Yoko Inui

Species-specific Leaf Volatile Compounds of Obligate Macaranga Myrmecophytes and Host-specific Aggressiveness of Symbiotic Crematogaster Ants

Macaranga myrmecophytes harbor species-specific Crematogaster ants that defend host trees from herbivores. We examined ant aggressive behaviors when artificially damaged leaf pieces from another tree were offered to four sympatric species of obligate Macaranga myrmecophytes. The ants showed aggressive behavior in response to leaf pieces regardless of the leaf species; however, aggressiveness was higher when conspecific leaf pieces were offered than when nonhost species were offered. Thus, ants can recognize leaf damage and distinguish among damaged leaf species. Chemical analyses of volatile compounds emitted from damaged leaves that may induce ant defense showed that the composition of the minor compounds differed among the four Macaranga species, although there were many compounds in common.

Feeding habits of Hymenoptera and Isoptera in a tropical rain forest as revealed by nitrogen and carbon isotope ratios

Despite the recognition of the functional role of Hymenoptera (ants, bees and wasps) and Isoptera (termites) in tropical ecosystems, their detailed feeding habits are not well known. To examine the feeding habits of these groups, we measured nitrogen (N) and carbon (C) stable isotope ratios (δ15N and δ13C) of hymenopterans (12 families, ≥16 genera and ≥32 species) and isopterans (one family and 10 species) collected in a tropical rain forest, Sarawak, Malaysia. We compared the isotopic signatures of these insects to those previously reported for other consumers collected in the same forest. The δ15N and δ13C values of these insects overlapped with those of the other consumers, indicating that they have access to diverse C and N sources in the forest. The δ15N values of ants and termites indicated that their feeding habits range along a continuum from herbivory (i.e. dependent on honeydew and nectar) to predation and from wood-feeders to soil-feeders, respectively. In addition, the δ15N values of wasps varied greatly from −0.1‰ (Braconidae sp.) to 8.6‰ (Bembix sp.), suggesting that their feeding habits also range from omnivory to predation. The ant species Camponotus gigas had δ13C values similar to those of invertebrate detritivores and omnivores rather than to those of invertebrate herbivores, although the diet of this species consists mostly of honeydew. This discrepancy suggests that the ant uses carbohydrates as an energy source, the isotopic signatures of which are not well retained in the body tissues. Values of both δ15N and δ13C of the predatory army ant Leptogenys diminuta and the soil-feeding termite Dicuspiditermes nemorosus did not differ significantly, indicating that both trophic level and the humification of feeding substrates can increase the isotopic signatures of terrestrial consumers.

Within‐nest abundance of a tropical cockroach Pseudoanaplectinia yumotoi associated with Crematogaster ants inhabiting epiphytic fern domatia in a Bornean dipterocarp forest

Many epiphytic ferns in tropical forest canopies have domatia that provide habitat for arboreal arthropods such as ants. Two species of fern that were predominantly occupied by Crematogaster difformis ants were collected from the canopy of a forest in Borneo. In the fern domatia, two cockroach species, Pseudoanaplectinia yumotoi and Blatta sp., were found to live with C. difformis. Pseudoanaplectinia yumotoi were found in large numbers and accounted for approximately 20% of the total arthropod population, while Blatta sp. comprised only 1% of the arthropods living in the domatia. Behavioural experiments were conducted to evaluate the response of C. difformis workers to newly introduced cockroaches of each species and to allospecific ant workers. Crematogaster difformis workers were highly aggressive and quickly attacked allospecific workers. Both cockroach species could fend off ant attacks; however, dead Blatta sp. were often attacked by the ants while dead P. yumotoi were infrequently bothered.

Secret Calls from under the Eaves: Acoustic Behavior of the Japanese House Gecko, Gekko japonicus

Several species of nocturnal gecko are known for their acoustic behavior in social contexts. The nocturnal house gecko, Gekko japonicus, was believed to be mute, except when threatened. We conducted behavioral encounter experiments involving same-sex and heterosexual pairs of G. japonicus. We also conducted two control experiments, chemical and blank, to confirm that acoustic behaviors are induced in the presence of conspecifics. Characteristic calls emitted by this gecko were recorded, providing the first evidence that G. japonicus uses acoustic signals for social communication. Geckos tended to call more frequently after the first physical contact with an opponent than before contact. In both control experiments, no gecko emitted calls. Whereas most characteristics of calls were similar to those reported for other gekkonid species, call intensity in this experiment was low. Males emitted calls toward both males and females, whereas females emitted calls almost exclusively toward other females. Several call variables differed significantly between intra- and intersexual calls. Our study demonstrated that G. japonicus uses acoustic signals in social contexts over short distances.

Various chemical strategies to deceive ants in three Arhopala species (lepidoptera: Lycaenidae) exploiting Macaranga myrmecophytes.

Macaranga myrmecophytes (ant-plants) are generally well protected from herbivore attacks by their symbiotic ants (plant-ants). However, larvae of Arhopala (Lepidoptera: Lycaenidae) species survive and develop on specific Macaranga ant-plant species without being attacked by the plant-ants of their host species. We hypothesized that Arhopala larvae chemically mimic or camouflage themselves with the ants on their host plant so that the larvae are accepted by the plant-ant species of their host. Chemical analyses of cuticular hydrocarbons showed that chemical congruency varied among Arhopala species; A. dajagaka matched well the host plant-ants, A. amphimuta did not match, and unexpectedly, A. zylda lacked hydrocarbons. Behaviorally, the larvae and dummies coated with cuticular chemicals of A. dajagaka were well attended by the plant-ants, especially by those of the host. A. amphimuta was often attacked by all plant-ants except for the host plant-ants toward the larvae, and those of A. zylda were ignored by all plant-ants. Our results suggested that conspicuous variations exist in the chemical strategies used by the myrmecophilous butterflies that allow them to avoid ant attack and be accepted by the plant-ant colonies.

Congruence of Microsatellite and Mitochondrial DNA Variation in Acrobat Ants (Crematogaster Subgenus Decacrema, Formicidae: Myrmicinae) Inhabiting Macaranga (Euphorbiaceae) Myrmecophytes

A previously reported mitochondrial DNA (mtDNA) phylogeny of Crematogaster (subgenus Decacrema) ants inhabiting Macaranga myrmecophytes indicated that the partners diversified synchronously and their specific association has been maintained for 20 million years. However, the mtDNA clades did not exactly match morphological species, probably owing to introgressive hybridization among younger species. In this study, we determined the congruence between nuclear simple sequence repeat (SSR, also called microsatellite) genotyping and mtDNA phylogeny to confirm the suitability of the mtDNA phylogeny for inferring the evolutionary history of Decacrema ants. Analyses of ant samples from Lambir Hills National park, northeastern Borneo, showed overall congruence between the SSR and mtDNA groupings, indicating that mtDNA markers are useful for delimiting species, at least at the local level. We also found overall high host-plant specificity of the SSR genotypes of Decacrema ants, consistent with the specificity based on the mtDNA phylogeny. Further, we detected cryptic genetic assemblages exhibiting high specificity toward particular plant species within a single mtDNA clade. This finding, which may be evidence for rapid ecological and genetic differentiation following a host shift, is a new insight into the previously suggested long-term codiversification of Decacrema ants and Macaranga plants.

Potential host range of myrmecophilous Arhopala butterflies (Lepidoptera: Lycaenidae) feeding on Macaranga myrmecophytes

“Plant-ants”, i.e. those symbiotic with myrmecophyte plants, defend their hosts against herbivores. Plant-ants are expected to affect the host-plant ranges of herbivores that feed on myrmecophytes. This study aimed to experimentally determine whether anti-herbivore defences by plant-ants restrict the larval host-plant ranges of four Arhopala (Lycaenidae) butterflies that feed on Macaranga (Euphorbiaceae) trees, some of which are myrmecophytes. We fed Arhopala larvae with the leaves of five Macaranga species under ant-excluded conditions to examine their potential host-plant ranges. Under ant-excluded conditions, three Arhopala species survived to the pupal stage when fed species not used in the field as well as their normal host species. Our data suggested that the aggressive behaviours of plant-ants towards leaf-feeding insects restrict the potential host-plant ranges of some Macaranga-feeding Arhopala butterflies.

Myrmecoxeny in Arhopala zylda (Lepidoptera, Lycaenidae) Larvae Feeding on Macaranga Myrmecophytes

ABSTRACT Some species in the tree genus Macaranga (Euphorbiaceae) in the Southeast Asian tropics are myrmecophytic; they have highly species-specific mutualisms with symbiotic ants (plant-ants), which defend them from herbivores. However, larvae of some Arhopala (Lycaenidae, Lycaeninae) species can elude the ants. Here we demonstrated that Arhopala zylda larvae showed myrmecoxeny on their myrmecophytic Macaranga host plants; they had no stable association with the plant-ants. Despite the presence of many plant-ants, A. zylda larvae were rarely attended or attacked by ants on their host plants. The plant-ants of three other myrmecophytic Macaranga species (non-hosts to A. zylda) also paid little attention to experimentally introduced A. zylda larvae. The myrmecoxeny seen in A. zylda is notable among lycaenid larvae that feed on myrmecophytes, because almost all are obligate intimate myrmecophiles.

Production of food bodies on the reproductive organs of myrmecophytic Macaranga species (Euphorbiaceae): effects on interactions with herbivores and pollinators

In protective ant–plant mutualisms, plants offer ants food (such as extrafloral nectar and/or food bodies) and ants protect plants from herbivores. However, ants often negatively affect plant reproduction by deterring pollinators. The aggressive protection that mutualistic ants provide to some myrmecophytes may enhance this negative effect in comparison to plant species that are facultatively protected by ants. Because little is known about the processes by which myrmecophytes are pollinated in the presence of ant guards, we examined ant interactions with herbivores and pollinators on plant reproductive organs. We examined eight myrmecophytic and three nonmyrmecophytic Macaranga species in Borneo. Most of the species studied are pollinated by thrips breeding in the inflorescences. Seven of eight myrmecophytic species produced food bodies on young inflorescences and/or immature fruits. Food body production was associated with increased ant abundance on inflorescences of the three species observed. The exclusion of ants from inflorescences of one species without food rewards resulted in increased herbivory damage. In contrast, ant exclusion had no effect on the number of pollinator thrips. The absence of thrips pollinator deterrence by ants may be due to the presence of protective bracteoles that limit ants, but not pollinators, from accessing flowers. This unique mechanism may account for simultaneous thrips pollination and ant defense of inflorescences.

Herbivore-constructed leaf shelters on Salix eriocarpa shoots affect arthropod communities

We observed the abundance of leaf shelters, aphids, other herbivores, and predators on willow trees, Salix eriocarpa, from May to October 2003. There was a positive correlation between the growth rate of aphids and the number of ants per shoot, suggesting ant attendance to aphids. Although the mean abundance of leaf shelters per shoot was rather low (1.7–2.2) throughout the observation period, aphids preferred to use shoots with leaf shelters compared with those without leaf shelters. The abundance of ants was positively influenced by the presence of leaf shelters and aphids from May to August. The abundance of other herbivores was positively influenced by leaf shelters, but negatively influenced by aphid presence from May to August. Furthermore, leaf shelters had a positive effect on the abundance of predators from July to October. These data suggest that a relatively low abundance of naturally occurring leaf shelters per shoot influenced the arthropod communities on S. eriocarpa, and the effect of those leaf shelters on each type of arthropod varied according to the season.