Yukihiko Toquenaga

Contrasting responses of bumble bees to feeding conspecifics on their familiar and unfamiliar flowers

Animals exploiting their familiar food items often avoid spatio-temporal aggregation with others by avoiding scents, less rewarding areas or visual contacts, thereby minimizing competition or interference when resources are replenished slowly in patches. When animals are searching or assessing available food sources, however, they may benefit from reducing sampling costs by following others at food sites. Therefore, animals may adjust their responses to others depending on their familiarity with foraging situations. Here, we conducted field experiments to test whether nectar-collecting bumble bees make this adjustment. We allowed free-foraging bees to choose between two inflorescences, one occupied by a conspecific bee and another unoccupied. When bees were presented with flowers of a familiar type, they avoided occupied inflorescences. In contrast, bees visited an occupied inflorescence when the flower type was unfamiliar. To our knowledge, this is the first report suggesting that animals adjust their responses to feeding conspecifics depending on their familiarity with food sources. Such behavioural flexibilities should allow foragers to both explore and exploit their environments efficiently.

Frpm artificial individuals to global patterns

Artificial Life is a model of biological systems that describes lives archived by computer simulation, chemical substrates or any other non-biological substrates. Artificial Life simulation adopts a bottom-up approach in which behavior of lower-level entities (e.g. molecules, cells and individuals) is all that is programed; global patterns (e.g. evolutionary patterns observed at the level of the population and the community) can emerge as a result of interaction among lower-level entities. Artificial Life simulations will be used not only to test ecological and evolutionary hypotheses explaining real organisms but also to show the validity of general theories, processes and concepts such as natural selection, theories of complexity, hierarchical relations and self-organization.

TEMPERATURE EFFECTS AND GENOTYPE-BY-ENVIRONMENT INTERACTIONS IN HYBRIDS: HALDANE'S RULE IN FLOUR BEETLES

When males of the flour beetle, Tribolium castaneum, are crossed to females of its close relative T. freemani, the sex ratio of the hybrids is female biased, owing in part to hybrid male mortality. Morphological abnormalities are also frequent in the surviving hybrid males, but not in the hybrid females. The finding that the heterogametic sex (male) is more adversely affected in interspecific crosses than the homogametic sex is consistent with Haldanes rule, which predicts that hybrid dysfunction should emerge as an indirect byproduct of divergent adaptation to differing environments. If so, environmental effects and genotype‐by‐environment interactions (GEI) should characterize the expression of Haldanes rule and interspecific hybrid traits in general. We used two wild‐collected populations of T. castaneum (from Infantes, Spain, and Madagascar) to investigate the effects of environmental variation on the expression of Haldanes rule. Males from each population were mated to several T. freemani females and the half‐sibling hybrid progenies were reared across a series of temperature regimes. For both populations, we found that hybrids raised at higher temperatures exhibited a more extreme expression of Haldanes rule: The hybrid sex ratios were more biased toward females and hybrid males had a much higher incidence of morphological abnormalities. The average response to temperature, the norm of reaction for Haldanes rule, varied between the two populations, and we found considerable and significant GEI for both hybrid traits within both populations. The evolutionary implications of these findings are discussed in the context of speciation arising as an indirect effect of local adaptation.

Contest and scramble competitions inCallosobruchus maculatus (Coleoptera: Bruchidae) II. Larval competition and interference mechanisms

Competition between contest and scramble strategists was examined using two strains ofCallosobruchus maculatus, the contest strain (iQ) and the scramble strain (tQ). The direct larval interference experiment within each strain showed that the contest strain performed strong direct interference throughout its larval stage against the conspecific opponent(s). The scramble strain also performed its interference, but only during the 3rd and the 4th larval stages. The inter-strain larval competition experiments inside a large and small mung bean (Vigna radiata) showed that the contest strain was, in general, superior to the scramble strain in competition, but the competition results were density- and frequency-dependent. In the large beans, the proportion of adult emergence of the contest strain increased with the increase of its own initial density but decreased as the initial density of the scramble strain increased. The shape of the proportion of adult emergence became nonlinear in the small beans. The cause of the complexity of inter-strain competition results was discussed in the light of the difference in larval interference ability between the two strains.

Contest and scramble competitions inCallosobruchus maculatus (Coleoptera: Bruchidae)

Larval competition curves and resource sharing patterns of 5 strains ofCallosobruchus maculatus (iQ, yQ, aaQ, wQ, and tQ) were examined. Offspring emergences as a function of the initial larval density were recorded to construct competition curves. Elytron length of emerged adults was used as the indicator of resource sharing patterns among competing larvae inside a bean. In the large beans, strain iQ showed a saturated competition curve and tQ strain showed a humped curve. Competition curves of the other 3 strains (yQ, aaQ, and wQ) were between those two extremes. In the small beans, strains iQ and tQ also showed a saturated and a humped competition curves, respectively, whereas the competition curves of the 3 intermediate scramble strains could not be distinguished from that of the iQ strain. Thus, the classification based on competition curves was sensitive to the resource condition (bean size). In both the large and the small beans, the elytron lengths of iQ strain remained constant irrespective of initial larval density. On the contrary, the elytron lengths of the 4 other strains decreased monotonically with higher initial larval density. Thus, the judgment based on the resource sharing pattern was shown to be robust. Only iQ strain should be designated as a contest type, and the remaining strains as scaramble types. Contest and scramble types inC. maculatus were also compared with those observed inC. analis andC. phaseoli using competition curves, resource sharing patterns, and other physiological characters.

Polymorphism of competition type and its genetics in Callosobruchus maculatus (Coleoptera: Bruchidae)

Callosobruchus maculatus has both contest and scramble competition strategies. The currently existing theoretical models using game theory suggest that the contest strategy should be selected for. However, most geographic strains of C. maculatus show scramble competition. We experimentally crossed the representative strains of contest and scramble. We expressed the degree of contest competition by a continuous value named the C-value, which ranges from zero (pure scramble) to unity (pure contest). The competition types expressed in the C-value were genetically additive. Their larval developmental rates were negatively correlated with C-values. Multiple-generation experiments of the mixed strains confirmed that there were no overwhelming advantages of contest over scramble type. Most of the mixed strains remained in the intermediate states. We discuss the results in terms of the resource size necessary for developmental success and developmental speed.

Global genetic differentiation in a cosmopolitan pest of stored beans: effects of geography, host-plant usage and anthropogenic factors.

Genetic differentiation can be promoted allopatrically by geographic isolation of populations due to limited dispersal ability and diversification over time or sympatrically through, for example, host-race formation. In crop pests, the trading of crops across the world can lead to intermixing of genetically distinct pest populations. However, our understanding of the importance of allopatric and sympatric genetic differentiation in the face of anthropogenic genetic intermixing is limited. Here, we examined global sequence variation in two mitochondrial and one nuclear genes in the seed beetle Callosobruchus maculatus that uses different legumes as hosts. We analyzed 180 samples from 42 populations of this stored bean pest from tropical and subtropical continents and archipelagos: Africa, the Middle East, South and Southeast Asia, Oceania and South America. For the mitochondrial genes, there was weak but significant genetic differentiation across continents/archipelagos. Further, we found pronounced differentiation among subregions within continents/archipelagos both globally and within Africa but not within Asia. We suggest that multiple introductions into Asia and subsequent intermixing within Asia have generated this pattern. The isolation by distance hypothesis was supported globally (with or without continents controlled) but not when host species was restricted to cowpeas Vigna unguiculata, the ancestral host of C. maculatus. We also document significant among-host differentiation both globally and within Asia, but not within Africa. We failed to reject a scenario of a constant population size in the recent past combined with selective neutrality for the mitochondrial genes. We conclude that mitochondrial DNA differentiation is primarily due to geographic isolation within Africa and to multiple invasions by different alleles, followed by host shifts, within Asia. The weak inter-continental differentiation is most likely due to frequent inter-continental gene flow mediated by human crop trade.

Egrets of a feather flock together

Group foraging and colonial formation in avian species are examined with an artificial life (alife) model using genetic algorithms (GA) and neural networks. Horns classical model predicts that colonial birds are more successful than territorial ones in a patchy resource environment, and the reverse is true when resources are evenly distributed. The weak point of the model is confusion between colony formation and flock foraging and implicit assumption of perfect knowledge of resource distribution by foraging birds. The authors made an alife model that realized both flock foraging and colonial formation simultaneously during evolution in a patchy environment. The reference organisms were egrets, which make colonial nests in the breeding season and colonial roosts in the nonbreeding season. In the computer model, artificial egrets used the presence of other egrets as an indicator to locate resource-rich patches in a resource-clumped environment. On the contrary, egrets were less reliant on other individuals as a source of foraging information in evenly distributed resource conditions. Colonial nesting was also induced only in a patchy environment where the foraging efficiency was always higher than the evenly distributed condition. Local enhancement played an important role in achieving colonial and flock foraging.

Do bumble bee queens choose nest sites to maximize foraging rate? Testing models of nest site selection

We proposed “foundress-max” hypothesis that a bumble bee foundress chooses her nest site to maximize her energy intake rate from nectar. To examine the hypothesis, we estimated the maximum energy intake rate at each site in the study area and compared the distribution of the maximum energy intake rates with those of actual nest sites. We also calculated rank correlations of the maximum energy intake rate with the number of nest-searching foundresses at 54 sites. The nest locations supported the foundress-max hypothesis, but the number of nest-searching foundresses did not. This could be attributed to the density of food sites: many food sites may attract many foundresses. Therefore, we subsequently proposed “foundress-sum” hypothesis that a foundress chooses her nest site to maximize the sum of energy intake rates. The nest locations supported the foundress-max hypothesis more than the foundress-sum hypothesis. A profitable food site would affect foundresses’ nest site selection.

Sewall wright meets artificial life: the origin and maintenance of evolutionary novelty

A decade ago, Langton coined the term Artificial Life (A-Life) to identify the new field of research that is attempting to create and characterize open-ended evolving systems using diverse computer-based methods. Fruitful interactions between A-Life research and that of conventional biological sciences (B-Life) are rare. Using the framework of molecular and evolutionary genetics, we discuss some of the reasons for this lack of conceptual cross-pollination between the disciplines and we identify some potential areas for interdisciplinary collaboration.