Takayoshi Nishida

Seasonal variation in elytral colour polymorphism in Harmonia axyridis (the ladybird beetle): the role of non-random mating

Seasonal variation in the frequency of elytral colour polymorphism in the ladybird beetle, Harmonia axyridis, was analysed through the combination of field observations and laboratory mate-choice experiments. Field observations involving daily mark-recapture of a local population revealed an overall preponderance of melanic morphs in the population, but the proportion of non-melanics notably increased from spring to summer due to their higher frequency of mating among different morphs of the spring generation. In the laboratory experiments, both melanic and non-melanic females preferentially chose non-melanic males in spring, but non-melanics were less successful at mating than melanics in summer. Therefore, the relative frequencies of melanics and non-melanics within a population vary throughout the seasons, with non-melanics increasing in proportion towards summer and decreasing thereafter. Laboratory experiments revealed that non-random mating among morphs is the basis of this variation, with females and, to a lesser degree, males being involved in the process of mate selection. The results of an artificial colour-change experiment indicate that male elytral colour is an important factor in female mate selection but other factors such as behaviour and chemical cues must also be involved.

Adaptive significance of death feigning posture as a specialized inducible defence against gape-limited predators

Death feigning is fairly common in a number of taxa, but the adaptive significance of this behaviour is still unclear and has seldom been tested. To date, all proposed hypotheses have assumed that prey manage to escape predation by sending a death-mimicking signal, although death-feigning postures are markedly different from those of dead animals. Moreover, the efficacy of this technique may largely depend on the foraging mode of the predator; death feigning seldom works with sit-and-wait predators that make the decision to attack and consume prey within a very brief time. We examined whether death feigning in the pygmy grasshopper Criotettix japonicus Haan was an inducible defence behaviour against the frog Rana nigromaculata, a sit-and-wait, gape-limited predator. The characteristic posture assumed by the grasshopper during death feigning enlarges its functional body size by stretching each of three body parts (pronotum, hind legs and lateral spines) in three different directions, thereby making it difficult for the predator to swallow the prey. Our result is the first consistent explanation for why death-mimicking animals do not always mimic the posture of dead animals.

Asymmetric reproductive interference between specialist and generalist predatory ladybirds

1. Closely related species often differ greatly in the quality and breadth of resources exploited, but the actual mechanisms causing these differences are poorly understood. Because in the laboratory specialized species often survive and perform as well or better on host species that are never utilized in nature, negative ecological interactions restricting host range must exist. Here, we focused on reproductive interference, which has been theoretically predicted to drive niche separation between closely related species with overlapping mating signals. 2. We examined the interspecific sexual interactions in relation to ecological specialization and generalization in two sibling ladybird species, Harmonia yedoensis and Harmonia axyridis. Harmonia yedoensis is a specialist predator that preys only on pine aphids, which are highly elusive prey for ladybird hatchlings, whereas H. axyridis is a generalist predator with a broad prey and habitat range. 3. We experimentally showed that conspecific sperm fertilized the vast majority of eggs regardless of mating order (i.e. conspecific sperm precedence) when a female of H. yedoensis or H. axyridis mated with both a conspecific and a heterospecific male. Moreover, we demonstrated that mating opportunities of H. yedoensis females strongly decreased as heterospecific density increased relative to conspecific density. In contrast, in H. axyridis, female mating success was high regardless of conspecific or heterospecific density. 4. Our results suggest that the generalist H. axyridis should be dominant to the specialist H. yedoensis in terms of reproductive interference. Our results support the hypothesis that asymmetric reproductive interference from the dominant species may force the non-dominant species to become a specialist predator that exclusively utilizes less preferred prey in nature.

The mechanism of the fecundity reduction in Callosobruchus maculatus caused by Callosobruchus chinensis males

Male promiscuity sometimes results in interspecific reproductive interaction, also known as reproductive interference. Reproductive interference entails costs for the individuals involved and affects the community structure by reducing the population growth rate. However, our understanding of the mechanisms generating reproductive interference is still insufficient. Two congeneric bean weevils, Callosobruchus chinensis and C. maculatus, show asymmetric reproductive interference; only C. chinensis males reduce the fecundity of the other species. Here we investigated the mechanism of reproductive interference by C. chinensis males on C. maculatus females in terms of lifetime fecundity. Callosobruchus chinensis males with ablated genitals, which could harass C. maculatus females but not copulate with them, did not reduce the C. maculatus fecundity, suggesting that interspecific copulation was necessary. However, a single interspecific copulation did not affect C. maculatus fecundity as long as the females also copulated with a conspecific male. Exposure to C. chinensis males for 24xa0h prior to oviposition significantly reduced C. maculatus fecundity, and fecundity was negatively correlated with the number of C. chinensis males the females were exposed to. Additionally, C. maculatus females experienced more interspecific copulations when they were housed with more C. chinensis males. Together these findings suggest that multiple interspecific copulations by C. chinensis males reduce the fecundity of C. maculatus females. Thus in general, even if a single interspecific copulation is apparently harmless, repeated interspecific copulations can be costly for the individuals involved. Furthermore, only by quantifying reproductive success were we able to identify the precise mechanism of reproductive interference.

Prey capture performance in hatchlings of two sibling Harmonia ladybird species in relation to maternal investment through sibling cannibalism

1. To elucidate the factors responsible for the evolution of host specialisation, prey capture performance in hatchlings of two sibling ladybird species, Harmonia yedoensis and H. axyridis, feeding on four aphid species was examined. Harmonia yedoensis is a specialist predator that preys only on pine aphids in the field, whereas H. axyridis is a generalist predator with a broad prey range.

Conflicting intersexual mate choices maintain interspecific sexual interactions

Reproductive interference, interspecific sexual interactions that affect reproductive success, is found in various taxa and has been considered as a fundamental source of reproductive character displacement (RCD). Once RCD has occurred, persistent interspecific sexual interactions between species pairs are expected to diminish. However, reproductive interference has been reported from some species pairs that sympatrically coexist. Thus, the question arises, can reproductive interference persist even after RCD? We modeled the evolutionary dynamics of signal traits and mate recognition that determine whether interspecific sexual interactions occur. Our models incorporate male decision making based on the recognition of signal traits, whereas most previous models incorporate only female decision making in mate selection. Our models predict the following: (1) even when male decision making is incorporated, males remain promiscuous; (2) nevertheless, the frequency of interspecific mating is maintained at a low level after trait divergence; (3) the rarity of interspecific mating is due to strict female mate recognition and the consequent refusal of interspecific courtship by females; and (4) the frequency of interspecific mating becomes higher as the cost to females of refusing interspecific courtship increases. These predictions are consistent with empirical observations that males of some species engage in infrequent heterospecific mating. Thus, our models predict that reproductive interference can persist even after RCD occurred.

Host specialization by reproductive interference between closely related herbivorous insects

Host specialization among closely related herbivores is common and is therefore a major theme in ecology. Despite its ecological importance, no general explanatory framework for host specialization is currently available. We introduce a new model of the evolution of host specialization in herbivorous insects. We use a two-dimensional lattice comprising 32xa0×xa032 cells. Moreover, the model incorporates reproductive interference, defined as any negative outcomes resulting from interspecific mating, between two species of herbivores feeding on two different host plants as well as ordinary resource competition and differences in host suitability. Our simulation showed that reproductive interference, together with other factors such as host-related performance and resource competition, can actually drive the evolution of host specialization and thereby host partitioning between herbivorous insects during secondary contact. Host specialization arises particularly when levels of both reproductive interference and resource competition are intermediate. In contrast, host specialization seldom occurs if reproductive interference is absent. Thus, reproductive interference is as key to specialization as is spatial partitioning, which was not regarded as a realistic outcome when only resource competition was considered.

Pollen–pistil interactions in reproductive interference: comparisons of heterospecific pollen tube growth from alien species between two native Taraxacum species

Summary nReproductive interference (RI), any negative interspecific interaction during the reproductive process, has been gaining increasing attention due to its potential explanatory power for the mutually exclusive distribution of closely related species. RI in plants may occur during any of three stages: pollen transfer, pollen–pistil interactions or hybridization. Pollen–pistil interactions may be especially important as most studies of RI have suggested the involvement at this stage. Details of these interactions are required to fully explore RI and are especially relevant in considering the impact of RI in the field. nWe present a plausible explanation of how RI functions in the pollen–pistil interaction stage using two Japanese native dandelions, Taraxacum japonicum and Taraxacum longeappendiculatum, of which only the former is vulnerable to RI from an alien congener, Taraxacum officinale. We conducted a series of hand pollinations in these native dandelions and compared pollen tube behaviour to examine differences associated with vulnerability and imperviousness to RI from the alien. nThe two native dandelions differed in terms of the absence/presence of pollen tube elongation after heterospecific pollination (pollination with only T.xa0officinale): pollen tubes grew through the ovaries of the vulnerable T.xa0japonicum, but not through those of the impervious T.xa0longeappendiculatum. In vitro hand pollination verified that the alien pollen tubes could extend into the ovaries of T.xa0japonicum. nOur results show that RI from the alien dandelion consumed ovules by heterospecific pollen deposition. The pistils of the impervious native species could prevent growth of the alien pollen tubes, thereby sparing the ovules for fertilization by conspecific pollen. The pistils of the vulnerable species lacked interspecific incompatibility against the alien, and thus, the alien pollen tube entered the ovary, eliminating an opportunity for conspecific pollen fertilization. This consumption of ovules by heterospecific pollen tubes would cause a seed set failure, leading to reduced abundance and a further exertion of RI in the next generation, which explains displacement of the vulnerable species by the alien in the field.

Flowering phenology and mating success of the heterodichogamous tree Machilus thunbergii Sieb. et Zucc (Lauraceae)

Heterodichogamy is defined as the presence of two flower morphs that exhibit male and female functions at different times among individuals within a population, and is regarded as an adaptation to promote outbreeding through enhanced intermorph pollination. In highly fragmented populations in which the morph frequency is biased, heterodichogamy may hamper population growth by reducing seed sets of the more numerous morph, and enhancing seed sets of the less numerous morph. In such situations, we hypothesize that individual plants experience greater seed sets if the opposite sexual morphs are nearby, and that individuals of a less numerous sexual morph have greater seed sets. After confirming heterodichogamy by observing flowering behavior and phenology, we tested these two hypotheses in a highly fragmented population of Machilus thunbergii, a putative heterodichogamous evergreen laurel tree. Our observations confirmed that M.u2009thunbergii is heterodichogamous, consisting of two types of protogynous and bisexual flowers: a morning female (MF)–afternoon male morph and a morning male (MM)–afternoon female morph at the individual level. Sexual expression of the two morphs was highly synchronized and reciprocal. Investigation of seed-set rates revealed greater rates of both morphs if the opposite morph was nearby. The less numerous sexual morph (MF) showed a greater seed-set rate than the more numerous sexual morph (MM).

Effects of environmental factors on the species composition of aquatic insects in irrigation ponds

Although irrigation ponds contribute to the conservation of aquatic biodiversity, they have experienced declines in recent years. We therefore examined the relationships between various environmental factors and the community composition of aquatic insects, specifically insect predators, in irrigation ponds to gain knowledge that would aid in the conservation and restoration of biodiversity. We selected Odonata, Hemiptera and Coleoptera as target taxonomic groups and conducted censuses of these groups in 21 ponds in Shiga, central Japan. In total, we collected 30 and 10 species (or species groups) of Odonata and Hemiptera, respectively, and 17 species of Coleoptera. A partial canonical correspondence analysis revealed that the following four environmental factors significantly affected the species composition of aquatic insect communities: the number of emergent plant species, percent concrete revetment, presence of litter and peripheral length. Among these variables, the number of emergent plant species was the most potent factor, perhaps because emergent plants serve as sites for oviposition and emergence, and provide refugia for aquatic insects (odonate nymphs in particular). In contrast, some species specifically inhabited sites poor in emergent plants. This study shows that reductions in concrete revetments are necessary for the conservation of biodiversity. This would lead to increases in the number of aquatic plant species, which provide habitats and oviposition sites for many aquatic insects. Furthermore, to enrich the local biodiversity of aquatic insects, groups of irrigation ponds with different environments are needed.