SciCombinator

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Concept: Wasp

175

It is thought that speciation in phytophagous insects is often due to colonization of novel host plants, because radiations of plant and insect lineages are typically asynchronous. Recent phylogenetic comparisons have supported this model of diversification for both insect herbivores and specialized pollinators. An exceptional case where contemporaneous plant-insect diversification might be expected is the obligate mutualism between fig trees (Ficus species, Moraceae) and their pollinating wasps (Agaonidae, Hymenoptera). The ubiquity and ecological significance of this mutualism in tropical and subtropical ecosystems has long intrigued biologists, but the systematic challenge posed by >750 interacting species pairs has hindered progress toward understanding its evolutionary history. In particular, taxon sampling and analytical tools have been insufficient for large-scale cophylogenetic analyses. Here, we sampled nearly 200 interacting pairs of fig and wasp species from across the globe. Two supermatrices were assembled: on an average, wasps had sequences from 77% of 6 genes (5.6 kb), figs had sequences from 60% of 5 genes (5.5 kb), and overall 850 new DNA sequences were generated for this study. We also developed a new analytical tool, Jane 2, for event-based phylogenetic reconciliation analysis of very large data sets. Separate Bayesian phylogenetic analyses for figs and fig wasps under relaxed molecular clock assumptions indicate Cretaceous diversification of crown groups and contemporaneous divergence for nearly half of all fig and pollinator lineages. Event-based cophylogenetic analyses further support the codiversification hypothesis. Biogeographic analyses indicate that the present-day distribution of fig and pollinator lineages is consistent with a Eurasian origin and subsequent dispersal, rather than with Gondwanan vicariance. Overall, our findings indicate that the fig-pollinator mutualism represents an extreme case among plant-insect interactions of coordinated dispersal and long-term codiversification. [Biogeography; coevolution; cospeciation; host switching; long-branch attraction; phylogeny.].

Concepts: Evolution, Biology, Insect, Phylogenetics, Pollination, Ficus, Wasp, Fig wasp

171

Guarding behavior is an important activity in sub-social insects, and this behavior is believed to improve the survival of offspring. Sclerodermus harmandi (Hymenoptera: Bethylidae) is one of most powerful epizoic parasitoid wasps, and it parasitizes Monochamus alternatus, a borer of wood and also the primary vector of the pinewood nematode Bursaphelenchus xylophilus. After laying eggs, S. harmandi exhibits sub-social behavior involving the female tending the clutch of eggs until emergence (guarding behavior). In this study, the benefits of this maternal care with regard to improvements in the survival of offspring were examined. During the developmental stages, only offspring in the egg and larval stages were sensitive to guarding behavior. A positive relationship between the survival of the offspring and the duration of guarding was detected with logistic regression analysis. A female replacement experiment demonstrated that multiparous S. harmandi stepmothers showed guarding behavior and that this behavior improved the survival of the immature offspring, whereas nulliparous stepmothers failed to exhibit the guarding behavior. These results indicate that S. harmandi females display maternal care and that this behavior improves the survival of offspring.

Concepts: Better, Regression analysis, Logistic regression, Improve, Insect, Nematodes, Wasp, Bursaphelenchus xylophilus

146

Mutualisms play a key role in most ecosystems, yet the mechanisms that prevent overexploitation of the mutualistic relationship are still poorly understood. In the mutualism between fig trees and their pollinating wasps both partners depend on each other. Fig trees benefit from female wasps that disperse their pollen, whereas wasps frequently benefit from a higher ratio of male offspring. Here we use manipulative field experiments to address whether host trees (Ficus racemosa) can influence the offspring sex ratio of the pollinator wasp. We controlled wasp matings; virgin wasps can lay only male eggs. We found that virgin foundress wasps had fewer offspring than mated foundresses. This was not caused by virgin wasps having a shorter lifespan, or laying fewer eggs. Instead, male wasp larvae were more likely to die during development. Additionally, male eggs were deposited in flowers of equal style length to those of female eggs, yet emerged from galls with shorter pedicels than those of female wasps. We suggest that male larvae are either allocated less resources by the tree, or are less able to attract resources, during development. If the tree orchestrates this difference it would promote a more female-biased wasp brood, thus increasing the tree’s fitness.

Concepts: Female, Insect, Symbiosis, Sex, Wasp, Fig wasp, Wasps, Resource allocation

49

The current study explored the role of valence and self-relevance in size perception of neutral and aversive animals. In Experiment 1, participants who were highly fearful of spiders and control (low fear of spiders) participants rated the size and unpleasantness of spiders and other neutral animals (birds and butterflies). We found that although individuals with both high and low fear of spiders rated spiders as highly unpleasant, only the highly fearful participants rated spiders as larger than butterflies. Experiment 2 included additional pictures of wasps (not self-relevant, but unpleasant) and beetles. The results of this experiment replicated those of Experiment 1 and showed a similar bias in size estimation for beetles, but not for wasps. Mediation analysis revealed that in the high-fear group both relevance and valence influenced perceived size, whereas in the control group only valence affected perceived size. These findings suggest that the effect of highly relevant stimuli on size perception is both direct and mediated by valence.

Concepts: Scientific method, Insect, Wasp, Spider, Spiders, Phobia, Itsy Bitsy Spider, Itsy Bitsy Teenie Weenie Yellow Polka Dot Bikini

42

Society is increasingly concerned with declining wild bee populations. Although most bees nest in the ground, considerable effort has centered on installing ‘bee hotels’-also known as nest boxes or trap nests-which artificially aggregate nest sites of above ground nesting bees. Campaigns to ‘save the bees’ often promote these devices despite the absence of data indicating they have a positive effect. From a survey of almost 600 bee hotels set up over a period of three years in Toronto, Canada, introduced bees nested at 32.9% of sites and represented 24.6% of more than 27,000 total bees and wasps recorded (47.1% of all bees recorded). Native bees were parasitized more than introduced bees and females of introduced bee species provisioned nests with significantly more female larva each year. Native wasps were significantly more abundant than both native and introduced bees and occupied almost ¾ of all bee hotels each year; further, introduced wasps were the only group to significantly increase in relative abundance year over year. More research is needed to elucidate the potential pitfalls and benefits of using bee hotels in the conservation and population dynamics of wild native bees.

Concepts: Insect, Population, Larva, Bee, Wasp, Nest, Abundance of the chemical elements, Nest box

38

Change in land cover is thought to be one of the key drivers of pollinator declines, and yet there is a dearth of studies exploring the relationships between historical changes in land cover and shifts in pollinator communities. Here, we explore, for the first time, land cover changes in England over more than 80 years, and relate them to concurrent shifts in bee and wasp species richness and community composition. Using historical data from 14 sites across four counties, we quantify the key land cover changes within and around these sites and estimate the changes in richness and composition of pollinators. Land cover changes within sites, as well as changes within a 1 km radius outside the sites, have significant effects on richness and composition of bee and wasp species, with changes in edge habitats between major land classes also having a key influence. Our results highlight not just the land cover changes that may be detrimental to pollinator communities, but also provide an insight into how increases in habitat diversity may benefit species diversity, and could thus help inform policy and practice for future land management.

Concepts: Pollinator decline, Bee, Wasp, Pollinator, Species diversity, Species richness

36

Hosts have numerous defenses against parasites, of which behavioral immune responses are an important but underappreciated component. Here we describe a behavioral immune response that Drosophila melanogaster uses against endoparasitoid wasps. We found that when flies see wasps, they switch to laying eggs in alcohol-laden food sources that protect hatched larvae from infection. This change in oviposition behavior, mediated by neuropeptide F, is retained long after wasps are removed. Flies respond to diverse female larval endoparasitoids but not to males or pupal endoparasitoids, showing that they maintain specific wasp search images. Furthermore, the response evolved multiple times across the genus Drosophila. Our data reveal a behavioral immune response based on anticipatory medication of offspring and outline a nonassociative memory paradigm based on innate parasite recognition by the host.

Concepts: Immune system, Bacteria, Insect, Larva, Parasitism, Drosophila, Wasp, Ant

29

Figs (Ficus) have a reciprocally obligate mutualism with tiny, short-lived (1-2 days) fig wasps (Agaonidae). The small size and short life of these pollinators is expected to make them more vulnerable to climate change than their larger and longer-lived hosts. We experimentally tested the thermal tolerances of four species of adult female fig wasp from equatorial Singapore. The results suggest that an increase of 3°C or more above the current temperatures experienced across much of the equatorial tropics would markedly decrease the active adult lifespan of all four species. Fig plants are the centre of an intricate web of specialist and generalist animals. Unless fig wasps can acclimate or adapt to warmer temperatures in time, these responses may disrupt the mutualism, potentially affecting multiple trophic levels.

Concepts: Plant, Ecology, Ficus, Wasp, Ant, Pollinators, Fig wasp, Global warming

29

The golden paper wasp is a social insect whose colony members have the remarkable ability to recognise each others' faces. New research shows that this species is singularly skilled at learning about faces, opening interesting perspectives on convergent evolution of specialist cognitive abilities in insects and vertebrates.

Concepts: Psychology, Evolution, Insect, Honey bee, Face perception, Wasp, Ant, Hymenoptera

27

Living in seasonally changing environments requires adaptation to seasonal cycles. Many insects use the change in day length as a reliable cue for upcoming winter and respond to shortened photoperiod through diapause. In this study, we report the clinal variation in photoperiodic diapause induction in populations of the parasitoid wasp Nasonia vitripennis collected along a latitudinal gradient in Europe. In this species, diapause occurs in the larval stage and is maternally induced. Adult Nasonia females were exposed to different photoperiodic cycles and lifetime production of diapausing offspring was scored. Females switched to the production of diapausing offspring after exposure to a threshold number of photoperiodic cycles. A latitudinal cline was found in the proportion of diapausing offspring, the switch point for diapause induction measured as the maternal age at which the female starts to produce diapausing larvae, and the critical photoperiod for diapause induction. Populations at northern latitudes show an earlier switch point, higher proportions of diapausing individuals and longer critical photoperiods. Since the photoperiodic response was measured under the same laboratory conditions, the observed differences between populations most likely reflect genetic differences in sensitivity to photoperiodic cues, resulting from local adaptation to environmental cycles. The observed variability in diapause response combined with the availability of genomic tools for N. vitripennis represent a good opportunity to further investigate the genetic basis of this adaptive trait.

Concepts: Genetics, Insect, Developmental biology, Larva, Lepidoptera, Polymorphism, Wasp, Photoperiodism