Gene duplication and the evolution of phenotypic diversity in insect societies

Gene duplication is an important evolutionary process thought to facilitate the evolution of phenotypic diversity. We investigated if gene duplication was associated with the evolution of phenotypic differences in a highly social insect, the honeybee Apis mellifera. We hypothesized that the genetic redundancy provided by gene duplication could promote the evolution of social and sexual phenotypes associated with advanced societies. We found a positive correlation between sociality and rate of gene duplications across the Apoidea, indicating that gene duplication may be associated with sociality. We also discovered that genes showing biased expression between A. mellifera alternative phenotypes tended to be found more frequently than expected among duplicated genes than singletons. Moreover, duplicated genes had higher levels of caste‐, sex‐, behavior‐, and tissue‐biased expression compared to singletons, as expected if gene duplication facilitated phenotypic differentiation. We also found that duplicated genes were maintained in the A. mellifera genome through the processes of conservation, neofunctionalization, and specialization, but not subfunctionalization. Overall, we conclude that gene duplication may have facilitated the evolution of social and sexual phenotypes, as well as tissue differentiation. Thus this study further supports the idea that gene duplication allows species to evolve an increased range of phenotypic diversity.     

Sampling methods for assessing social wasps species diversity in a eucalyptus plantation

Social wasps were collected in a eucalyptus (Eucalyptus urograndis, a genetically enhanced eucalyptus) plantation in Juiz de Fora, state of Minas Gerais, southwestern of Brazil, by using a variety of sampling methods, including active capture and baited traps set at two microhabitats. Six new records of social wasps for this region were obtained during the study: Brachygastra lecheguana Latreille, Polybia ignobilis Haliday, Polybia occidentalis Olivier, Polybia platycephala (Richards), Protonectarina sylveirae Saussure, and Protopolybia exigua Saussure (Hymenoptera: Vespidae). The active capture method sampled the greatest number of species. Baited traps installed in the canopy sampled agreater number of individuals of social wasps than those set on the trunks of eucalyptus trees. Fruit based baits captured a greater number of social wasp species than those baited with a protein derivative. Once no method was able to sample all species, we concluded that the efficiency of a survey can be maximized by using traps with a variety of different baits, in combination with active capture methods.     

Gene expression and molecular evolution

The combination of complete genome sequence information and estimates of mRNA abundances have begun to reveal causes of both silent and protein sequence evolution. Translational selection appears to explain patterns of synonymous codon usage in many prokaryotes as well as a number of eukaryotic model organisms (with the notable exception of vertebrates). Relationships between gene length and codon usage bias, however, remain unexplained. Intriguing correlations between expression patterns and protein divergence suggest some general mechanisms underlying protein evolution.     

Recombination and the evolution of coordinated phenotypic expression in a frequency-dependent game

A long standing question in evolutionary biology concerns the maintenance of adaptive combinations of traits in the presence of recombination. This problem may be solved if positive epistasis selects for reducing the rate of recombination between such traits, but this requires sufficiently strong epistasis. Here we use a model that we developed previously to analyze a frequency-dependent strategy game in asexual populations, to study how adaptive combinations of traits may be maintained in the presence of recombination when epistasis is too weak to select for genetic linkage. Previously, in the asexual case, our model demonstrated the evolution of adaptive associations between social foraging strategies and learning rules. We verify that these adaptive associations, which are represented by different two-locus haplotypes, can easily be broken by genetic recombination. We also confirm that a modifier allele that reduces the rate of recombination fails to evolve (due to weak epistasis). However, we find that under the same conditions of weak epistasis, there is an alternative mechanism that allows an association between traits to evolve. This is based on a genetic switch that responds to the presence of one social foraging allele by activating one of the two alternative learning alleles that are carried by all individuals. We suggest that such coordinated phenotypic expression by genetic switches offers a general and robust mechanism for the evolution of adaptive combinations of traits in the presence of recombination.     

The role of post-natal ontogeny in the evolution of phenotypic diversity in Podarcis lizards

Understanding the role of the developmental pathways in shaping phenotypic diversity allows appreciating in full the processes influencing and constraining morphological change. Podarcis lizards demonstrate extraordinary morphological variability that likely originated in short evolutionary time. Using geometric morphometrics and a broad suite of statistical tests, we explored the role of developmental mechanisms such as growth rate change, ontogenetic divergence/convergence/parallelism as well as morphological expression of heterochronic processes in mediating the formation of their phenotypic diversity during the post-natal ontogeny. We identified hypermorphosis - the prolongation of growth along the same trajectory - as the process responsible for both intersexual and interspecific morphological differentiation. Albeit the common allometric pattern observed in both sexes of any species constrains and canalizes their cephalic scales variation in a fixed portion of the phenotypic space, the extended growth experienced by males and some species allows them to achieve peramorphic morphologies. Conversely, the intrasexual phenotypic diversity is accounted for by non-allometric processes that drive the extensive morphological dispersion throughout their ontogenetic trajectories. This study suggests a model of how simple heterochronic perturbations can produce phenotypic variation, and thus potential for further evolutionary change, even within a strictly constrained developmental pathway.     

The evolution of worker caste diversity in social insects

Morphological diversification of workers is predicted to improve the division of labor within social insect colonies, yet many species have monomorphic workers. Individual-level selection on the reproductive capacities of workers may counter colony-level selection for diversification, and life-history differences between species (timing of caste determination, colony size, genetic variation available) may mediate the strength of this selection. We tested this through phylogenetically independent contrast analyses on a new data set for 35 ant species. Evidence was found that early divergence of queen-worker developmental pathways may facilitate the evolution of worker diversity because queen-worker dimorphism was strongly positively associated with diversity. By contrast, risks for colonies that invest in specialized workers and colony size effects on costs of worker reproduction seem unlikely to strongly affect the evolution of worker diversity because there was no significant association between colony size and diversity when controlling statistically for queen-worker dimorphism. Finally, worker diversity was greater in species with multiple lineages per colony, and it was negatively associated with relatedness in monogynous species. This could be due to high intracolonial genetic variance favoring the expression and evolution of great worker diversity or to diversity evolving more easily when there is selection for repression of worker reproduction (worker policing).     

Gene expression and the evolution of insect polyphenisms

Polyphenic differences between individuals arise not through differences at the genome level but as a result of specific cues received during development. Polyphenisms often involve entire suites of characters, as shown dramatically by the polyphenic castes found in many social insect colonies. An understanding of the genetic architecture behind polyphenisms provides a novel means of studying the interplay between genomes, gene expression and phenotypes. Here we discuss polyphenisms and molecular genetic tools now available to unravel their developmental bases in insects. We focus on several recent studies that have tracked gene-expression patterns during social insect caste determination. BioEssays 23:62-68, 2001. Published 2001 John Wiley & Sons, Inc.     

Visual sensitivity and foraging in social wasps

Summary While there is a distinction between that intensity of illumination which permits social wasps to forage, and that to which a sessile worker can respond, nevertheless illumination is the most critical of the environmental factors which control the activity of wasps. Low temperatures, high winds, and heavy rain all reduce activity but unless exceptionally severe do not wholly stop it. At dawn, when the critical level of illumination is attained, workers leave the nest, but at dusk they will not leave should the same critical level be due in the course of the foraging flight, after which they could not return.The three species of wasp,Vespula vulgaris, V. rufa, andV. germanica have a common threshold of illumination, although the hornet,Vespa crabro can forage in moonlight at an altogether lower illumination. Honey-bees normally need a still higher illumination than do wasps.In all these species, the thresholds of illumination are related to the length of the compound eyes, so that species with large eyes need less light by which to forage. Moreover, there is a slight difference between the threshold at dawn when workers leave the nest, and that at dusk, when they must needs have sufficient light by which to return. This difference is almost constant for each species, when, as is customary, one measures it on a logarithmic scale.Lastly, the estimates, which these experiments provide, of the threshold illuminations depend stochastically on the number of workers foraging. A correction for this bias is given.

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Sommaire Parmi les facteurs du milieu qui contrôlent l'activité des guêpes, celui de l'intensité de lumière est le plus important; toutefois on note une différence entre l'intensité de lumière qui permet aux guêpes sociales de fourrager, et celle qui produit une réponse des ouvrières sessiles.En général, les basses températures, les vents forts, et les grandes pluies réduisent leur activité, mais ces facteurs ne l'arrêtent pas complètement, à moins qu'ils ne soient très marqués.A l'aube, quand le niveau critique de lumière est atteint, les ouvrières quittent le guêpier, mais, le soir, si elles s'attendent à ce que la lumière vienne à s'abaisser au cours de leur sortie au-dessous du niveau critique, elles ne sortent pas.Les trois espèces de guêpe,Vespula vulgaris, V. rufa, etV. germanica, réagissent au même seuil de lumière, mais le frelon,Vespa crabro, est capable de fourrager au clair de lune par une lumière moins intense. Normalement, les abeilles exigent une lumière plus intense que les guêpes.Dans toutes ces espèces, le seuil de lumière se rapporte à la hauteur des yeux composés, par conséquent les espèces pourvues de grands yeux sont à même de fourrager par une lumière moins intense. De plus, il y a une légère différence entre le seuil de lumière à l'aube, quand les ouvrières quittent le guêpier, et celui du soir lorsqu'elles ont besoin d'une lumière suffisante pour rentrer. Cette différence, quand elle est mesurée à l'échelle logarithmique, comme il est d'usage, est presque constante pour chaque espèce.Enfin, les évaluations du seuil de lumière dans ces expériences dépendent stochastiquement du nombre d'ouvrières en train de fourrager. On a tenu compte de ce fait.

     

Euphorine phylogeny: the evolution of diversity in host-utilization by parasitoid wasps (Hymenoptera: Braconidae)

ABSTRACT.

• 1 New data on the phylogeny of the braconid subfamily Euphorinae supports the hypothesis that parasitism of adult insects by Euphorinae originated during parasitism of Chrysomelidae, a group whose larvae are ecologically coincident with adults.
• 2 Evolution of the habit of attacking the adult stage opened a new adaptive zone; subsequently the Euphorinae have diversified on to a phylogenetically greater variety of hosts than any other braconid subfamily.
• 3 Parasitism of eumastacid grasshoppers evolved from beetle parasitism in the tribe Perilitini.
• 4 The tribe Euphorini shows the greatest diversity of hosts utilized. Most attack Heteroptera; however, Chrysopopthorus diversified on to adult Chrysopidae, Euphoriella on to Psocoptera, and Cryptoxilos on to Scolytidae.
• 5 Parasitism of bark beetles (Scolytidae) has evolved independently in three genera: Cosmophorus, Cryptoxilos and Ropalophorus. This is the most specialized form of beetle parasitism by euphorines, since it involves direct parasitism of concealed hosts.
• 6 Parasitism of adult hymenopterans by the tribe Syntretini may be related to attacking hosts while they are foraging at flowers.
• 7 The pattern of diversification in the Euphorinae indicates several adaptive radiations within host orders, as well as a history of major host-shifts between phylogenetically distantly-related host groups: Coleoptera to Orthoptera; Coleoptera to Hymenoptera; Coleoptera to Heteroptera; Heteroptera to Neuroptera, Psocoptera, and back to Coleoptera. Both the‘host taxonomy’and‘host habitat’hypotheses of host-shifting are supported. Host-shifts have involved hosts occurring in the same micro-habitat and usually having similar feeding habits. This is consistent with current theory of host-location by means of host-produced kairomones and visual cues.

     

Preferential phenotypic association linked with cooperation in paper wasps

Animals can influence their social environment by preferentially associating with certain conspecifics. Such preferential association has gained increasing theoretical attention, as it may influence social evolution and population dynamics. However, relatively little empirical work has examined the occurrence of preferential association and its effects on cooperative group formation. Here, we test the factors associated with cooperative group formation in Polistes dominulus nest‐founding queen wasps. P. dominulus are a good system to study preferential association, as foundresses can nest alone or in groups and group membership is flexible. We found that both social and environmental factors were associated with partner choice. First, facial patterns were associated with cooperation. Wasps with more similar facial patterns were more likely to cooperate than wasps with less similar facial patterns. This preferential phenotypic association fits the theoretical criteria for the evolution of tag‐based cooperation. Season was also associated with cooperation; wasps on early‐season nests were more likely to cooperate than wasps on late‐season nests. High levels of aggression by nest owners during initial interactions were also correlated with lower probabilities of subsequent cooperation, suggesting that nest owners have some control over group membership. Other factors including body weight, weight similarity and nest productivity were not linked with cooperation. Overall, multiple factors influence cooperation in paper wasps, including facial pattern similarity. The occurrence of preferential phenotypic association in paper wasps is quite interesting and may influence the evolution of cooperation and population divergence in this group.     

Recognition and social dominance inPolistes wasps

It has been known for a long time that relationships among femalePolistes wasps are controlled by dominance through linear hierarchy. A classical model of the development of social hierarchies suggests that individual recognition intervenes to maintain the hierarchy established after the initial confrontation that takes place when two animals meet for the first time. This hypothesis was tested in the present study onPolistes gallicus (L.) foundresses. A tolerance index was calculated from the reactions of a dominant female when already known or unacquainted females were successively encountered. Results show that individualised social recognition is possible and that it takes more than 4 h of inter-individual contact for individual characteristics to be memorised.     

The geographic structure of selection on a coevolving interaction between social parasitic wasps and their hosts hampers social evolution

Social parasites exploit societies, rather than organisms, and rear their brood in social insect colonies at the expense of their hosts, triggering a coevolutionary process that may affect host social structure. The resulting coevolutionary trajectories may be further altered by selection imposed by predators, which exploit the abundant resources concentrated in these nests. Here, we show that geographic differences in selection imposed by predators affects the structure of selection on coevolving hosts and their social parasites. In a multiyear study, we monitored the fate of the annual breeding attempts of the solitary nesting foundresses of Polistes biglumis wasps in four geographically distinct populations that varied in levels of attack by the congeneric social parasite, P. atrimandibularis. Foundress fitness depended mostly on whether, during the long founding phase, a colony was invaded by social parasites or attacked by predators. Foundresses from each population differed in morphological traits and reproductive tactics that were consistent with selection imposed by their natural enemies and in ways that may affect host sociality. In turn, parasite traits were consistent with selection imposed locally by hosts, implying a geographic mosaic of coevolution in this brood parasitic interaction.     

Trade-offs between gene expression,growth and phenotypic diversity in microbial populations

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The evolution of gregariousness in parasitoid wasps

Data are assembled on the clutch-size strategies adopted by extant species of parasitoid wasp. These data are used to reconstruct the history of clutch-size evolution in the group using a series of plausible evolutionary assumptions. Extant families are either entirely solitary, both solitary and gregarious, or else clutch size is unknown. Parsimony analysis suggests that the ancestors of most families were solitary, a result which is robust to different phylogenetic relationships and likely data inadequacies. This implies that solitariness was ubiquitous throughout the initial radiation of the group, and that transitions to gregariousness have subsequently occurred a minimum of 43 times in several, but not all lineages. Current data suggest that species-rich and small-bodied lineages are more likely to have evolved gregariousness, and contain more species with small gregarious brood sizes. I discuss the implications of these data for clutch-size theory.     

Reconstruction ofRopalidia fasciata nests by progeny females after a typhoon and its significance in the social evolution of wasps

A strong typhoon, Typhoon 13, which swept through Okinawa on 2 and 3 September 1993, knocked down 57 to 61 of 97Ropalidia fasciata nests. Thirty-five colonies soon reconstructed their nests, and these nests produced 54±46 adults per nest during the remaining two and a half months before winter. The number of females involved in nest reconstruction was 21.3±15.7 per nest. The frequency of foraging activity was significantly higher in nests being reconstructed than in normal nests. As only one out of 205 foundresses marked in spring was found in early September, most cases of nest reconstruction were considered to be made exclusively by females that emerged during the spring and summer of 1993 (progeny females). All of the reconstructed nests but one produced adult females. Many females (31%) collected from the two reconstructed nests were inseminated. As all the reconstructed nests became vacant by early January, indicating completion of the colony cycle, and a large number of nests were established in the spring of 1994, females emerging from reconstructed nests in 1993 probably become foundresses in 1994.Ropalidia fasciata is considered to be, at least partially, bivoltine in Okinawa. The significance of these facts for the evolution of multi-queen social systems in the Polistinae is discussed.     

Gene expression and chemical diversity in hypothalamic neurosecretory neurons

Hypothalamic neurosecretory neurons transcribe, translate, store, and secrete a large number of chemical messengers. The neurons contain hypothalamic signal substances that regulate the secretion of anterior pituitary hormones as well as the neurohypophysial peptides vasopressin and oxytocin. In addition to the classical hypophysiotropic hormones, a large number of neuropeptides and classical transmitters of amine and amino acid nature are present in the same cells. This is particularly evident in the magnocellular neurons of the supraoptic and paraventricular nuclei, and in parvocellular neurons of the arcuate and paraventricular nuclei. The changes in gene expression induced by experimental manipulations and the colocalization chemical messengers in hypothalamic neurosecretory neurons and its possible significance is summarized in this review.     

Nonsiblicidal behavior and the evolution of clutch size in bethylid wasps

Parent-offspring conflict over clutch size may lead to siblicidal behavior between juveniles. In parasitoid wasps, selection for siblicide in small broods is predicted to produce a dearth of gregarious broods with few eggs. Here we document the clutch size distribution in the Bethylidae, a large family of aculeate parasitoids. Small gregarious clutches are the most common. Further data suggest that the most common gregarious clutches in the parasitoid Hymenoptera as a whole contain only a few eggs. Across bethylid species, both clutch size and wasp size increase with host size. Within genera clutch size is more closely related to host size, but between genera or larger clades wasp size is more closely related to host size. The volume of the emerging wasp brood does not depend on whether a species lays single- or multiple-egg clutches once host size is taken into account. These data suggest that clutch size in bethylid wasps is best described by traditional optimality models and that siblicide plays little role in this and possibly other families. We propose several ecological reasons for the rarity of siblicide in bethylids: ectoparasitism, idiobiosis, and a suite of characteristics associated with high within-brood relatedness.     

Figs and fig wasps: evolution in a microcosm

Fig wasps are tiny insects that both pollinate and feed upon fig plants. Each species requires the other in order to complete its reproductive cycle. The interaction centres on the unique structure of syconium (fig inflorescence), which provides an intriguing and convenient microcosm for studying the action of both natural and sexual selection.     

Evolutionary trends in the chromosome numbers of swarm-founding social wasps

Chromosome number and morphology are relevant aspects of genomic organization of eukaryotes and are considered key components to comprehend evolutionary mechanisms and karyotypic differentiation. Social wasps belonging to the tribe Epiponini show complex social characteristics that make these insects interesting models for genetic and evolutionary studies. However, there is a paucity of genetic information in social wasps as a whole. Aiming to investigate the process of chromosomal evolution of the social Epiponini wasps, chromosomes of ten species of this group were analyzed using Giemsa and fluorochrome staining techniques and fluorescence in situ hybridization in two species. In this study a high variation in the chromosome number and morphology was found and the previous range of chromosome number of Epiponini was broadened from n = 8–32 to n = 5–33. We also suggest that chromosomal segments with high GC content must have had a key role in the karyotype diversification of these wasps. Moreover, based on a phylogenetic background we find evidence of a main role of chromosomal fusion in the occurrence of gradual decrease of chromosome number in Epiponini during its chromosomal evolutionary history.