Evolution of Eusociality

The haplodiploid genetic system found in all Hymenopterans creates an asymmetry in genetic relatedness so that full-sistersare more closely related to each other than a mother is to her daughters. Thus Hymenopteran workers who rear siblings can. obtain higher inclusive fitness compared to individuals who rear offspring. However, polyandry and polygyny reduce relatedness between workers and their .;jsters and'thus tend to break down the genetic asymmetry created by haplodiploidy. Since the advent of electrophoretic analysis of variability at enzyme loci, several estimates of intra-colony genetic relatedness in the Hymenoptera have been published. To test the role 'of the genetic asymmetry created by haplodiploidy in the evolution of eusociality, I assume that workers are capable of investing in their brothers and sisters in their ratio of relatedness to them. I then compute a haplodiploidy threshold, which is the threshold relatedness to sisters required for workers to obtain a weighted mean relatedness of 0,5 to siblings and thus break even with solitary foundresses. When workers rear mixtures of sisters and brothers in an outbred population, the value of this threshold is 0,604. An examination of the distribution of 185 estimates of mean genetic relatedness between sisters in Hymenopteran colonies shows that the values are wellbelow the expected 0,75 for full sisters, both in higly eusocial as well as in primitively eusocial species although relatedness values in the latter are higher than in the former. Of the 177 estimates with standard error, 49 are significantly lower than the haplodiploidy threshold and 22 are significantly higher. Of the 35 species studied only 6 have one or more estimates that are significantly higher than the haplodiploidy threshold. For more than half the estimates, the probability of the relatedness value being above the haplodiploidy threshold is less than 0,5. Reanalysis of these data using 0,5 as the threshold does not drastically alter these conclusions. I conclude that the genetic asymmetry created by haplodiploidy is, in most cases, insufficientby itself either to promote the origin of eusocialityor to maintain the highly eusocialstate.

Eusocial insects are those that show overlap of generations, cooperative brood care and reproductive caste differentiation. Of these, primitively eusocial insects show no morphological differences between reproductive and worker castes and exhibit considerable flexibility in the social roles that adult females may adopt. This makes them attractive model systems for investigations concerning the origin of eusociality. The rapidly accumulating information on primitively eusocial wasps suggests that haplodiploidy is unlikely to have an important role in the origin of eusociality. General kin selection (without help from haplodiploidy) could however have been an important factor due to the many advantages of group living. Pre-imaginal caste bias leading to variations in fertility is also likely to have some role. Because workers often have some chance of becoming reproductives in future, mutualism and other individual selection models suggest themselves as important factors. A hypothesis for the route to eusociality which focuses on the factors selecting for group living at different stages in social evolution is presented. It is argued that group living originates owing to the benefit of mutualism (the 'Gambling Stage') but parental manipulation and subfertility soon become important (the 'Manipulation Stage') and finally the highly eusocial state is maintained because genetic asymmetries created by haplodiploidy are exploited by kin recognition (the 'Recognition Stage').

This has long been a species with close proximity of mankind (the species of Apis are distributed all over the Old World), with economic interest because of honey production (pollination became a hotly debated topic more recently, e.g., Oldroyd, 2007; Cameron et al., 2010; Potts et al.,

SignificanceIn eusocial insect societies, such as ants and some bees and wasps, phenotypes are highly plastic, generating alternative phenotypes (queens and workers) from the same genome. The greatest plasticity is found in simple insect societies, in which individuals can switch between phenotypes as adults. The genomic, transcriptional, and epigenetic underpinnings of such plasticity are largely unknown. In contrast to the complex societies of the honeybee, we find that simple insect societies lack distinct transcriptional differentiation between phenotypes and coherently patterned DNA methylomes. Instead, alternative phenotypes are largely defined by subtle transcriptional network organization. These traits may facilitate genomic plasticity. These insights and resources will stimulate new approaches and hypotheses that will help to unravel the genomic processes that create phenotypic plasticity.

Phylogenetic relationships of corbiculate bees have been a well-known focus of controversies over the past 30 years. The majority of the morphological datasets support the monophyly of Apina + Meliponina, whereas molecular datasets recover Meliponina as sister to Bombina. This issue is especially critical to the proper understanding of the evolution of clusters of traits that define the corbiculate eusocial behaviour. This work provides a description and discussion on characters of the head capsule, mandibles and sitophore of bees. Thirty-three characters are proposed and optimized within concurrent phylogenetic hypotheses for corbiculate bees, which results in seven derived character-states supporting the monophyly of the Apina + Meliponina + Bombina and nine supporting the Apina + Meliponina clades. Although some striking synapomorphies (e.g. tentorial bridge, pleurostomal condyle, hypopharyngeal lobe) support the former clade, most characters supporting the latter (i.e. Apina + Meliponina) were losses/reductions. Moreover, two previously undescribed character transformations on hypostoma and sitophore favour the Bombina + Meliponina clade. Internal head capsule characters are useful for phylogenetic analysis in comparative studies of bees, and corbiculate bees in particular, when efforts are made to solve the 'corbiculate controversy'. An attempt is made to standardize the current terminology used for bees and for other Hymenoptera.

SignificanceIn eusocial insect societies, such as ants and some bees and wasps, phenotypes are highly plastic, generating alternative phenotypes (queens and workers) from the same genome. The greatest plasticity is found in simple insect societies, in which individuals can switch between phenotypes as adults. The genomic, transcriptional, and epigenetic underpinnings of such plasticity are largely unknown. In contrast to the complex societies of the honeybee, we find that simple insect societies lack distinct transcriptional differentiation between phenotypes and coherently patterned DNA methylomes. Instead, alternative phenotypes are largely defined by subtle transcriptional network organization. These traits may facilitate genomic plasticity. These insights and resources will stimulate new approaches and hypotheses that will help to unravel the genomic processes that create phenotypic plasticity.

ABSTBACT The secretion of cuticular antimicrobial compounds is an important defensive mechanism for social insects and recent studies have demonstrated their role in the evolution of sociality. However, the factors that might affect their production and effectiveness are not well understood. For example, natural variation in colony size and genetic diversity are predicted to influence susceptibility to disease and, as a consequence, we hypothesized that colonies adjust by varying antimicrobial production. We examined the effects of colony size and genetic diversity on the antimicrobial properties of cuticular compounds in 10 colonies of the Australian paper wasp, Polistes humilis F. ( Vespidae: Polistinae). The effectiveness of antimicrobial compounds produced by a colony was found to increase with genetic diversity but decrease with colony size. Further, genetic diversity appears to be the stronger factor in determining effectiveness of antimicrobial compounds and may compensate fo...

Unlike the queens of other primitively eusocial species, Ropalidia marginata queens are strikingly docile and non-aggressive individuals, never at the top of the behavioural dominance hierarchy of their colonies. Nevertheless, these queens are completely successful at suppressing worker reproduction, suggesting that they do not use aggression but employ some other mechanism (e.g. pheromones) to do so. Upon removal of the queen from a colony, a single worker, the 'potential queen', immediately begins to display highly elevated levels of aggression towards her nest mates. This individual becomes the next docile queen if the original queen is not returned. We attempt to understand the function of the temporary and amplified dominance behaviour displayed by the potential queen. We find that the dominance behaviour shown by the potential queen is unrelated to the number of her nest mates, their dominance ranks or ovarian condition. This suggests that aggression may not be used to actively suppress other workers and counter threat. Instead we find evidence that dominance behaviour is required for the potential queen's rapid ovarian development, facilitating her speedy establishment as the sole reproductive individual in the colony.

Here, I explore the mechanisms and social phenomena of quorum sensing in both ant colonies and human groups. Quorum sensing refers to the process by which organisms coordinate behavior based on population density through chemical communication. Ants engage in quorum sensing through the use of pheromones to make collective decisions around tasks like foraging, nest selection, and defense. As more ants encounter resources and release pheromones, they achieve a "quorum" at which the entire colony responds in a coordinated manner without centralized control.

The paper examines examples of quorum sensing in human behaviors as well. During group decision-making, individuals exchange ideas through language and non-verbal cues, with consensus emerging once a threshold of agreement is reached. Other examples include the menstrual cycle, immune response coordination, and insulin regulation in the body. Concepts from the theory of eusociality are also discussed, highlighting characteristics like reproductive division of labor and cooperative care common to both ant colonies and human societies.

Analogies are drawn between ant and human behaviors mediated by quorum sensing, such as collective decision-making in boardrooms mirroring ant choices around nest relocation. The efficient and adaptable organization of ant colonies also informs our understanding of human social structures and dynamics. Understanding quorum sensing provides insights into controlling pest populations and improving coordination in challenging environments. Studying communication and cooperation in ant societies enhances comprehension of parallel processes in humans, with implications for addressing social and health issues.

http://www.eje.cz bee, Apis mellifera (Linnaeus, 1758) (Hymenoptera: Apidae), was fi rst chosen for the sequencing of the whole genome after the fruit fl y Drosophila melanogaster (Diptera: Drosophilidae) (Meigen, 1830), mosquito Anopheles gambiae (Diptera: Anophelinae) (Giles, 1902) and silkworm The draft genome sequence of the Japanese honey bee, Apis cerana japonica (Hymenoptera: Apidae

Apis cerana and Apis mellifera, are very important honey species for agriculture in Asian countries. In recent decades, A. cerana populations have sharply declined in all Asian countries as a result of Sacbrood Virus infection and have now recovered to their original size. It can change the genetic structure of local populations of A. cerana. We used the nuclear gene Vitellogenin VG to assess the genetic structure of local populations of A. cerana and the signature of adaptive selection. We performed a population genetic analysis of the honey bees A. cerana from South Korea in comparison with A. cerana samples from Russia, Japan, Nepal, and China. The sequences of the gene VG of a closely related honey bee species, A. mellifera, from India and Poland were used as outgroup samples. A comparative analysis of northern and southern A. cerana populations was performed. The signatures of positive adaptive selection were found in the local population of A. cerana. We performed the Tajima&#...

We argue that attempts to extrapolate moral motives for non-egoistic behavior in organizational behavior often interpret results empathically or deontically, while leaving other moral motivational frames, such as the utilitarian and virtue ethical, under-examined. We encourage the creation of experimental measures to distinguish various philosophical frames.

The Dufour gland in workers of vespine wasps appears as an unpaired tubiform gland that opens in close proximity to the sting base. The epithelial cells that line the central reservoir are characterized by apical microvillus-like projections and deep basal invaginations. Their cytoplasm contains a well-developed Golgi apparatus, numerous mitochondria, as well as strands of smooth endoplasmic reticulum. The Dufour gland duct occurs ventrally to the venom gland duct, and bends downward near the sting base to open in the dorsal vaginal wall. In this region, the duct is dorsoventrally flattened, and shows conspicuous bundles of parallel microtubules in the epithelial cells, that transmit the pulling forces of the myofilaments of the underlying muscular supply to the cuticle. This results in an active opening mechanism regulated by muscular contraction, while passive closure probably results from the return of the cuticular intima to a rest position.

Significance Most hypotheses explaining the evolution of sociality in insects assume that positive selection drives the evolution of worker traits. Yet we know little about the extent of natural selection acting on social insects. We produced a map of positive selection for the honey bee through analysis of 40 individual genomes. We found strong evidence of positive selection acting on genes and regulatory sequences, and we discovered that mutations in worker-biased proteins tend to have greater fitness effects than mutations in queen-biased proteins. We also found many instances of positive selection acting on genes that influence worker traits, suggesting that worker phenotypes represent a major vector for adaptation in social insects.

The fauna of termites (Isoptera) preserved in Early Eocene amber from the Cambay Basin (Gujarat, India) are described and figured. Three new genera and four new species are recognized, all of them Neoisoptera-Parastylotermes krishnai Engel & Grimaldi, sp. n. (Stylotermitidae); Prostylotermes kamboja Engel & Grimaldi, gen. et sp. n. (Stylotermitidae?); Zophotermes Engel, gen. n., with Zophotermes ashoki Engel & Singh, sp. n. (Rhinotermitidae: Prorhinotermitinae); and Nanotermes isaacae Engel & Grimaldi, gen. et sp. n. (Termitidae: Termitinae?). Together these species represent the earliest Tertiary records of the Neoisoptera and the oldest definitive record of Termitidae, a family that comprises >75% of the living species of Isoptera. Interestingly, the affinities of the Cambay amber termites are with largely Laurasian lineages, in this regard paralleling relationships seen between the fauna of bees and some flies. Diversity of Neoisoptera in Indian amber may reflect origin of the amber deposit in Dipterocarpaceae forests formed at or near the paleoequator.

Significance In eusocial insect societies, such as ants and some bees and wasps, phenotypes are highly plastic, generating alternative phenotypes (queens and workers) from the same genome. The greatest plasticity is found in simple insect societies, in which individuals can switch between phenotypes as adults. The genomic, transcriptional, and epigenetic underpinnings of such plasticity are largely unknown. In contrast to the complex societies of the honeybee, we find that simple insect societies lack distinct transcriptional differentiation between phenotypes and coherently patterned DNA methylomes. Instead, alternative phenotypes are largely defined by subtle transcriptional network organization. These traits may facilitate genomic plasticity. These insights and resources will stimulate new approaches and hypotheses that will help to unravel the genomic processes that create phenotypic plasticity.

Background: The first generation of genome sequence assemblies and annotations have had a significant impact upon our understanding of the biology of the sequenced species, the phylogenetic relationships among species, the study of populations within and across species, and have informed the biology of humans. As only a few Metazoan genomes are approaching finished quality (human, mouse, fly and worm), there is room for improvement of most genome assemblies. The honey bee (Apis mellifera) genome, published in 2006, was noted for its bimodal GC content distribution that affected the quality of the assembly in some regions and for fewer genes in the initial gene set (OGSv1.0) compared to what would be expected based on other sequenced insect genomes. Results: Here, we report an improved honey bee genome assembly (Amel_4.5) with a new gene annotation set (OGSv3.2), and show that the honey bee genome contains a number of genes similar to that of other insect genomes, contrary to what was suggested in OGSv1.0. The new genome assembly is more contiguous and complete and the new gene set includes~5000 more protein-coding genes, 50% more than previously reported. About 1/6 of the additional genes were due to improvements to the assembly, and the remaining were inferred based on new RNAseq and protein data. Conclusions: Lessons learned from this genome upgrade have important implications for future genome sequencing projects. Furthermore, the improvements significantly enhance genomic resources for the honey bee, a key model for social behavior and essential to global ecology through pollination.

Around 150 million years ago, eusocial termites evolved from within the cockroaches, 50 million years before eusocial Hymenoptera, such as bees and ants, appeared. Here, we report the first, 2GB genome of a cockroach, Blattella germanica, and the 1.3GB genome of the drywood termite, Cryptotermes secundus. We show evolutionary signatures of termite eusociality by comparing the genomes and transcriptomes of three termites and the cockroach against the background of 16 other eusocial and non-eusocial insects. Dramatic adaptive changes in genes underlying the production and perception of pheromones confirm the importance of chemical communication in the termites. These are accompanied by major changes in gene regulation and the molecular evolution of caste determination. Many of these results parallel molecular mechanisms of eusocial evolution in Hymenoptera. However, the specific solutions are remarkably different, thus revealing a striking case of convergence in one of the major evolu...

Around 150 million years ago, eusocial termites evolved from within the cockroaches, 50 million years before eusocial Hymenoptera, such as bees and ants, appeared. Here, we report the first, 2GB genome of a cockroach, Blattella germanica, and the 1.3GB genome of the drywood termite, Cryptotermes secundus. We show evolutionary signatures of termite eusociality by comparing the genomes and transcriptomes of three termites and the cockroach against the background of 16 other eusocial and non-eusocial insects. Dramatic adaptive changes in genes underlying the production and perception of pheromones confirm the importance of chemical communication in the termites. These are accompanied by major changes in gene regulation and the molecular evolution of caste determination. Many of these results parallel molecular mechanisms of eusocial evolution in Hymenoptera. However, the specific solutions are remarkably different, thus revealing a striking case of convergence in one of the major evolu...

Eusocial insects are those that show overlap of generations, cooperative brood care and reproductive caste differentiation. Of these, primitively eusocial insects show no morphological differences between reproductive and worker castes and exhibit considerable flexibility in the social roles that adult females may adopt. This makes them attractive model systems for investigations concerning the origin of eusociality. The rapidly accumulating information on primitively eusocial wasps suggests that haplodiploidy is unlikely to have an important role in the origin of eusociality. General kin selection (without help from haplodiploidy) could however have been an important factor due to the many advantages of group living. Pre-imaginal caste bias leading to variations in fertility is also likely to have some role. Because workers often have some chance of becoming reproductives in future, mutualism and other individual selection models suggest themselves as important factors. A hypothesis for the route to eusociality which focuses on the factors selecting for group living at different stages in social evolution is presented. It is argued that group living originates owing to the benefit of mutualism (the 'Gambling Stage') but parental manipulation and subfertility soon become important (the 'Manipulation Stage') and finally the highly eusocial state is maintained because genetic asymmetries created by haplodiploidy are exploited by kin recognition (the 'Recognition Stage').

Unlike the queens of other primitively eusocial species, Ropalidia marginata queens are strikingly docile and non-aggressive individuals, never at the top of the behavioural dominance hierarchy of their colonies. Nevertheless, these queens are completely successful at suppressing worker reproduction, suggesting that they do not use aggression but employ some other mechanism (e.g. pheromones) to do so. Upon removal of the queen from a colony, a single worker, the 'potential queen', immediately begins to display highly elevated levels of aggression towards her nest mates. This individual becomes the next docile queen if the original queen is not returned. We attempt to understand the function of the temporary and amplified dominance behaviour displayed by the potential queen. We find that the dominance behaviour shown by the potential queen is unrelated to the number of her nest mates, their dominance ranks or ovarian condition. This suggests that aggression may not be used to actively suppress other workers and counter threat. Instead we find evidence that dominance behaviour is required for the potential queen's rapid ovarian development, facilitating her speedy establishment as the sole reproductive individual in the colony.

Significance Most hypotheses explaining the evolution of sociality in insects assume that positive selection drives the evolution of worker traits. Yet we know little about the extent of natural selection acting on social insects. We produced a map of positive selection for the honey bee through analysis of 40 individual genomes. We found strong evidence of positive selection acting on genes and regulatory sequences, and we discovered that mutations in worker-biased proteins tend to have greater fitness effects than mutations in queen-biased proteins. We also found many instances of positive selection acting on genes that influence worker traits, suggesting that worker phenotypes represent a major vector for adaptation in social insects.

M.9 and M.26 rootstocks were planted in Estonia, Latvia and Lithuania in 2005. After five growing season, the strongest growth and the highest yield were recorded in Lithuania. The growth rate of trees on B.396, B.9, P62, P67, P66 and Pure 1 was similar to those of trees M.9. The rootstocks P22, P59 and PB-4 appeared to be more dwarfing than M.9. The highest cumulative yield of 'Auksis' was obtained from trees grafted on M.9, M.26, P62 and P67. The least productive were trees on PB-4 rootstock at all the places. Effects of rootstock on fruit weight were modest. Rootstock and location interaction was recorded for P61 in growth vigour control, and P22 and Pure 1 in cumulative yield.

Changes in gene regulation that underlie phenotypic evolution can be encoded directly in the DNA sequence or mediated by chromatin modifications such as DNA methylation. It has been hypothesized that the evolution of eusocial division of labor is associated with enhanced gene regulatory potential, which may include expansions in DNA methylation in the genomes of Hymenoptera (bees, ants, wasps, and sawflies). Recently, this hypothesis garnered support from analyses of a commonly used metric to estimate DNA methylation in silico, CpG content. Here, we test this hypothesis using direct, nucleotide-level measures of DNA methylation across nine species of Hymenoptera. In doing so, we generated new DNA methylomes for three species of interest, including one solitary and one facultatively eusocial halictid bee and a sawfly. We demonstrate that the strength of correlation between CpG content and DNA methylation varies widely among hymenopteran taxa, highlighting shortcomings in the utility of CpG content as a proxy for DNA methylation in comparative studies of taxa with sparse DNA methylomes. We observed strikingly high levels of DNA methylation in the sawfly relative to other investigated hymenopterans. Analyses of molecular evolution suggest the relatively distinct sawfly DNA methylome may be associated with positive selection on functional DNMT3 domains. Sawflies are an outgroup to all ants, bees, and wasps, and no sawfly species are eusocial. We find no evidence that either global expansions or variation within individual ortholog groups in DNA methylation are consistently associated with the evolution of social behavior.

Around 150 million years ago, eusocial termites evolved from within the cockroaches, 50 million years before eusocial Hymenoptera, such as bees and ants, appeared. Here, we report the first, 2GB genome of a cockroach, Blattella germanica, and the 1.3GB genome of the drywood termite, Cryptotermes secundus. We show evolutionary signatures of termite eusociality by comparing the genomes and transcriptomes of three termites and the cockroach against the background of 16 other eusocial and non-eusocial insects. Dramatic adaptive changes in genes underlying the production and perception of pheromones confirm the importance of chemical communication in the termites. These are accompanied by major changes in gene regulation and the molecular evolution of caste determination. Many of these results parallel molecular mechanisms of eusocial evolution in Hymenoptera. However, the specific solutions are remarkably different, thus revealing a striking case of convergence in one of the major evolu...

Termites have developed a wide array of defensive mechanisms. One of them is the mandibulate soldier caste that crushes or pierces their enemies. However, in several lineages of Termitinae, soldiers have long and slender mandibles that cannot bite but, instead, snap and deliver powerful strikes to their opponents. Here, we use morphological and molecular evidence to describe Roisinitermesebogoensis Scheffrahn, gen. & sp. n. from near Mbalmayo, Cameroon. Soldiers of R.ebogoensis are unique among all other kalotermitid soldiers in that they possess snapping mandibles. The imago of R.ebogoensis is also easily distinguished from all other Kalotermitidae by the lack of ocelli. Our study reveals a new case of parallel evolution of snapping mandibles in termites, a complex apparatus responsible of one of the fastest biological acceleration rates measured to date.

Olfactory/odorant receptors (ORs) probably govern eusocial behaviour in honey bees through detection of cuticular hydrocarbons (CHCs) and queen mandibular gland pheromones (QMP). CHCs are involved in nest-mate recognition whereas QMP acts as sex pheromone for drones and as retinue pheromone for female workers. Further studies on the effect of eusociality on the evolution of ORs are hindered by the non-availability of comprehensive OR sets of solitary species. We report complete OR repertoires from two solitary bees Dufourea novaeangliae (112 ORs) and Habropoda laboriosa (151 ORs). We classify these ORs into 34 phylogenetic clades/subfamilies. Differences in the OR sets of solitary and eusocial bees are observed in individual subfamilies like subfamily 9-exon (putative CHC receptors) and L (contains putative QMP receptor group). A subfamily (H) including putative floral scent receptors is expanded in the generalist honey bees only, but not in the specialists. On the contrary, subfami...

"Extreme Geography." 1997. California Geographer. 37:10-31. http://scholarworks.csun.edu/handle/10211.2/2690 URI: http://hdl.handle.net/10211.2/2690

A recent editorial in the Annals of the Association of American Geographers (Dixon and Jones, 1996) presented an entertaining deconstructive reading of scientific geography. This innovative essay, titled "For a Supercalifragilistkexpialidodous Scientific Geography" demonstrates how a popular deconstructive method adapted from the humanities might be deployed to facilitate the "self-redemption" of scientific geography from its constraining positivist closure, perhaps paving the way for its alignment with the open-ended and pluralistic world-view of an emerging critical poststructuralism.

Stylistic and methodological attributes of a postmodern attitude permeate the editorial: its postmodern style is a refreshing departure from the usual flat, denotative prose of scientific reporting; the blatant refusal of the authors to marshal validity and reference authority in support of their arguments is a portent of bold, new directions for the discipline's flagship journal; finally, the topic is thought-provoking and timely, motivated in part by the authors' concern about "the absence of a considered debate over the challenges and relative merits of poststructuralism" (Dixon and Jones, 1996,767). The timing of this debate cannot be separated from events and conditions that characterize the postmodern-as-epoch (Dear and Wassmansdorf, 1993). Perhaps that time has already passed: at least one observer suggests that the impact of information technology and time-space convergence on human communication seems to render Dixon and Jones' "considered debate" less relevant in the postmodern epoch (Nemeth, 1997).

Postmodern skepticism thrives on information surfeits. Ambiguous truths are bred wholesale by the speed in the span of cyberspace. This new flexibility in the accumulation of information now feeds a frenzy of popular skepticism about absolute truths that is much deeper and darker than the methodological skepticism in positivist science that continues to invest heavily in their credibilities.

Keyword: quantum thinking;
California Geographer ~ URI: http://hdl.handle.net/10211.2/2690

Within an insect society individuals are not equally related, which often leads to reproductive conflicts among them. This thesis investigates reproductive conflict in the honey bee. The main conflict investigated is that over male production. This is an important conflict as workers can lay unfertilised male eggs. In honey bees, this conflict is resolved by worker policing in which workers eat worker-laid eggs. Policing is selectively favoured because of the low average relatedness among workers due to polyandry. A naturally occurring anarchistic colony, in which workers evade normal policing and many workers' sons are reared, was investigated. About half the patrilines produced males, which caused a loss of inclusive fitness for the worker collective, contrary to previously described anarchistic colonies. In hopelessly queenless colonies, worker policing usually stops and the workers rear a last batch of males before the colony dies. Another atypical colony was investigated in which worker policing was not switched off despite being queenless. This caused a maladaptive phenotype in which many workers activated their ovaries and laid eggs that were not reared, leading to the colony producing no males before dying. Variation in worker policing was also investigated in normal queenright colonies. Large between colony variations occurred, and a lower egg policing rate in worker cells, where workers normally do not lay, could reduce the cost of mistaken removal of queen-laid eggs. Another potential conflict in honey bee societies concerns queen rearing. Workers may increase their inclusive fitness by favouring full-sister queens. Here I show that nepotism was absent in a previously unstudied context, the confinement of queens in their cells by workers. This confirms the general absence of nepotism in queen replacement, which is a largely cooperative process. Last, I investigated the learning abilities of bees towards cuticular hydrocarbons, which are thought to play a role in kin and nestmate recognition. Unsaturated alkenes are shown to be the most likely candidate compounds for recognition in honey bees. iii Biographical Sketch Nicolas Châline was born on the 2nd of March 1973 in Mantes la Jolie, France. His early education took place in Lisieux at the Marie Curie School and the Lycée Gambier. After his Baccalauréat in 1990, he spent three years in Rouen where he did his preparatory classes for the Biology "Grandes Ecoles" at the Lycée Pierre Corneille. He was then admitted to the National Agronomy Institute of Paris-Grignon in 1993, which trains Life engineers, and in the third year he chose to do an MSc in Population Biology, Genetics and Eco-Ethology (DEA) at Tours University. In 1997, he was called for his national service which he did in Martinique as an IT engineer. In August 1998, he moved to Luton, Beds, and then started working at the University of Sheffield in June 1999 in the Department of Animal and Plant Sciences as a research technician in the Sheffield Molecular Genetics Facility. He officially started his PhD on the 1 st of

The fauna of termites (Isoptera) preserved in Early Eocene amber from the Cambay Basin (Gujarat, India) are described and figured. Three new genera and four new species are recognized, all of them Neoisoptera-Parastylotermes krishnai Engel & Grimaldi, sp. n. (Stylotermitidae); Prostylotermes kamboja Engel & Grimaldi, gen. et sp. n. (Stylotermitidae?); Zophotermes Engel, gen. n., with Zophotermes ashoki Engel & Singh, sp. n. (Rhinotermitidae: Prorhinotermitinae); and Nanotermes isaacae Engel & Grimaldi, gen. et sp. n. (Termitidae: Termitinae?). Together these species represent the earliest Tertiary records of the Neoisoptera and the oldest definitive record of Termitidae, a family that comprises >75% of the living species of Isoptera. Interestingly, the affinities of the Cambay amber termites are with largely Laurasian lineages, in this regard paralleling relationships seen between the fauna of bees and some flies. Diversity of Neoisoptera in Indian amber may reflect origin of the amber deposit in Dipterocarpaceae forests formed at or near the paleoequator.

The fauna of termites (Isoptera) preserved in Early Eocene amber from the Cambay Basin (Gujarat, India) are described and figured. Three new genera and four new species are recognized, all of them Neoisoptera - Parastylotermes krishnai Engel & Grimaldi, sp. n. (Stylotermitidae); Prostylotermes kamboja Engel & Grimaldi, gen. et sp. n. (Stylotermitidae?); Zophotermes Engel, gen. n., with Zophotermes ashoki Engel & Singh, sp. n. (Rhinotermitidae: Prorhinotermitinae); and Nanotermes isaacae Engel & Grimaldi, gen. et sp. n. (Termitidae: Termitinae?). Together these species represent the earliest Tertiary records of the Neoisoptera and the oldest definitive record of Termitidae, a family that comprises >75% of the living species of Isoptera. Interestingly, the affinities of the Cambay amber termites are with largely Laurasian lineages, in this regard paralleling relationships seen between the fauna of bees and some flies. Diversity of Neoisoptera in Indian amber may reflect origin of t...

In eusocial insects, the ability to discriminate nest-mates from non-nest-mates is widespread and ensures that altruistic actions are directed towards kin and agonistic actions are directed towards non-relatives. Most tests of nest-mate recognition have focused on hymenopterans, and suggest that cooperation typically evolves in tandem with strong antagonism towards non-nest-mates. Here, we present evidence from a phylogenetically and behaviourally basal termite species that workers discriminate members of foreign colonies. However, contrary to our expectations, foreign intruders were the recipients of more rather than less cooperative behaviour and were not subjected to elevated aggression. We suggest that relationships between groups may be much more peaceable in basal termites compared with eusocial hymenoptera, owing to energetic and temporal constraints on colony growth, and the reduced incentive that totipotent workers (who may inherit breeding status) have to contribute to sel...

The Dufour gland in workers of vespine wasps appears as an unpaired tubiform gland that opens in close proximity to the sting base. The epithelial cells that line the central reservoir are characterized by apical microvillus-like projections and deep basal invaginations. Their cytoplasm contains a well-developed Golgi apparatus, numerous mitochondria, as well as strands of smooth endoplasmic reticulum. The Dufour gland duct occurs ventrally to the venom gland duct, and bends downward near the sting base to open in the dorsal vaginal wall. In this region, the duct is dorsoventrally flattened, and shows conspicuous bundles of parallel microtubules in the epithelial cells, that transmit the pulling forces of the myofilaments of the underlying muscular supply to the cuticle. This results in an active opening mechanism regulated by muscular contraction, while passive closure probably results from the return of the cuticular intima to a rest position.

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Extant research suggests that individuals employ traditional moral heuristics to support their observed altruistic behavior; yet findings have largely been limited to inductive extrapolation and rely on relatively few traditional frames in so doing, namely, deontology in organizational behavior and virtue theory in law and economics. Given that these and competing moral frames such as utilitarianism can manifest as identical behavior, we develop a moral framing instrument—the Philosophical Moral-Framing Measure (PMFM)—to expand and distinguish traditional frames associated and disassociated with observed altruistic behavior. The validation of our instrument based on 1015 subjects in 3 separate real stakes scenarios indicates that heuristic forms of deontology, virtue-theory, and utilitarianism are strongly related to such behavior, and that egoism is an inhibitor. It also suggests that deontic and virtue-theoretical frames may be commonly perceived as intertwined and opens the door ...

Phenotypic plasticity is important in adaptation and shapes the evolution of organisms. However, we understand little about what aspects of the genome are important in facilitating plasticity. Eusocial insect societies produce plastic phenotypes from the same genome, as reproductives (queens) and nonreproductives (workers). The greatest plasticity is found in the simple eusocial insect societies in which individuals retain the ability to switch between reproductive and nonreproductive phenotypes as adults. We lack comprehensive data on the molecular basis of plastic phenotypes. Here, we sequenced genomes, microRNAs (miRNAs), and multiple transcriptomes and methylomes from individual brains in a wasp (Polistes canadensis) and an ant (Dinoponera quadriceps) that live in simple eusocial societies. In both species, we found few differences between phenotypes at the transcriptional level, with little functional specialization, and no evidence that phenotype-specific gene expression is dr...

Queens of many social insect species are known to maintain reproductive monopoly by pheromonal signalling of fecundity. Queens of the primitively eusocial wasp Ropalidia marginata appear to do so using secretions from their Dufour's glands, whose hydrocarbon composition is correlated with fertility. Solitary nest foundresses of R. marginata are without nestmates; hence expressing a queen signal can be redundant, since there is no one to receive the signal. But if queen pheromone is an honest signal inextricably linked with fertility, it should correlate with fertility and be expressed irrespective of the presence or absence of receivers of the signal, by virtue of being a byproduct of the state of fertility. Hence we compared the Dufour's gland hydrocarbons and ovaries of solitary foundresses with queens and workers of post-emergence nests. Our results suggest that queen pheromone composition in R. marginata is a byproduct of fertility and hence can honestly signal fertility. This provides important new evidence for the honest signalling hypothesis.

The DAVID Gene Functional Classification Tool http://david.abcc.ncifcrf.gov uses a novel agglomeration algorithm to condense a list of genes or associated biological terms into organized classes of related genes or biology, called biological modules. This organization is accomplished by mining the complex biological co-occurrences found in multiple sources of functional annotation. It is a powerful method to group functionally related genes and terms into a manageable number of biological modules for efficient interpretation of gene lists in a network context.

This article is part of the 'More Fun Than Fun' column by Prof Raghavendra Gadagkar. He will explore interesting research papers or books and, while placing them in context, make them accessible to a wide readership. I recently had great fun reading a fascinating book, called Experiencing the Impossible: The Science of Magic (2019), by Gustav Kuhn. I might not have read the book but for the remarkable fact that Gustav Kuhn is both a professional scientist and a professional magician. He is a reader in psychology at Goldsmiths, a constituent college of the University of London, and has publishedin addition to this bookover 60 scholarly articles on the psychology and neurobiology of magic. He and his collaborators were the first to put volunteers into an fMRI machine while they were enjoying magic. They discovered that areas in the brains of their subjects involved in monitoring and managing conflict lit up preferentially. And of course, magic, as Kuhn explains in his book, involves a conflict between what we think is possible and what we think we have seen. In other studies, he has used eye-tracking technology to observe volunteers 'succumbing' to magic. His credentials as a magician are equally impressive: he is the president of the Science of Magic Association, a member of the Experimental Psychology Society and the Magic Circle. A young Gustav was hooked onto magic at 13 years, when his school-friend performed the trick of making an egg appear from behind his ear, and hasn't looked back. All science should be fun, but Kuhn and his students seem to have more than their fair share. This happens when you combine your hobby with your profession. His students learn to perform magic and show off their skills before children, fellow students and the public, all the while gathering data for their scientific papers and theses. I read the following excerpt (trimmed for length) from one of their recent papers with envy: "Sixty participants recruited on Goldsmiths University campus took

Eusocial insects are those that show overlap of generations, cooperative brood care and reproductive caste differentiation. Of these, primitively eusocial insects show no morphological differences between reproductive and worker castes and exhibit considerable flexibility in the social roles that adult females may adopt. This makes them attractive model systems for investigations concerning the origin of eusociality. The rapidly accumulating information on primitively eusocial wasps suggests that haplodiploidy is unlikely to have an important role in the origin of eusociality. General kin selection (without help from haplodiploidy) could however have been an important factor due to the many advantages of group living. Pre-imaginal caste bias leading to variations in fertility is also likely to have some role. Because workers often have some chance of becoming reproductives in future, mutualism and other individual selection models suggest themselves as important factors. A hypothesis for the route to eusociality which focuses on the factors selecting for group living at different stages in social evolution is presented. It is argued that group living originates owing to the benefit of mutualism (the 'Gambling Stage') but parental manipulation and subfertility soon become important (the 'Manipulation Stage') and finally the highly eusocial state is maintained because genetic asymmetries created by haplodiploidy are exploited by kin recognition (the 'Recognition Stage').