Hereditary nature of death-feigning frequency in a parasitoid wasp Heterospilus prosopidis using sib analysis.

Death feigning, a state of immobility observed in many animals in response to external stimuli, is an anti-predator behaviour. Although previous studies showed that death-feigning behaviours are quantitative genetic traits, the knowledge of the heritable basis of death-feigning behaviour is lacking. To investigate the heritable basis of death-feigning behaviour, we used three laboratory strains of a braconid parasitoid wasp, Heterospilus prosopidis. The heritable basis using half-sib analysis, and the effects of different geographical backgrounds, rearing conditions in the laboratory, and host age were evaluated. The results of the half-sib analysis showed that the frequency of death feigning varied among sires, suggesting a certain extent of additive genetic variance. Also, the frequency of death feigning varied between geographical backgrounds and among strains. Death-feigning frequency was not affected by the age of the host. Our findings highlight the importance of genetic factors underlying the basis of the death-feigning behaviour and provide support for the genetic alterations of traits from the perspective of evolution in various animal species.

Foraging predicts the evolution of warning coloration and mimicry in snakes.

Warning coloration and Batesian mimicry are classic examples of Darwinian evolution, but empirical evolutionary patterns are often paradoxical. We test whether foraging costs predict the evolution of striking coloration by integrating genetic and ecological data for aposematic and mimetic snakes (Elapidae and Dipsadidae). Our phylogenetic comparison on a total of 432 species demonstrated that dramatic changes in coloration were well predicted by foraging strategy. Multiple tests consistently indicated that warning coloration and conspicuous mimicry were more likely to evolve in species where foraging costs of conspicuous appearance were relaxed by poor vision of their prey, concealed habitat, or nocturnal activity. Reversion to crypsis was also well predicted by ecology for elapids but not for dipsadids. In contrast to a theoretical prediction and general trends, snakes' conspicuous coloration was correlated with secretive ecology, suggesting that a selection regime underlies evolutionary patterns. We also found evidence that mimicry of inconspicuous models (pitvipers) may have evolved in association with foraging demand for crypsis. These findings demonstrate that foraging is an important factor necessary to understand the evolution, persistence, and diversity of warning coloration and mimicry of snakes, highlighting the significance of additional selective factors in solving the warning coloration paradox.

Heritable effects on caste determination and colony-level sex allocation in termites under field conditions.

The ecological success of social insects is attributed to the division of labor, where newly hatched offspring differentiate into either fertile progeny or functionally sterile worker castes. There is growing evidence for the heritable (genetic or epigenetic) effects on caste determination based on laboratory experiments. Here, we indirectly demonstrate that heritable factors have the principal role in caste determination and strongly affect colony-level production of both sexes of fertile dispersers (i.e., alates) in field colonies of the termite Reticulitermes speratus. An egg-fostering experiment suggests that the colony-dependent sex-specific caste fates were almost entirely determined before oviposition. Our investigation of field colonies revealed that such colony-dependent sex-specific caste fates result in the intercolonial variation in the numerical sex ratio of differentiated fertile offspring and, eventually, that of alates. This study contributes to better understanding the mechanisms underlying the division of labor and life-history traits in social insects.

A novel phylogenetic comparative method for evaluating the strength of branch-specific directional selection.

Phylogenetic comparative methods (PCMs) have played a central role in studying the evolution of phenotypic traits. However, when a trait experienced directional selection, previous PCMs have faced a dilemma between mathematically tractable but restrictive models (i.e., simple Gaussian process models) and flexible but intractable approaches (i.e., a simulation-based process model of phenotype evolution built on population genetics frameworks). This paper proposes a novel Gaussian process macroevolutionary model, called the "branch-specific directional selection (BSDS)," for evaluating the strength of directional selection to reconcile these two approaches. This model is based on a second-order approximation of a previous simulation-based process model but has a closed-form likelihood function. This can also be extended to incorporate intraspecies variations and to linear mixed models, which are necessary for meta-analysis. We conduct numerical experiments to validate the proposed method and apply it to the brain volume of Hominidae species. The results show that the proposed methods yield statistically more reliable inferences and computational time is about hundred thousand times faster than the previous simulation-based methods. Further extensions of the BSDS model are expected to provide a clearer picture of the connection of microevolutionary processes and macroevolutionary patterns.

Spillover and spillback risks of ectoparasites by an invasive squirrel Callosciurus erythraeus in Kanto region of Japan.

Invasive organisms can alter host-parasite relationships in a given ecosystem by spreading exotic parasites and/or becoming a new reservoir for native ones. Since these problems affect management programs of the invasive host organisms, it is necessary to monitor them individually. The Pallas's squirrel Callosciurus erythraeus is an invasive arboreal mammal introduced into Japan that threatens to exacerbate ecological and public health problems by spreading native and exotic parasites. However, only limited surveys have been available especially for ectoparasites, using the traditional combing method in which the possibility of oversight is inherent. Here, we evaluated the ectoparasite occurrences in Kanto region of Japan, using the whole-shaving method as an alternative approach. As a result of examining 52 hosts from two invaded districts (Yokohama and Yokosuka), chigger mites (Leptotrombidium spp.) and fleas (Ceratophyllus anisus and Ceratophyllus indages indages) were newly recovered in addition to the previously reported tick (Haemaphysalis flava) and exotic lice (Enderleinellus kumadai and Neophaematoponis callosciuri). The parasite burdens were higher in Yokosuka and in male host individuals, affecting infracommunity richness and composition. Our findings on the variety of native and exotic ectoparasites, at higher abundances in some cases than previously known, may suggest that both the spillover and spillback risks need to be adjusted upwards.

The benefits of grouping as a main driver of social evolution in a halictine bee.

Over the past decade, the cause of sociality has been much debated. Inclusive fitness [br in Hamilton's rule (br - c > 0)] has been criticized but is still useful in the organization of a framework by elucidating mechanisms through which br (benefit × relatedness) becomes larger than c (cost). The bee Lasioglossum baleicum is suitable for investigation of this issue because of the sympatric occurrence of both social and solitary nesting in its populations. We show that a large part (approximately 92%) of the inclusive fitness of a eusocial worker can be attributed to the benefits of grouping. A 1.5-fold relatedness asymmetry benefit in singly mated haplo-diploids explains a small part (approximately 8.5%) of the observed inclusive fitness. Sociality enables this species to conduct foraging and nest defense simultaneously, which is not the case in solitary nests. Our results indicate that this benefit of grouping is the main source of the increased inclusive fitness of eusocial workers.

Activity of invasive slug Limax maximus in relation to climate conditions based on citizen's observations and novel regularization based statistical approaches.

Citizen science is a powerful tool that can be used to resolve the problems of introduced species. An amateur naturalist and author of this paper, S. Watanabe, recorded the total number of Limax maximus (Limacidae, Pulmonata) individuals along a fixed census route almost every day for two years on Hokkaido Island, Japan. L. maximus is an invasive slug considered a pest species of horticultural and agricultural crops. We investigated how weather conditions were correlated to the intensity of slug activity using for the first time in ecology the recently developed statistical analyses, Bayesian regularization regression with comparisons among Laplace, Horseshoe and Horseshoe+ priors for the first time in ecology. The slug counts were compared with meteorological data from 5:00 in the morning on the day of observation (OT- and OD-models) and the day before observation (DBOD-models). The OT- and OD-models were more supported than the DBOD-models based on the WAIC scores, and the meteorological predictors selected in the OT-, OD- and DBOD-models were different. The probability of slug appearance was increased on mornings with higher than 20-year-average humidity (%) and lower than average wind velocity (m/s) and precipitation (mm) values in the OT-models. OD-models showed a pattern similar to OT-models in the probability of slug appearance, but also suggested other meteorological predictors for slug activities; positive effect of solar radiation (MJ) for example. Five meteorological predictors, mean and highest temperature (°C), wind velocity (m/s), precipitation amount (mm) and atmospheric pressure (hPa), were selected as the effective factors for the counts in the DBOD-models. Therefore, the DBOD-models will be valuable for the prediction of slug activity in the future, much like a weather forecast.

Difference in evolutionary patterns of strongly or weakly selected characters among ant populations.

Despite being a central issue in evolutionary biology, few studies have examined the stasis of characters in populations with no gene flow. A possible mechanism of such stasis is stabilizing selection with similar peaks in each population. This study examined the evolutionary patterns of morphological characters with and without strong selection in ant populations. We show that compared to a character that seems to be less important, characters that are more important were less variable within and among populations. Microsatellite analyses showed significant genetic differences between populations, implying limited gene flow between them. The observed levels of genetic differentiation cannot be attributed to recent population separations. Thus, the observed differences in morphological variance seem to reflect the degree of selection on each character. The less important character changed proportionately with time, but such a pattern was not observed in more important characters. These results suggest that stabilizing selection maintains morphological stasis between populations of the same species with minimal gene flow independent of divergence times.