Wataru Kojima

Physiological adaptation of the Asian corn borer Ostrinia furnacalis to chemical defenses of its host plant, maize.

A number of gramineous plants such as maize contain cyclic hydroxamic acids (cHx) that are toxic to many herbivores. Among the Ostrinia species found in Japan, the Asian corn borer Ostrinia furnacalis is the only one that utilizes maize, a gramineous plant. We used O. furnacalis and two congeners, Ostrinia scapulalis and Ostrinia latipennis, to obtain insights into the physiological adaptation of O. furnacalis to cHx. When an artificial diet containing a low concentration (0.3mg/g diet) of cHx was fed to the larvae of O. furnacalis and O. scapulalis, larval growth and survival were significantly less affected in O. furnacalis than O. scapulalis. An artificial diet containing a high level (0.7mg/g diet) of cHx was found to severely retard the growth of both species, albeit to different degrees. In an assay in vitro, homogenate of the digestive tract of O. furnacalis larvae degraded cHx more rapidly than that of O. scapulalis or O. latipennis. The degradation was found to be enzymatic and dependent on a cofactor, UDP-glucose, suggesting that UDP-glucosyltransferase or other UDP-glucose-dependent enzymes were involved. This enzymatic adaptation probably has enabled O. furnacalis to utilize plants containing cHx.

Low Level of Extra-Pair Paternity in a Population of the Barn Swallow Hirundo Rustica Gutturalis

Abstract On average, male birds other than social father sire more than 10% of all offspring. Levels of extra-pair paternity below 5% of offspring are rarely found and are now considered worthy of explanation in monogamous birds. We recorded the lowest levels of paternity loss ever reported in a population of Barn Swallows Hirundo rustica. The levels of extra-pair paternity were below 5% of offspring (7/243 in 2005 and 1/53 in 2006). We discuss our results in relation to the density-dependence of extra-pair paternity.

Pupal vibratory signals of a group-living beetle that deter larvae: Are they mimics of predator cues?

Pupae of some insects produce sounds or vibrations, but the function of the sounds/vibrations has not been clarified in most cases. Recently, we found vibratory communication between pupae and larvae of a group-living beetle Trypoxylus dichotoma, which live in humus soil. The vibratory signals produced by pupae were shown to deter approaching larvae, thereby protecting themselves. In the present study, we tested our hypothesis that pupal signals are mimics of vibratory noises associated with foraging of moles, the most common predators of T. dichotoma. Mole vibrations played back in laboratory experiments deterred larval approaches in the same way as pupal signals. These findings suggest that to deter conspecific larvae, pupae of T. dichotoma may have exploited a preexisting response of larvae to predator vibrations by emitting deceptive signals.

Attraction to Carbon Dioxide from Feeding Resources and Conspecific Neighbours in Larvae of the Rhinoceros Beetle Trypoxylus dichotomus.

Saprophagous (feeding on decaying matter) insects often use carbon dioxide (CO2) as a cue for finding food. Humus-feeding larvae of the giant rhinoceros beetle Trypoxylus dichotomus exhibit a clumped distribution in natural microhabitats, but the mechanisms driving the distribution were unknown. Herein, I examined whether larvae use CO2 as a cue for fermented humus and aggregate in the vicinity of the food. I found that (i) larvae of T. dichotomus are strongly attracted to CO2, (ii) larvae orient toward highly fermented humus when given a choice between highly and poorly fermented humus, (iii) the highly fermented humus emits more CO2 than the poorly fermented humus, and (iv) larvae grow larger when fed highly fermented humus rather than poorly fermented humus. The clumped distribution of larvae is probably formed along the concentration gradient of CO2 induced by heterogeneity of fermented organic materials in soil. My laboratory experiments also revealed that larvae are chemically attracted to each other. Moreover, CO2 concentrations in soil were increased by the larval respiration, and small amounts of CO2 (much less than emitted during respiration by a single larva) were sufficient for larval attraction. These results suggest that not only response to fermented food resources, but also respiratory CO2 from conspecifics may lead to aggregation. Enhanced densities resulted in reduced weight gain under experimental conditions. However, exploiting a high-value resource at enhanced densities still led to greater body weight compared to individually exploiting a low-value resource. This demonstrates the adaptive value of the response to CO2 sources in this species.

Variation in body size in the giant rhinoceros beetle Trypoxylus dichotomus is mediated by maternal effects on egg size

1. The egg size of insects can vary depending on maternal body size or resource status, and it may influence offspring body size by determining initial resource level.

Rhinoceros beetles suffer male-biased predation by mammalian and avian predators.

Male sexually-selected traits often impose an increased risk of predation on their bearers, causing male-biased predation. We investigated whether males of the sap-feeding Japanese rhinoceros beetle Trypoxylus dichotomus were more susceptible to predation than females by comparing the morphology of beetles caught in bait traps with the remains of beetles found on the ground. The males of this species are larger than the females and have a horn on the head. We found that predation pressure was greater for males than for females, and that larger individuals of both sexes were more vulnerable to predation. We identified two predators, the raccoon dog Nyctereutes procyonoides and jungle crow Corvus macrorhynchos, by monitoring sap-site trees with infrared video cameras. Raccoon dogs visited sap-site trees at night, while crows came after daybreak. The highest frequency of visits by both predators was observed in the first half of August, which matches the peak season of T. dichotomus. Raccoon dogs often left bite marks on the remains of prey, whereas crows did not. Bite marks were found on most of the remains collected at two distant localities, which suggested that predation by raccoon dogs is common. Size- and sex-dependent differences in the conspicuousness and active period of T. dichotomus probably explain these biased predation patterns. Our results suggest that having a large horn/body is costly in terms of the increased risk of predation. Predation cost may act as a stabilizing selection pressure against the further exaggeration of male sexual traits.

Deceptive vibratory communication: pupae of a beetle exploit the freeze response of larvae to protect themselves

It is argued that animal signals may have evolved so as to manipulate the response of receivers in a way that increases the fitness of the signallers. In deceptive communication, receivers incur costs by responding to false signals. Recently, we reported that pupae of the soil-inhabiting Japanese rhinoceros beetle Trypoxylus dichotoma produce vibratory signals to deter burrowing larvae, thereby protecting themselves. In the present study, monitoring of vibrations associated with larval movement revealed that T. dichotoma larvae remained motionless for ca 10 min when pupal vibratory signals were played back transiently (freeze response). Furthermore, pupal signals of T. dichotoma elicited a freeze response in three other scarabaeid species, whose pupae do not produce vibratory signals. This indicates that the freeze response to certain types of vibration evolved before the divergence of these species and has been evolutionarily conserved, presumably because of the fitness advantage in avoiding predators. Pupae of T. dichotoma have probably exploited pre-existing anti-predator responses of conspecific larvae to protect themselves by emitting deceptive vibratory signals.

Chemically mediated group formation in soil-dwelling larvae and pupae of the beetle Trypoxylus dichotomus

Many insects form groups through interactions among individuals, and these are often mediated by chemical, acoustic, or visual cues and signals. In spite of the diversity of soil-dwelling insects, their aggregation behaviour has not been examined as extensively as that of aboveground species. We investigated the aggregation mechanisms of larvae of the Japanese rhinoceros beetle Trypoxylus dichotomus, which live in groups in humus soil. In two-choice laboratory tests, 2nd- and 3rd-instar larvae gathered at conspecific larvae irrespective of the kinship. The ablation of maxillae, which bear chemosensilla, abolished aggregation behaviour. Intact larvae also exhibited aggregation behaviour towards a larval homogenate. These results suggest that larval aggregation is mediated by chemical cues. We also demonstrated that the mature larvae of T. dichotomus built their pupal cells close to a mesh bag containing a conspecific pupal cell, which indicated that larvae utilize chemical cues emanating from these cells to select the pupation site. Thus, the larvae of T. dichotomus may use chemical cues from the conspecifics in two different contexts, i.e. larval aggregation and pupation site selection. Using conspecific cues, larvae may be able to choose suitable locations for foraging or building pupal cells. The results of the present study highlight the importance of chemical information in belowground ecology.

Mechanism of synchronous metamorphosis: larvae of a rhinoceros beetle alter the timing of pupation depending on maturity of their neighbours

Synchronous development is widespread throughout the animal kingdom, but how synchrony is achieved remains largely unknown. I examined whether (1) a group-living rhinoceros beetle Trypoxylus dichotomus prepares for pupation (i.e. prepupates) synchronously in the field, (2) whether the synchrony occurs through social interactions, and (3) whether the synchrony incurs physiological costs. I found that larvae prepupate synchronously within natural humus sites. Laboratory experiments show that, when pairs of larvae are placed in the same cage, they prepupate on almost the same day, while two larvae chosen from different cages are expected to prepupate at 6-day intervals. I examined the mechanism of synchronous prepupation by inducing maturity asynchrony between two individuals. Less advanced larvae shortened the larval period in the presence of more advanced neighbours, whilst advanced individuals prolonged the larval periods in the presence of less advanced neighbours. However, variations in the prolongation or shortening of the larval period were dependant on the sites from which the larvae were collected, and in no site did both prolongation and shortening occur together. When the larval periods were prolonged or shortened, the body weight of the resulting pupae decreased. These data show that larvae of this species alter the timing of prepupation depending on the maturity of their neighbours. This developmental plasticity is likely to incur physiological costs due to pupation in suboptimal timing. The pupae and prepupae of this species may gain some benefits such as predator avoidance through the synchrony, which outweigh the cost in terms of reduced body weight.

Female Condition-dependent Allocation of Nuptial Gifts by Males in the Moth Ostrinia scapulalis (Lepidoptera: Crambidae)

ABSTRACT The males of many insect species transfer a spermatophore, i.e., a proteinaceous capsule containing sperm, to females during copulation, and this may also function as a nuptial gift. If production of the spermatophore is costly and variations in the quality of females are large, males may strategically allocate their investment based on the quality of the mate to maximize their own reproductive success. We examined the size and protein content of spermatophores transferred to females of different ages and body sizes, and also to water-deprived and water-replete females in the moth Ostrinia scapulalis (Walker). Males transferred a spermatophore of a smaller size or with less protein to older females, smaller females, and water-deprived females. These results indicated that O. scapulalis males manipulated their reproductive investment based on the conditions of the mate. We also demonstrated that older males varied their resource allocation to a greater extent in response to female conditions than younger males. Thus, resource allocation by the males of this species is modulated by both female conditions and the age of the males.