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PubMed2005

A secondary structural model of the 28S rRNA expansion segments D2 and D3 for Chalcidoid wasps (Hymenoptera: Chalcidoidea).

We analyze the secondary structure of two expansion segments (D2, D3) of the 28S ribosomal (rRNA)-encoding gene region from 527 chalcidoid wasp taxa (Hymenoptera: Chalcidoidea) representing 18 of the 19 extant families. The sequences are compared in a multiple sequence alignment, with secondary structure inferred primarily from the evidence of compensatory base changes in conserved helices of the rRNA molecules. This covariation analysis yielded 36 helices that are composed of base pairs exhibiting positional covariation. Several additional regions are also involved in hydrogen bonding, and they form highly variable base-pairing patterns across the alignment. These are identified as regions of expansion and contraction or regions of slipped-strand compensation. Additionally, 31 single-stranded locales are characterized as regions of ambiguous alignment based on the difficulty in assigning positional homology in the presence of multiple adjacent indels. Based on comparative analysis of these sequences, the largest genetic study on any hymenopteran group to date, we report an annotated secondary structural model for the D2, D3 expansion segments that will prove useful in assigning positional nucleotide homology for phylogeny reconstruction in these and closely related apocritan taxa.

Animals↗
PubMed2001

The phylogenetic analysis of variable-length sequence data: elongation factor-1alpha introns in European populations of the parasitoid wasp genus Pauesia (Hymenoptera: Braconidae: Aphidiinae).

Elongation factor-1alpha (EF-1alpha) is a highly conserved nuclear coding gene that can be used to investigate recent divergences due to the presence of rapidly evolving introns. However, a universal feature of intron sequences is that even closely related species exhibit insertion and deletion events, which cause variation in the lengths of the sequences. Indels are frequently rich in evolutionary information, but most investigators ignore sites that fall within these variable regions, largely because the analytical tools and theory are not well developed. We examined this problem in the taxonomically problematic parasitoid wasp genus Pauesia (Hymenoptera: Braconidae: Aphidiinae) using congruence as a criterion for assessing a range of methods for aligning such variable-length EF-1alpha intron sequences. These methods included distance- and parsimony-based multiple-alignment programs (CLUSTAL W and MALIGN), direct optimization (POY), and two "by eye" alignment strategies. Furthermore, with one method (CLUSTAL W) we explored in detail the robustness of results to changes in the gap cost parameters. Phenetic-based alignments ("by eye" and CLUSTAL W) appeared, under our criterion, to perform as well as more readily defensible, but computationally more demanding, methods. In general, all of our alignment and tree-building strategies recovered the same basic topological structure, which means that an underlying phylogenetic signal remained regardless of the strategy chosen. However, several relationships between clades were sensitive both to alignment and to tree-building protocol. Further alignments, considering only sequences belonging to the same group, allowed us to infer a range of phylogenetic relationships that were highly robust to tree-building protocol. By comparing these topologies with those obtained by varying the CLUSTAL parameters, we generated the distribution area of congruence and taxonomic compatibility. Finally, we present the first robust estimate of the European Pauesia phylogeny by using two EF-1alpha introns and 38 taxa (plus 3 outgroups). This estimate conflicts markedly with the traditional subgeneric classification. We recommend that this classification be abandoned, and we propose a series of monophyletic species groups.

Animals↗
PubMed1993

DNA insertions as a component of the evolution of unique satellite DNA families in two genera of parasitoid wasps: Diadromus and Eupelmus (Hymenoptera).

In the two parasitoid wasps, Diadromus collaris and Eupelmus orientalis, the satellite DNAs were each found to consist wholly or largely of a single family (5%-7% of the genome). Several clones of each family were obtained and sequenced. The repeat unit in each species is characterized by both the repetition of a basic motif and the presence of an inserted sequence. Sequence comparisons with satellite DNA from D. pulchellus and E. vuilleti provide plausible scenarios for the evolution of the satellite DNA in each genus. Palindromes and A-rich tracts in each consensus sequence suggest the formation, in vivo, of hairpin structures and bend centers that may play a role in heterochromatin condensation in insects. The insertions in the repeat units of each species also contain these structural features, suggesting that maintenance of these insertions requires constraints similar to those pertaining to the rest of the satellite-DNA unit.

Animals↗
PubMed2004

Queen and worker policing in monogynous and monandrous colonies of a primitively eusocial wasp.

Kin-selection theory predicts that a worker prefers to produce her own sons in a colony with monandry and monogyny because relatedness to her sons (0.5) and nephews (0.375) exceeds that to brothers (0.25). In spite of this prediction, recent studies reveal that workers police each other (mutual-worker egg removal) even in monandrous and monogynous colonies. We conducted field and laboratory studies to evaluate queen and worker policing in queen-right colonies of the primitively eusocial wasp Polistes chinensis antennalis. Genetic studies using microsatellite markers, as well as extensive observations of natural colonies, revealed that both queen and workers removed both queen- and worker-laid eggs in monogynous and monandrous colonies. The queen's eggs survived to hatching more successfully than those of the workers (88.5% versus 1.4%). We discuss the likely factors to explain these worker-policing behaviours.

Animals↗
PubMed2006

Kin competition promotes dispersal in a male pollinating fig wasp.

Despite theoretical predictions, there is little empirical evidence that kin competition avoidance promotes dispersal. We show that dispersal by male Platyscapa awekei pollinating fig wasps is promoted by both low returns in the natal fig and kin competition avoidance, with strategies depending on the interaction between phenotype (body size) and local conditions. We discuss the paucity of similar work, how males might assess conditions, and then contrast male dispersal and fighting behaviour. This indicates that differences in the scale at which behaviours affect competition can mean that they are the product of dissimilar selective forces even when they have the same recipients. More generally, this could explain why other social interactions are often mixtures of cooperation and conflict.

Animals↗
PubMed1995

Maternity assignment and queen replacement in a social wasp.

Assigning offspring to parents is important for understanding the evolution of reproductive conflicts and cooperation, particularly in the model systems represented by social insects. Molecular genetic markers are often used to exclude, and occasionally used to assign, candidate parents. However, their use in social insects has been unsatisfactory so far because candidate mothers are often highly related and candidate fathers are unknown. Here, we show that microsatellite loci can be scored from each mother's stored sperm permitting effective maternity assignment. The theoretical power of this method is huge, and we demonstrate its practical utilization in this large-scale study of the wasp, Polistes annularis. All 219 genotyped daughters were either assigned to a unique mother or shown to be the progeny of an uncollected dead mother. The data reveal an unexpectedly high number of changes in reproductive dominance. Maternity assignments using this method should help solve many difficult questions in social evolution.

Alleles↗
PubMed1997

Horizontal transmission of parthenogenesis-inducing microbes in Trichogramma wasps.

Complete parthenogenesis (thelytoky) in species of the parasitic wasp Trichogramma is usually caused by the cytoplasmically inherited bacterium Wolbachia. This symbiont induces gamete duplication, which, in these haplodiploid organisms, results in all-female broods. Antibiotic treatment "cures' this condition, restoring normal sexual reproduction. Phylogenetic analysis of Wolbachia has shown that, in contrast with the strains in other host organisms (where the symbiont also induces different reproductive alterations), those in Trichogramma form a monophyletic group. This might be an indication of symbiont-host cocladogenesis. To test this, we performed comparative molecular phylogenetics on 20 parthenogenetic Trichogramma cultures and their Wolbachiae. We conclude that there is, in fact, little evidence for cocladogenesis. Instead, the phylogenetic distribution of the symbionts appears to result from occasional horizontal transmission, which probably takes place inside the hosts of Trichogramma parasitoids (usually lepidopteran eggs). This study therefore suggests that parthenogenesis is not only curable, it can sometimes be contagious also.

Alphaproteobacteria↗
PubMed2000

Genetic support for the evolutionary theory of reproductive transactions in social wasps.

Recent evolutionary models of reproductive partitioning within animal societies (known as 'optimal skew', 'concessions' or 'transactional' models) predict that a dominant individual will often yield some fraction of the group's reproduction to a subordinate as an incentive to stay in the group and help rear the dominant's offspring. These models quantitatively predict how the magnitude of the subordinate's 'staying incentive' will vary with the genetic relatedness between dominant and subordinate, the overall expected group output and the subordinate's expected output if it breeds solitarily. We report that these predictions accord remarkably well with the observed reproductive partitioning between conesting dominant and subordinate queens in the social paper wasp Polistes fuscatus. In particular, the theory correctly predicts that (i) the dominant's share of reproduction, i.e. the skew, increases as the colony cycle progresses and (ii) the skew is positively associated both with the colony's productivity and with the relatedness between dominant and subordinate. Moreover, aggression between foundresses positively correlated with the skew, as predicted by transactional but not alternative tug-of-war models of societal evolution. Thus, our results provide the strongest (quantitative support yet for a unifying model of social evolution.

Animals↗
PubMed2001

Integration of a B chromosome into the A genome of a wasp.

B chromosomes are genome symbionts, the presence of which in many eukaryote species is explained, in most cases, by their violation of Mendelian rules, usually based on meiotic or mitotic instability, leading to their accumulation in the germ line (drive). However, B chromosome integration into the genome as a regular member of the chromosome set should imply the loss of drive. A possible way of bypassing this difficulty is to regularize meiosis when the B chromosome is frequent in the population, in order to yield gametes with one B chromosome. In diploid organisms, this task needs to be achieved in the two sexes, but in haplodiploids the problem simplifies to only the diploid sex. We have found, to the authors' knowledge, the first evidence of a B chromosome that is regularizing its meiotic behaviour and limiting its number to one B chromosome per haploid genome, the same dosage as the standard (A) chromosomes, in the solitary wasp Trypoxylon albitarse. It suggests a possible mechanism for B chromosome integration as a regular member of the chromosome complement.

Animals↗
PubMed2001

Recognition of social parasites as nest-mates: adoption of colony-specific host cuticular odours by the paper wasp parasite Polistes sulcifer.

Colonies of the polistine wasp Polistes dominulus are parasitized by the permanent worker-less social parasite Polistes sulcifer. After usurpation of the host colony, parasite females are characterized by a change in the relative proportions of their cuticular hydrocarbons to match those of the host species. In this paper we present evidence from field data and laboratory experiments that P. sulcifer females adopt a colony-specific host odour that facilitates their acceptance by host females of the usurped colony. Presentation experiments demonstrate that parasite females are recognized as foreign individuals by workers of other parasitized nests. We show that the modification of parasite cuticular compounds is sufficient for this recognition. This provides evidence that, after invasion, P. sulcifer queens do not require appeasement or propaganda substances for their acceptance by host colonies. Furthermore, multivariate discriminant analysis of the cuticular hydrocarbon proportions of the parasites after usurpation assigns the parasites together with P. dominulus females of their own host colony. To the authors' knowledge, this is the first confirmation that social parasites adopt colony-specific host odours.

Animals↗
PubMed2002

High reproductive skew in tropical hover wasps.

A plethora of recent models examines how genetic and environmental factors might influence partitioning of reproduction ('skew') in animal societies, but empirical data are sparse. We used three microsatellite loci to estimate skew on 13 nests of the Malaysian hover wasp, Liostenogaster flavolineata. Groups are small in L. flavolineata (1-10 females) and all females are capable of mating and laying eggs. Despite considerable variation between nests in parameters expected to influence skew, skew was uniformly high. On 11 of the 13 nests, all female eggs had been laid by a single dominant female. A second female had laid one to two out of 5-10 eggs respectively on the two remaining nests. A likelihood analysis suggested that on average, 90% of the male eggs had also been laid by the dominant. The slightly lower skew among male eggs might reflect the lower average relatedness of subordinates to male versus female offspring of the dominant. We suggest that high skew in L. flavolineata may result from strong ecological constraints and a relatively high probability that a subordinate will eventually inherit the dominant, egg-laying position.

Animals↗
PubMed2002

Integration of a B chromosome into the A genome of a wasp, revisited.

A previous study showed that in the haplodiploid solitary wasp Trypoxylon albitarse, most individuals carry one B chromosome per haploid genome, the same dosage as the standard (A) chromosomes, indicating a possible regularization of B-chromosome meiotic behaviour and its integration into the A genome. In a new sampling, we have analysed 15 populations (including 9 out of the 10 previously analysed) to test the evolution of this integration process. The new results provide a direct report of the invasion process in the Porto Firme population, where B frequency has dramatically increased in only four generations. In the populations from the Viçosa region, however, B frequency has remained stable, although the principal B type, the metacentric one, has increased in frequency at the expense of the acrocentric one in several populations. The implications of these new results on the hypothesis of the integration of these B chromosomes, as regular members of the A genome, are discussed.

Animals↗
PubMed2002

Mating system and sex ratios of a pollinating fig wasp with dispersing males.

Recent studies have used sex ratios to quantify the mating systems of organisms, the argument behind it being that more female-biased sex ratios are an indication of higher local mate competition, which goes hand-in-hand with higher levels of inbreeding. Although qualitative tests of the effects of mating systems on sex ratios abound, there is a dearth of studies that quantify both the mating system and the sex ratio. I use a colour dimorphism with a simple Mendelian inheritance to quantify the mating system of an unusual fig-pollinating wasp in which males disperse to obtain matings on non-natal mating patches. In qualitative agreement with initial expectations, the sex ratios of single foundresses are found to be higher than those of regular species. However, by quantifying the mating system, it is shown that the initial expectation is incorrect and this species' sex ratio is a poor predictor of its mating system (it underestimates the frequency of sib-mating). The species has a very high variance in sex ratio suggesting that excess males can simply avoid local mate competition (and hence a lowered fitness to their mother) by dispersing to other patches.

Animals↗
PubMed2003

Ultrastructure meets reproductive success: performance of a sphecid wasp is correlated with the fine structure of the flight-muscle mitochondria.

Organisms show a remarkable inter-individual variation in reproductive success. The proximate causes of this variation are not well understood. We hypothesized that the ultrastructure of costly or complex tissues or organelles might affect reproductive performance. We tested this hypothesis in females of a sphecid wasp, the European beewolf, Philanthus triangulum (Hymenoptera, Sphecidae), that show considerable variation in reproductive success. The most critical component of reproduction in beewolf females is flying with paralysed honeybees, which more than double their weight. Because of the high energetic requirements for flight, we predicted that the ultrastructure of the flight-muscle mitochondria might influence female success. We determined the density of mitochondria and the density of the inner mitochondrial membranes (DIMM) of the flight muscles as well as age, body size and fat content. Only DIMM had a significant influence on female reproductive success, which might be mediated by an elevated adenosine triphosphate (ATP) supply. The variation in DIMM might result from differences in larval provisions or from an accumulation of mutations in the mitochondrial genome. Our results support the hypothesis that the organization of complex structures contributes to inter-individual variation in reproductive success.

Adenosine Triphosphate↗
PubMed2005

Direct assessment of queen quality and lack of worker suppression in a paper wasp.

Assessing a conspecific's potential is often crucial to increase one's fitness, e.g. in female choice, contests with rivals or reproductive conflicts in animal societies. In the latter, helpers benefit from accurately assessing the fertility of the breeder as an indication of inclusive fitness. There is evidence that this can be achieved using chemical correlates of reproductive activity. Here, we show that queen quality can be assessed by directly monitoring her reproductive output. In the paper wasp Polistes dominulus, we mimicked a decrease in queen fertility by regularly removing brood. This triggered ovarian development and egg-laying by many workers, which strongly suggests that brood abundance is a reliable cue of queen quality. Brood abundance can be monitored when workers perform regular brood care in small size societies where each brood element is kept in a separate cell. Our results also show that although the queen was not manipulated, and thus remained healthy and fully fertile, she did not control worker egg-laying. Nevertheless, when workers laid eggs, the queen secured a near reproductive monopoly by selectively destroying these eggs, a mechanism known as 'queen policing'. By contrast, workers destroyed comparatively few queen-laid eggs, but did destroy each other's eggs.

Animals↗
PubMed2005

60 million years of co-divergence in the fig-wasp symbiosis.

Figs (Ficus; ca 750 species) and fig wasps (Agaoninae) are obligate mutualists: all figs are pollinated by agaonines that feed exclusively on figs. This extraordinary symbiosis is the most extreme example of specialization in a plant-pollinator interaction and has fuelled much speculation about co-divergence. The hypothesis that pollinator specialization led to the parallel diversification of fig and pollinator lineages (co-divergence) has so far not been tested due to the lack of robust and comprehensive phylogenetic hypotheses for both partners. We produced and combined the most comprehensive molecular phylogenetic trees to date with fossil data to generate independent age estimates for fig and pollinator lineages, using both non-parametric rate smoothing and penalized likelihood dating methods. Molecular dating of ten pairs of interacting lineages provides an unparalleled example of plant-insect co-divergence over a geological time frame spanning at least 60 million years.

Animals↗
PubMed2006

Differential gene expression and phenotypic plasticity in behavioural castes of the primitively eusocial wasp, Polistes canadensis.

Understanding how a single genome can produce a variety of different phenotypes is of fundamental importance in evolutionary and developmental biology. One of the most striking examples of phenotypic plasticity is the female caste system found in eusocial insects, where variation in reproductive (queens) and non-reproductive (workers) phenotypes results in a broad spectrum of caste types, ranging from behavioural through to morphological castes. Recent advances in genomic techniques allow novel comparisons on the nature of caste phenotypes to be made at the level of the genes in organisms for which there is little genome information, facilitating new approaches in studying social evolution and behaviour. Using the paper wasp Polistes canadensis as a model system, we investigated for the first time how behavioural castes in primitively eusocial insect societies are associated with differential expression of shared genes. We found that queens and newly emerged females express gene expression patterns that are distinct from each other whilst workers generally expressed intermediate patterns, as predicted by Polistes biology. We compared caste-associated genes in P. canadensis with those expressed in adult queens and workers of more advanced eusocial societies, which represent four independent origins of eusociality. Nine genes were conserved across the four taxa, although their patterns of expression and putative functions varied. Thus, we identify several genes that are putatively of evolutionary importance in the molecular biology that underlies a number of caste systems of independent evolutionary origin.

Adaptation, Physiological↗