Hiroshi Makihara

Biological invasion into the nested assemblage of tree–beetle associations on the oceanic Ogasawara Islands

Invasion by alien organisms is a common worldwide phenomenon, and many alien species invade native communities. Invasion by alien species is especially likely to occur on oceanic islands. To determine how alien species become integrated into island plant–insect associations, we analyzed the structure of tree–beetle associations using host plant records for larval feeding by wood-feeding beetles (Coleoptera: Cerambycidae) on the oceanic Ogasawara Islands in the northwestern Pacific Ocean. The host plant records comprised 109 associations among 28 tree (including 8 alien) and 26 cerambycid (including 5 alien) species. Of these associations, 41.3% involved at least one alien species. Most native cerambycid species feed on host trees that have recently died. Alien trees were used by as many native cerambycid species (but by significantly more alien cerambycid species) as were native trees. Native cerambycid species used as many alien tree species (but significantly more native tree species) as did alien cerambycids. Thus, we observed many types of interactions among native and alien species. A network analysis revealed a significant nested structure in tree–cerambycid associations regardless of whether alien species were excluded from the analysis. The original nested associations on the Ogasawara Islands may thus have accepted alien species.

Beetle responses to artificial gaps in an oceanic island forest: implications for invasive tree management to conserve endemic species diversity

Natural forests are often replaced by invasive alien trees on isolated oceanic islands. Adequate eradication of invasive trees should be conducted with the goal of biodiversity conservation, because islands support many endemic organisms that depend on native forests. An invasive alien tree, Bischofia javanica Blume (Euphorbiaceae), has invaded and replaced natural forests on the oceanic Ogasawara (Bonin) Islands, Japan, in the northwestern Pacific Ocean. To determine how the removal of B. javanica trees affects insect diversity, we examined flying beetles captured using Malaise traps in B. javanica forests on Hahajima. The abundance, species density, and species composition of wood-boring beetles (Coleoptera: Cerambycidae, Elateridae, Mordellidae, and Scolytidae) were compared between closed-canopy sites and gaps created by girdling B. javanica trees in alien forests during two seasons (June–July and October–November 2005). Of the collected beetles, 75.8, 87.5, 90.0, and 0.0% of cerambycid, elaterid, mordellid, and scolytid beetle species, respectively, were endemic to the Ogasawara Islands. More cerambycid, elaterid, and mordellid individuals were captured in June–July than in October–November; the number of scolytid individuals did not differ between seasons. More cerambycid, elaterid, and scolytid individuals were captured in artificial gaps than on the closed-canopy forest floor. Although fewer mordellid individuals were captured in gaps, more endemic mordellids were captured in gaps. More cerambycid and scolytid species were captured in artificial gaps than in closed-canopy areas. The positive responses of beetles to artificial gaps suggest that the removal of B. javanica increases beetle diversity and the abundance of endemic beetles.

Differences in endemic insect assemblages among vegetation types on a small island of the oceanic Ogasawara Islands

Natural vegetation is often replaced by invasive alien plants on isolated oceanic islands. To determine how invasive alien plants affect insect diversity, we compared flying insects captured using Malaise traps among different vegetation types on a small island (Nishijima; 0.49 km2) in the oceanic Ogasawara (Bonin) Islands in the north‐western Pacific. The numbers of individuals and species, and the species composition of pollinators (bees), predators (wasps) and wood borers (cerambycid, mordellid and elaterid beetles) were compared among three vegetation types: Casuarina equisetifolia (an invasive alien tree) forest, natural forest and natural grassland (forest edge), during two seasons (June and October–November 2005). In traps, 80.0, 66.7, 87.5, 85.7 and 100.0% of bee, wasp, cerambycid, mordellid and elaterid beetle species, respectively, were endemic to the Ogasawara Islands. Grassland had the highest wasp and bee species richness, whereas natural forest had the highest species richness of wood‐boring beetles. The C. equisetifolia forest had the poorest species richness for most insect groups (except mordellid beetles). More individuals of most insect groups (except bees) were captured in June than in October–November. More individual bees and wasps were captured in grassland than in forests, whereas more individual mordellid and elaterid beetles were captured in forests than in grassland. The number of cerambycid individuals did not differ among vegetation types. Redundancy analysis suggested that most insect species preferred natural forest or grassland to alien forest. Therefore, further invasion of natural grassland and forest by the alien tree C. equisetifolia may negatively affect the endemic insect fauna of Nishijima.

Effects of fires on butterfly assemblages in lowland dipterocarp forest in East Kalimantan

The post‐fire butterfly fauna in lowland dipterocarp forest of the Bukit Soeharto Education Forest (BSEF), East Kalimantan, Indonesia, was assessed during the period November 1998–April 2000 by means of consecutive Malaise trap samples, with supplementary field observations for March–April 1999. A total of 514 butterflies belonging to 61 species and representing six families were caught in the traps. Melanitis leda (Nymphalidae: Satyrinae), Charaxes bernardus (Nymphalidae: Charaxinae), and Danaus genutia (Nymphalidae: Danainae) were the species most frequently caught (60, 52 and 47 individuals, respectively), representing 31% of the total. These three species are generalists and “disturbance indicators” for tropical rainforest, being characteristic of disturbed or secondary forests, being distributed widely, and having larvae that feed on a wide range of host plants. In contrast, other species, such as Trogonoptera brookiana and Troides amphrysus, were recorded before the fires but were not recorded again afterwards. The pre‐ and post‐fire butterfly fauna of East Kalimantan were compared on the basis of butterfly specimens deposited in the Tropical Rain Forest Research Center that were collected in and around the Bukit Soeharto Education Forest before the fires (1988–1995). On the basis of the post‐fire survey, based on Malaise trap samples and field observations, only 43% of the butterfly species (not including Lycaenidae and Hesperiidae) were confirmed to have persisted. The data suggest that refugia that are not affected by fire are necessary for the conservation of specialist butterflies, as well as many other forms of wildlife.

Evaluation of secondary forests as alternative habitats to primary forests for flower-visiting insects

Although primary forests are important for biological conservation, the value of secondary forests for forest-dependent organisms needs to be evaluated when habitat restoration is required. We examined whether flower-visiting insects can use secondary forests as alternative habitats to primary forests. In particular, we compared assemblages of bees (Anthophila) and flower longhorn beetles (Lepturinae: Cerambycidae) in young secondary, mature secondary, and primary forests. Our results showed that more bee species were captured in primary and mature secondary forests than in young secondary forests, and flower longhorn beetle species were captured more frequently in primary forests than in mature and young secondary forests. Ordination showed that the communities in the three forest types were not statistically identical, which indicated that secondary forests cannot provide an absolute alternative habitat to primary forests for bees and flower longhorn beetles. However, the results also suggest that as secondary forests mature, more primary forest species would be able to use secondary forests as habitats. This implies that restoration from other land uses, such as monoculture plantations, to secondary forests could help to promote the faunal biodiversity of primary forests.

Sexual and Male Horn Dimorphism in Copris ochus (Coleoptera: Scarabaeidae)

Abstract Copris ochus (Coleoptera: Scarabaeidae), an endangered species, is the largest dung beetle in Japan. In C. ochus, males have a long head horn, while females lack this long horn (sexual dimorphism). Very large males of C. ochus have disproportionately longer head horns than small males, suggesting male horn dimorphism, although the dimorphism has not been investigated quantitatively. To clarify sexual and male horn dimorphism in C. ochus quantitatively, we examined the scaling relationship between body size (prothorax width) and head horn length in 94 females and 76 males. These beetles were captured during July 1978 from a natural population on Mt. Aso in southwestern Japan using a light trap. Although the horn length of the females and males scaled with prothorax width, the scaling relationship differed between the sexes, i.e., the relationship was linear in females and nonlinear in males. Statistical tests for dimorphism in male horn length showed a significant discontinuous relationship, thus indicating distinct sexual and male dimorphism in head horns. Long- and short-horned C. ochus males may have different reproductive behaviors, as described in other horned dung beetles.

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.

Chemical composition of the defensive secretion of the longhorned beetle, Chloridolum loochooanum.

Adults of the longhorned beetle, Chloridolum loochooanum Gressitt (Coleoptera: Cerambycidae) emit a white frothy secretion from their metasternal glands. This defensive substance contains cyclopentanoid monoterpenoids (iridodials), whose structures were elucidated by gas chromatography-mass spectrometry (GC-MS) and nuclear magnetic resonance (NMR) analyses that compared the naturally occurring structures with synthesized versions. Optically active citronellals, [(S)-, (R)-, and (S)/(R)- mixture], were used as starting materials for synthesizing the corresponding iridodials for the determination of the absolute configuration of the natural product. The retention time of (2S)-iridodial, derived from (S)-citronellal, corresponded to that of C. loochooanum iridodial by enantioselective GC analysis. Thus, we suggest that the absolute configuration of C. loochooanum iridodial is (1R,2S,5S)-iridodial.

Revisitation of sites surveyed 19 years ago reveals impoverishment of longhorned beetles in natural and planted forests

As planted forests expand in area, they are beginning to dominate landscapes as a matrix and cause the fragmentation of remaining natural forests. To understand and predict the responses of biological assemblages to maturing planted landscapes, examining the effects of forest type (natural vs planted) and forest age on such assemblages is particularly important. Therefore, to document the effects of forest type and age on longhorned beetle assemblages, in 2008 we collected beetles in broad‐leaved natural and cedar planted forests where beetles had also been collected in 1989. Beetle species composition differed greatly between the two forest types in 1989, whereas this difference was less pronounced in 2008. Species richness and total abundance were higher in natural forests than in planted forests in 1989. In 2008, species richness had decreased in both forest types, but the difference between the two forest types had been maintained. Total abundance was also markedly lower in 2008, and the difference between forest types was much smaller. Although larval host plants were not associated with the responses of species to year (forest age or maturation), beetle species whose larvae fed on either broad‐leaved or coniferous trees (or both) exhibited slight preferences for natural forests. These results suggest that longhorned beetle assemblages become impoverished in planted landscapes as the planted matrix matures. Changes in species composition with forest maturation may be difficult to predict based on larval host plants. However, consideration of larval host plants may enable the prediction of changes in species composition caused by the replacement of natural forests by planted forests.