices, and bedding. Given an adequate supply of food, larvae should pupate and weave a silken cocoon within 1-2 weeks after 3 larval stages. After another week or two, the adult flea is fully developed and ready to emerge from the cocoon. They may however remain resting during this period until they receive a signal that a host is near - vibrations (including sound), heat, and carbon dioxideare all stimuli indicating the probable presence of a host. Fleas are known to overwinter in the larval or pupal stages.

Fleas lay tiny white oval shaped eggs. Their larvae are small and pale with bristles covering their worm-like body. They lack eyes, and have mouthparts adapted to chewing. While the adult flea's diet consists solely of blood, the larvae feed on various organic matter, including the feces of mature fleas. In the pupal phase the larvae are enclosed in a silken, debris-covered cocoon.

Life cycle and habitat
Fleas are holometabolous insects, going through the three life cycle stages of larva, pupa, and imago(adult). The flea life cycle begins when the female lays after feeding. Adult fleas must feed on blood before they can become capable of reproduction. Eggs are laid in batches of up to 20 or so, usually on the host itself, which easily roll onto the ground. As such, areas where the host rests and sleeps become one of the primary habitats of eggs and developing fleas. The eggs take around two days to two weeks to hatch.Flea larvae emerge from the eggs to feed on any available organic material such as dead insects, feces, and vegetable matter. They are blind and avoid sunlight, keeping to dark places like sand, cracks and crevices, and bedding. Given an adequate supply of food, larvae should pupate and weave a silken cocoon within 1-2 weeks after 3 larval stages. After another week or two, the adult flea is fully developed and ready to emerge from the cocoon. They may however remain resting during this period until they receive a signal that a host is near - vibrations (including sound), heat, and carbon dioxideare all stimuli indicating the probable presence of a host. Fleas are known to overwinter in the larval or pupal stages.

Once the flea reaches adulthood its primary goal is to find blood - adult fleas must feed on blood in order to reproduce. Adult fleas only have around a week to find food once they emerge, though they can survive two months to a year between meals. A flea population is unevenly distributed, with 50 percent eggs, 35 percent larvae, 10 percent pupae, and 5 percent adults. Their total life cycle can take as little as two weeks, but may be lengthened to many months if conditions are favorable. Female fleas can lay 500 or more eggs over their life, allowing for phenomenal growth rates.

The best-known bee species is the European honey bee, which, as its name suggests, produceshoney, as do a few other types of bee. Human management of this species is known as beekeeping or apiculture.

The true honey bees (genus Apis) have arguably the most complex social behavior among the bees. TheEuropean (or Western) honey bee, Apis mellifera, is the best known bee species and one of the best known of all insects.

Dolichovespula maculata is a North American insect which, despite commonly being called thebald-faced hornet (or white-faced hornet), is not a true hornet at all. It belongs to a genus ofwasps called yellowjackets in North America, and is more distantly related to true hornets like the Asian giant hornet or European hornet, but the term "hornet" is often used colloquially to refer to any vespine with an exposed aerial nest.

The bald-faced hornet lives throughout North America, including southern Canada, the Rocky Mountains, the western coast of the United States, and most of the eastern US. They are most common in the southeastern United States. They are best known for their large football-shapedpaper nest, which they build in the spring for raising their young. These nests can sometimes reach 3 feet tall. Like the median wasp Dolichovespula media in Europe, bald-faced hornets are extremely protective of their nests and will sting repeatedly if disturbed.

Every year young queens that were born and fertilized the previous year start a new colony and raise their young. The workers expand the nest by chewing up wood that mixes with a starch in their saliva, which they spread with their mandibles and legs to dry into paper. The workers also guard the nest and collect nectar and arthropods to feed the larvae. This continues through summer and into fall. As winter approaches, the wasps die, except for young fertilized queens which hibernate underground or in hollow trees. The nest is generally abandoned by winter, and will most likely not be reused. When spring arrives the young queens emerge, and the cycle begins again.
Bald-faced hornets visit flowers, especially in late summer, and can be minor pollinators. Like other social wasps, bald-faced hornets have a caste system made up of the following:
1. Queens — fertile females which begin the colonies and lay eggs.
2. Workers — infertile females which do the manual labor.
3. Drones — males, which have no stingers, and are born from unfertilized eggs.

Jumping spiders are generally diurnal, active hunters. Their well developed internal hydraulicsystem extends their limbs by altering the pressure of body fluid (blood) within them. This enables the spiders to jump without having large muscular legs like a grasshopper. The jumping spider can therefore jump 20 to 60 or even 75-80 times the length of their body. When a jumping spider is moving from place to place, and especially just before it jumps, it tethers a filament of silk to whatever it is standing on. Should it fall for one reason or another, it climbs back up the silk tether.

Jumping spiders are Scopula bearing spiders, which means that they have a very interesting Tarsal section. And the end of each leg they have hundreds of tiny hairs, which each then split into hundreds more tiny hairs, each tipped with an "end foot". These thousands of tiny feet allow them to climb up and across virtually any terrain. They can even climb up glass by gripping onto the tiny imperfections, usually an impossible task for any spider.

Jumping spiders also use their silk to weave small tent-like dwellings where females can protect their eggs, and which also serve as a shelter while moulting. Jumping spiders are known for their curiosity. If approached by a human hand, instead of scuttling away to safety as most spiders do, the jumping spider will usually leap and turn to face the hand. Further approach may result in the spider jumping backwards while still eyeing the hand. The tiny creature will even raise its forelimbs and "hold its ground". Because of this contrast to other arachnids, the jumping spider is regarded as inquisitive as it is seemingly interested in whatever approaches it.

Mice range in size from 12 to 21 cm (4 to 8 inches) long (including a long tail). They weigh from .25 to 2 oz (7.1 to 57 g). The coat color ranges from white to brown to gray. Most mice have a pointed snout with long whiskers, round ears, and thin tails. Many mice scurry along the ground, but some can hop or jump.

Although mice may live up to two and a half years in captivity, the average mouse in the wild lives only about four months, primarily owing to heavy predation. Cats, wild dogs, foxes, birds of prey, snakes and even certain kinds of insects have been known to prey heavily upon mice. Nevertheless, because of its remarkable adaptability to almost any environment, and its ability to live commensally with humans, the mouse is regarded to be the second most successful mammalian genus living on Earth today, after humans.

Mice can at times be harmful pests, damaging and eating crops and spreading diseasesthrough their parasites and feces. In western North America, breathing dust that has come in contact with mouse feces has been linked to the deadly hantavirus.

Mud dauber (sometimes "dirt dauber," "dirt dobber," or "dirt diver" in the southern U.S.) is a name commonly applied to a number of wasps from either the family Sphecidae orCrabronidae that build their nests from mud. Mud dauber may refer to any of the following common species:

- The solid black organ pipe mud dauber, Trypoxylon politum (family Crabronidae)
- The black and yellow mud dauber, Sceliphron caementarium (family Sphecidae)
- The irridescent blue mud dauber, Chalybion californicum (family Sphecidae)
Mud daubers are long, slender wasps, the latter two species above with thread-like waists. The name of this wasp group comes from the nests that are made by the females, which consist of mud molded into place by the wasp's mandibles.

The organ-pipe mud dauber, as the name implies, builds nests in the shape of a cylindrical tube resembling an organ pipe or pan flute. The black and yellow mud dauber's nest is composed of a series of cylindrical cells that are plastered over to form a smooth nest about the size of a lemon. The metallic-blue mud dauber foregoes building a nest altogether and simply uses the abandoned nests of the other two species and preys primarily on black widow spiders.

Mud daubers are rarely aggressive. They do however pose a special risk to aircraft operation, as they are prone to nest in the small openings and tubes that compose aircraftpitot-static systems. Their presence in these systems can disable or impair the function of the airspeed indicator, the altimeter, and/or the vertical speed indicator. It is thought that mud dauber wasps were ultimately responsible for the crash of Birgenair Flight 301, which killed 189 passengers and crew.

Pillbugs (woodlice of the family Armadillidiidae) can be confused with pill millipedes although they are only very distantly related to one another.

Both of these groups of terrestrial segmented arthropods are about the same size. They live in very similar habitats, and they can both roll up into a ball. Both pill millipedes and pillbugs appear superficially similar to the naked eye. This is an example of convergent evolution.

The larvae of these beetles reduce timbers to a mass of very fine, powder-like substance.

Adults do little damage, it is the larvae that does the major part of the damage.They go through a complete metamorphosis: adults, eggs, larvae and pupae.

You can easily recognize the work of powder post beetles. When the adults emerge, usually in June, some species leave small holes about the size of a pin in the surface of the wood others make holes the size of pencil lead. From these holes, a fine, powder like brood of larvae carry on their destructive feeding. Normally, these insects have a 1-year life cycle this means that the adults will appear only once each year. And because of this habit the larvae have a feeding period of many months.

True Powderpost Beetles(Lyctidae):
The adults are very small, less than 1/4" in size. They are flattened and reddish-brown to black in color. Larvae are white, cream colored, shaped with dark brown heads. Larvae create tunnels in the wood and become pupae. As adults they bore out through the wood, pushing a fine powdery dust out.The shape of their holes are round ,about 1/32-1/16 pinholes.

They attack hardwoods depositing their eggs. True Powder post beetles breed in dead and dried hardwoods such as the dead branches and limbs of trees. Their presence is overlooked until they are discovered in stored lumber, rafters, joists, finished wood, and furniture products. As a rule, they enter lumber while it is being stored and cured, then later, emerge from the finished product. Old items of furniture and wood antiques are especially vulnerable to attack by the beetles.

Damage is usually to the starch-rich sapwood of large-pored hardwoods such as ash, hickory, oak, walnut and cherry. The hardwood floors of new homes are commonly attacked.

Their diet is starch, sugar and protein in the sapwood of hardwoods Wood that is less than 6% moisture content is seldom attacked. The life cycle averages one year to complete. This wood-boring beetle is the most widespread in the United States. Many times infestations are built into structures from infested lumber. They can re infest.
Lycid damage is characterized by:
- Presence of extremely fine, flour like powder falling from the surface holes.
- The frass left by other wood borers usually contains pellets, has a course texture and a tendency to stick together.
- When inspecting damage, be sure to distinguish old damage from active beetle infestations.
- Recently formed holes and frass(sawdust like) are light in color and clear in appearance - old holes and frass are dark in color.

Termites remain hidden within wood and are often difficult to detect. However, subterranean termites may be detected by the presence of winged reproductives, mud tubes, and wood damage.

Biology
Subterranean termites are social insects that live in colonies consisting of many individuals. The colonies are composed of workers, soldiers and reproductives. The workers, which are about 1/8 inch long, have no wings, are white to cream colored and very numerous. Soldiers defend the colony against insects, like ants, that can attack the colony. Soldiers are wingless and white in color with large brown heads and mandibles (jaws). King and queen termites perform the reproductive functions of the colony. They are dark brown to black in color and have two pairs of wings about twice the length of their body.

Subterranean termites feed on wood or other items that contain cellulose, such as paper, fiberboard, and some fabrics derived from cotton or plant fibers. Termites have protozoa in their digestive tracts that can convert cellulose into usable food.

Subterranean termites nest in the soil to obtain moisture, but they also nest in wood that is often wet. They easily attack any wood in contact with the ground. If the wood does not contact the soil, they can build mud tunnels or tubes to reach wood several feet above the ground. These tunnels can extend for 50-60 feet to reach wood and often enter a structure through expansion joints in concrete slabs or where utilities enter the house.

The Formosan subterranean termite is often nicknamed the super-termite because of its destructive habits. This is because of the large size of its colonies, and the termites' ability to consume wood at a rapid rate. A single colony may contain several million (compared with several hundred thousand termites for other subterranean termite species) that forage up to 300 feet (100 m) in soil. A mature Formosan colony can consume as much as 13 ounces of wood a day and severely damage a structure in as little as three months. Because of its population size and foraging range, the presence of colonies poses serious threats to nearby structures. Once established, Formosan subterranean termite has never been eradicated from an area.

Formosan subterranean termites infest a wide variety of structures (including boats and high-risecondominiums) and can damage trees. In the United States it is responsible for tremendous property damage resulting in large treatment and repair costs.

Winged Reproductives
Winged reproductives emerge from colonies in great numbers usually in the spring and during the daylight hours. Usually termites are first noticed by the presence of winged reproductives. Mating occurs during these flights, and males and females form new colonies. Winged termites can be distinguished from flying ants by their thick-waist, straight antennae and wings of equal size.

A wasp is a predatory flying stinging insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. A narrower and simpler but popular definition of the term wasp is any member of the aculeate family Vespidae. Wasps are critically important in natural biocontrol as almost every pest insect species has at least one wasp species that is a predator upon it.

Categorization
The various species of wasp fall into one of two main categories: solitary wasps and social wasps. Adult solitary wasps generally live and operate alone, and most do not construct nests (below) all adult solitary wasps are fertile. By contrast, social wasps exist in colonies numbering up to several thousand strong and build nests—but in some cases not all of the colony can reproduce. In the more advanced species, just the wasp queen and male wasps can mate, whilst the majority of the colony is made up of sterile female workers.

Wolf spiders are members of the family Lycosidae, from the Greek word "λύκος" meaning "wolf". They are robust and agile hunters, and have good eyesight. They live mostly solitary lives and hunt alone. Some are opportunistic wanderer hunters, pouncing upon prey as they find it or chasing it over short distances. Others lie in wait for passing prey, often from or near the mouth of a burrow.

Like all species, wolf spiders have a primitive body structure, with a head used mainly for eating and breathing, and an abdomen, which carries all the spider's organs, including the spinnerets. Many of the sub- species, which include the large and common grey wolf spider, are a mixture of greys and light browns, hence the name.

Yellowjacket or yellow-jacket is the common name in North America for predatory wasps of the genera Vespula and Dolichovespula. Members of these genera are known simply as "wasps" in other English-speaking countries. Most of these are black-and-yellow some are black-and-white (such as the bald-faced hornet, Dolichovespula maculata), while others may have the abdomen background color red instead of black. They can be identified by their distinctive markings, small size (similar to or slightly smaller or larger than a honey bee), their occurrence only in colonies, and a characteristic, rapid, side to side flight pattern prior to landing. They are often mistakenly called "bees". All females are capable ofstinging. Yellowjackets are important predators of pest insects.

Identification
A typical yellowjacket worker is about 12 mm (0.5 inches) long, with alternating bands on the abdomen while the queen is larger, about 19 mm (0.75 inches) long (the different patterns on the abdomen help separate various species). Workers are sometimes confused with honey bees, especially when flying in and out of their nests. Yellowjackets, in contrast to honey bees, are not covered with tan-brown dense hair on their bodies and lack the flattened hairy hind legs used to carry pollen. Yellowjackets have a lance-like stinger with small barbs and typically sting repeatedly, though occasionally the sting becomes lodged and pulls free of the wasp's body the venom, like most bee/wasp venoms, is primarily only dangerous to those who are allergic, unless a victim receives a large number of stings (main article: Bee sting). All species have yellow or white on the face. Mouthparts are well-developed for capturing and chewing insects, with a proboscis for sucking nectar, fruit and other juices. Nests are built in trees, shrubs or in protected places such as inside human-made structures (attics, hollow walls or flooring, in sheds, under porches and eaves of houses), or in soil cavities, mouse burrows, etc. Nests are made from wood fiber chewed into a paper-like pulp. Yellowjackets have two antennae and two wings. These two wings are distinctive because they fold in half length-wise.

Due to their aggressive behavior, including stinging, many other insects exhibit mimicry of yellowjackets in addition to numerous bees andwasps (Müllerian mimicry), the list includes some flies, moths, and beetles (Batesian mimicry). Yellowjackets' closest relatives, the hornets, closely resemble them but have a much bigger head, seen especially in the large distance from the eyes to the back of the head.


Coals to Newcastle

It seems like Gaia really went on a bender in the late Carboniferous, getting drunk on oxygen. By some estimates, the atmosphere was over 30% oxygen back then, compared to 21% today. Living things took advantage of the opportunity. Insects apparently face an upper limit in size because they rely on diffusion through tracheas instead of forced respiration through lungs to get oxygen into their bodies. With more oxygen in the air, this limit was raised. The Carboniferous saw dragonflies with a wingspan up to 70 centimeters, and body lengths up to 30 centimeters, comparable to a seagull.

This happened because plants were turning carbon dioxide into organic matter and free oxygen, and the organic matter was accumulating. With carbon dioxide being removed from the atmosphere, the late Carboniferous and subsequent early Permian saw a reduced greenhouse effect, and global cooling. This was another Ice Age, with ice caps around the southern pole.

A lot of organic carbon ended up being buried. Much of the world’s coal, especially high quality anthracite, has its origin in Carboniferous tropical forests. Western Europe and eastern North America lay in the tropics at the time, and got a particularly generous allotment of coal. Three hundred million years later this bounty would fuel the early Industrial Revolution. (Thanks partly to some of my Welsh ancestors, who helped dig it up back in the day.)


12864_2011_3549_MOESM1_ESM.DOC

Additional file 1: PCR strategy. The PCR amplification strategy and collection records for the lice sequenced in this study. (DOC 56 KB)

Full phylogenetic tree of lice and relatives

Additional file 2: . Full tree generated after [31] which was pruned to produce figure 2. (EPS 869 KB)

Ne-mt genes found in

Additional file 3: Pediculus. Complete list of gene homologues and blast statistics for the annotation of ne-mt genes found in Pediculus. (XLS 111 KB)

12864_2011_3549_MOESM4_ESM.DOC

Additional file 4: Novel mt gene boundaries in lice. Complete list of novel mt gene boundaries found in lice. (DOC 62 KB)

12864_2011_3549_MOESM5_ESM.DOC

Additional file 5: mtSSB alignment. Alignment of the amino acid sequences of mtSSB genes annotated from insect nuclear genomes. (DOC 70 KB)



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