Teruyoshi Nagamitsu

Pollination biology in a lowland dipterocarp forest in Sarawak, Malaysia. I. Characteristics of the plant-pollinator community in a lowland dipterocarp forest

Flowerings and flower visitors were observed continuously in alowland dipterocarp forest in Sarawak, Malaysia, for 53 mo in1992-1996. Flower visitors of 270 plant species were observed orcollected, and pollinators were assessed by observing body contact tostigmas and anthers. We recognized 12 categories of pollination systems.Among them, plants pollinated by social bees included the largest numberof species (32%) and were followed by beetle-pollinated species(20%). Pollination systems were significantly related with somefloral characters (flowering time of day, reward, and floral shape), butnot with floral color. Based on the relationships between pollinatorsand floral characters, we described pollination syndromes found in alowland dipterocarp forest. The dominance of social bees and beetlesamong pollinators is discussed in relation to the general floweringobserved in dipterocarp forests of West Malesia. In spite of high plantspecies diversity and consequent low population densities of lowlanddipterocarp forests, long-distance-specific pollinators were uncommoncompared with theNeotropics.

Seasonality and vertical structure of light-attracted insect communities in a dipterocarp forest in Sarawak

Nocturnal flying insects were collected monthly for 13 months using ultra violet light-traps set at various vertical levels in a weakly-seasonal, tropical lowland dipterocarp forest in Sarawak, Malaysia. Abundance, faunal composition, size distribution and guild structure of these samples were analyzed with respect to temperal and vertical distributions. The nocturnal flying insect community in the canopy level was highly dominated by fig wasps (84%) in individual number, and by scarabaeid beetles (28%) in weight. A principal component analysis on monthly catches detected non-random, seasonal trends of insect abundance. The first two principal trends were an alternation of wetter (September to January) and less wet seasons (February to August) and an alternation between the least wet (January to March) and the other seasons. Many insect groups were less abundant in the least wet season than the other seasons, whilst inverse patterns were found in Scarabaeidae and Tenebrionidae. Significantly positive and negative correlations between monthly catch and rainfall were detected only in ovule-feeders and in phloem-feeders, respectively. Delayed, significant negative correlations between monthly catch and 1–3 month preceding rainfall were more frequently detected in phytophages, phloem-feeders, seed-feeders, wood-borers and scavengers. The peak in abundance along vertical levels were found at the canopy level (35 m) for phloem-, ovule-, seed-, root-, fungal-feeders and nectar collectors, at an upper subcanopy level (25 m) for scavengers and aquatic predators, and at a middle subcanopy level (17 m) for ants. Catches at the emergent level (45 m) did not exceed those at the canopy level.

Aggressive foraging of social bees as a mechanism of floral resource partitioning in an Asian tropical rainforest

Abstract Interference competition by aggressive foraging often explains resource partitioning, but mechanisms contributing to partitioning have rarely been studied in Asian social bee guilds. Foraging of social bees at canopy flowers of Santiria laevigata (Burseraceae) and honey-water feeders was studied in a lowland mixed-dipterocarp forest in Sarawak, Malaysia. Four stingless bee species (Apidae, Meliponinae), Trigona canifrons, T.␣fimbriata, T. apicalis and T. melina, aggressively defended flower patches and feeders. At the flowers, T.␣canifrons excluded other bees only in the morning when nectar flow peaked. At the feeders, the aggression resulted in asymmetric interference competition, which produced a dominance hierarchy among seven social bee species. Interspecific partitioning of the feeders was detected in time and height but not quality. Only time of the first arrival after feeder presentation was negatively correlated with the dominance hierarchy: more aggressive species arrived at the feeders later than less aggressive species. This result suggests that a trade-off between searching ability and defensive ability at flower patches gives rise to resource partitioning in the social bee guild.

Cockroach pollination and breeding system of Uvaria elmeri (Annonaceae) in a lowland mixed-dipterocarp forest in Sarawak.

Tropical forest plants are known to be pollinated by a diverse array of animals. Here we report on the pollination of a woody climber species, Uvaria elmeri (Annonaceae), by cockroaches in a lowland mixed-dipterocarp forest in Sarawak, Malaysia. To the best of our knowledge, this is the first report of pollination by cockroaches. The cauliflorous flowers in the understory are protogynous and bloomed for 50 h. An odor similar to decayed wood or a mushroom was secreted by flowers and was stronger during the male stage. Pollinators were cockroaches (Blattellidae) and drosophilid flies (Drosophilidae). Cockroaches, the main pollinators, visited flowers during both female and male stages at night, feeding on stigmatic exudate and pollen. Drosophilids, the secondary pollinators. mainly visited female-stage flowers during daytime, fed on stigmatic exudate. and laid eggs on stigmas. Neither autogamy nor self-compatibility was observed. Fruit production appeared to be pollen-limited. The fruit set, which was 2% of flowers in natural condition, was significantly lower than the 30% fruit set obtained by artificial cross-pollination. We discuss the traits of cockroaches as pollinators and the breeding system of U. elmeri.

Development and polymorphism of simple sequence repeat DNA markers for Shorea curtisii and other Dipterocarpaceae species

Nine simple sequence repeat (SSR) markers were developed from Shorea curtisii using two different methods. One SSR locus was isolated by the commonly used method of screening by colony hybridization, and the other eight loci were isolated by a vectorette PCR method. Primer pairs were designed based on the sequences of all these SSR loci. Analysis of 40 individuals of S. curtisii from natural forest in Malaysia revealed that all SSR loci were polymorphic. Four SSR markers, Shc01, Shc04, Shc07 and Shc09, were highly polymorphic. We have also tested the applicability of these SSR printers to other species of Dipterocarpaceae using PCR amplification. Because the flanking region sequences of the S. curtisii SSRs were well conserved within this family, the SSR primers for S. curtisii can be applied to almost all species of Dipterocarpaceae.

Six-Year Population Fluctuation of the Giant Honey Bee Apis dorsata (Hymenoptera: Apidae) in a Tropical Lowland Dipterocarp Forest in Sarawak

Abstract The giant honey bee Apis dorsata F. inhabits lowland tropical rainforests in Southeast Asia, where a general, community-wide flowering occurs at intervals of 4 yr on average. The numerical response by the honey bee population to the drastic increase of flower resources during general flowering was investigated for 6 yr by monthly light-trapping and by nest counts in a lowland dipterocarp forest in Borneo. The numbers of A. dorsata workers obtained by light-trapping were highest during general flowering periods, whereas very few workers were trapped in other periods. The abundance of A. dorsata nests showed temporal correspondence with the abundance of trapped workers, and the nests disappeared in the nonflowering periods. These data suggest that the A. dorsata population increases rapidly in response to general flowering and that this is initiated by nonseasonal, long-distance migration. Drones of A. dorsata were present during the general flowering period, but there is no evidence that reproduction by A. dorsata occurs only in general flowering periods. Fluctuation in abundance by the honey bee A. koschevnikovi Enderlein was also observed by monthly light-trapping. The temporal trend of this species was similar to that of A. dorsata, but sightings persisted even in the nonflowering periods. Both honey bees responded numerically to floral resources, but long-distance migration in A. koschevnikovi was unlikely.

Preference in flower visits and partitioning in pollen diets of stingless bees in an Asian tropical rain forest

Floral resource partitioning among stingless bees (Trigona, Meliponini, Apidae) in a lowland rain forest in Sarawak, Malaysia, was investigated using tree towers and walkways in a 4-year study that included a general flowering period. We obtained 100 collections of insect visitors to flowers of varying floral location and shape representing 81 plant species. The tendency of 11 species of stingless bees to visit specific flowers with a particular floral location and shape was analyzed by logistic regression analysis. This analysis showed that the proportion of flower visitor collections containing Trigona fuscobalteata and T. melanocephala differed according to floral location. The former was frequently collected at canopy and gap flowers, whereas the latter was most often collected at understory flowers. The analysis also suggested that T. erythrogastra was more rarely collected at shallow flowers than at deep flowers. Analysis of the pollen diets of T. collina, T. fuscobalteata, T. melanocephala, and T. melina revealed that similarity of pollen sources differed among the six permutated pairs of the four species. The lowest mean rank of similarity found was between T. fuscobalteata and T. melanocephala. This result supports the hypothesis that preference in visiting flowers in different locations leads to pollen resource partitioning.

MICROSATELLITE ANALYSIS OF THE BREEDING SYSTEM AND SEED DISPERSAL IN SHOREA LEPROSULA (DIPTEROCARPACEAE)

To understand the breeding system and seed dispersal in Shorea leprosula (Dipterocarpaceae) in Peninsular Malaysia, a microsatellite analysis was conducted of embryos of immature and mature fruits fallen in litter traps under the crowns of five trees. Outcrossed and selfed progeny mothered by the trees and those dispersed from other trees were distinguished by genotypes of three polymorphic microsatellite loci. The mean outcrossing rate of mature fruit embryos in S. leprosula pollinated by thrips (0.91) was not lower than those previously reported from Shorea species pollinated by bees, even though thrips seem to be less efficient pollinators than bees. Although four of the five trees showed high and stable outcrossing rates during fruit maturation, the outcrossing rate increased in one tree with highly selfed embryos of immature fruits. These results suggest that inbreeding depression during fruit maturation as well as self‐incompatibility reduce the proportion of inbred embryos. The proportion of fruits dispersed from neighbor trees in fruits trapped under a tree crown had a mean value of 0.20 and was lowest under the tree with highly selfed embryos of immature fruits. This low fraction of dispersed fruits under this tree suggests long distances from this tree to reproductive neighbors, which may reduce cross pollination.

Differences in pollen sources of Apis cerana and Apis mellifera at a primary beech forest in central Japan

SUMMARYPlant taxa of pollen sources for Apis cerana japonica and A. mellifera ligustica at a primary beech forest in Ashiu (Kyoto, Japan) were compared. Most pollen sources were shared between the two honey bee species. Seasonal variation in the pollen utilization of the two bee species were similar from April to October. Pollen source overlap in interspecific pairs of colonies was smaller than that in intraspecific pairs at the start and end of the season. Pollen collection from seven plant taxa differed between A. cerana and A. mellifera; A. cerana preferred tall trees, while A. mellifera favoured short herbs. Dioecious and andromonoecious flowers with green petals were preferred by A. cerana. These findings suggest differences in both resource location and visual attractiveness of pollen sources between the two honey bee species, which were emphasized when air temperature was low.

Abundance, body size, and morphology of bumblebees in an area where an exotic species, Bombus terrestris, has colonized in Japan

An exotic bumblebee species, Bombus terrestris, has colonized in Japan and becomes dominant in some local communities. We examined the effects of land use and bumblebee abundance on the number and body size of bumblebees collected using window traps in a lowland area in the southern Ishikari district, Hokkaido. In 2004, we collected 922 bumblebees of six species using 70 traps at 17 sites. A statistical model fitted to the data demonstrated that dispersion from commercial B. terrestris colonies used in greenhouses positively affected the number of B. terrestris caught by each trap. This exotic species was abundant in sites where paddy fields were prevalent, but three native species, B. hypocrita, B. ardens, and B. diversus, were abundant in sites where farms and woodlands were widespread. The local abundance of B. terrestris was not associated negatively with the number and body size of native bumblebees. Thus, we did not find any competitive interactions between exotic and native bumblebees although habitat conditions seem to be common determinants of the bumblebee populations. A morphological analysis showed that B. terrestris had intermediate tongue length between B. hypocrita and B. ardens.