Kuniyasu Momose

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.

Irregular droughts trigger mass flowering in aseasonal tropical forests in asia

General flowering is a community-wide masting phenomenon, which is thus far documented only in aseasonal tropical forests in Asia. Although the canopy and emergent layers of forests in this region are dominated by species of a single family, Dipterocarpaceae, general flowering involves various plant groups. Studying proximate factors and estimating the flowering patterns of the past and future may aid our understanding of the ecological significance and evolutionary factors behind this phenomenon. Here we show that this phenomenon is most likely triggered by irregular droughts based on 10 years of observations. In the aseasonal forests of SE Asia, droughts tend to occur during transition periods from La Niña to El Niño, which results in an irregular 6-7-yr cycle involving a dry period with several droughts and a wet period without droughts. The magnitude of a flowering event also depends on the timing of droughts associated with the El Niño southern oscillation (ENSO) cycle, with the largest events occurring after an interval of several years with no flowering. Because most plant species can only reproduce successfully during large flowering events, changes in the ENSO cycle resulting from global warming, may have serious ramifications for forest regeneration in this region.

Beetle pollination of Shorea parvifolia (section Mutica, Dipterocarpaceae) in a general flowering period in Sarawak, Malaysia

Pollination ecology of an emergent tree species, Shorea (section Mutica) parvifolia (Dipterocarpaceae), was studied using the canopy observation system in a lowland dipterocarp forest in Sarawak, Malaysia, during a general flowering period in 1996. Although the species has been reported to be pollinated by thrips in Peninsular Malaysia, our observations of flower visitors and pollination experiments indicated that beetles (Chrysomelidae and Curculionidae, Coleoptera) contributed to pollination of S. parvifolia in Sarawak. Beetles accounted for 74% of the flower visitors collected by net-sweeping, and 30% of the beetles carried pollen, while thrips accounted for 16% of the visitors, and 12% of the thrips carried pollen. The apical parts of the petals and pollen served as a reward for the beetles. Thrips stayed inside the flower almost continuously after arrival, and movements among flowers were rare. Fruit set was significantly increased by introduction of beetles to bagged flowers, but not by introduction of thrips. Hand-pollination experiments and comparison of fruit set in untreated, bagged, and open flowers suggested that S. parvifolia was mainly outbreeding.

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.

Various population fluctuation patterns of light-attracted beetles in a tropical lowland dipterocarp forest in Sarawak

Abstract The population fluctuation pattern of light-attracted beetles was studied from August 1992 to September 1998 (for 73 months) using ultraviolet light-traps set at three vertical levels in a tropical lowland dipterocarp forest in Sarawak, Malaysia. During our study, a general flowering occurred from April to July in 1996, and flowering on a small scale in 1997 and 1998. We analyzed the data for eight scarabaeid and six meloid species, some of which were anthophilous species. Various fluctuation patterns were observed among the beetle species in aspects of both seasonality and correlation with the supraannual phenological pattern. Three large chafer species (Scarabaeidae, Melolonthini) showed a clear seasonal fluctuation pattern with a peak once from March to May every year, the peak monthly catch greatly fluctuating annually. Other scarabaeid beetles did not show such a clear seasonal population pattern and hardly fluctuated annually. Populations of an anthophilous scarabaeid species, Parastasia bimaculata, a specific pollinator of Homalomena propinqua (Araceae), hardly fluctuated, probably because of its response to the constant flowering of its floral hosts. Monthly catches of an anthophilous scarabaeid, Anomala sp., and meloid beetles showed clear supraannual patterns in response to the general flowering and were significantly correlated with the flowering intensity with or without a lag of a month. The fluctuation pattern of meloids suggests a supraannual population fluctuation pattern of their hosts, i.e., megachilid/anthophorid bees.

Resource use of insect seed predators during general flowering and seeding events in a Bornean dipterocarp rain forest.

Insect seed predators of 24 dipterocarp species (including the genera ot Dipterocarpus, Dryobalanops and Shorea) and five species belonging to the Moraceae, Myrtaceae, Celastraceae and Sapotaceae were investigated. In a tropical lowland dipterocarp forest in Sarawak, Malaysia, these trees produces seeds irregularly by intensely during general flowering and seeding events in 1996 and/or 1998. Dipterocarp seeds were preyed on by 51 insect species (11 families), which were roughly classified into three taxonomic groups: smaller moths (Trotricidae, Pyralidae, Crambidae, Immidae, Sesiidae, and Cosmopterigidae), scolytids (Scolydae) and weevils (Curdulionidae, Apionidae, Anthribidae, and Attelabidae). Although the host-specificity of invertebrate seed predators has been assumed to be high in tropical forests, it was found that the diet ranges of some insect predators were relatively wide and overlapped one another. Most seed predators that were collected in both study years changes their diets between general flowering and seeding events. The results of cluster analyses based on the number of adult of each predator species that emerged from 100 seeds of each tree species, suggested that the dominant species was not consistent, alternating between the two years.

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.

Population fluctuations of light-attracted chrysomelid beetles in relation to supra-annual environmental changes in a Bornean rainforest.

In Southeast Asian tropical rainforests, two events, severe droughts associated with the El Niño-Southern Oscillation and general flowering, a type of community-wide mass flowering, occur at irregular, supra-annual intervals. The relationship between these two supra-annual events and patterns of insect population fluctuations has yet to be clearly elucidated. Leaf beetles (Chrysomelidae) are major herbivores and flower-visitors of canopy trees, affecting their growth and reproduction and, in turn, affected by tree phenology; but their population fluctuations in the Southeast Asian tropics have not been extensively investigated. We examined population fluctuation patterns of the 34 most dominant chrysomelid species in relation to the two supra-annual events by conducting monthly light-trapping over seven years in a lowland dipterocarp forest in Borneo. Our results showed large community variation in population fluctuation patterns and a supra-annual (between-year) variation in abundance for most of the dominant chrysomelids that was significantly larger than the annual (within-year) variation. Specifically, in response to a severe drought in 1998, chrysomelid species exhibited different population responses. These results show that population fluctuations of individual species, rather than the entire assemblage, must be analyzed to determine the effects of changes in environmental conditions on the structure of insect assemblages in the tropics, especially in regions where supra-annual environmental changes are relatively more important than seasonal changes.

Plant reproductive intervals and pollinators in the aseasonal tropics: a new model

What factors determine reproductive intervals and modes of pollination in plants of the aseasonal tropics? To answer this general question, we present a new explanation for some community patterns of plant reproductive intervals and pollinators observed in a lowland dipterocarp forest in Sarawak, Malaysia, using a mathematical model featuring different display effects for different types of pollinators. Predictions from the model matched with the following observed patterns: (i) flowering intervals were different among forest strata (forest floor < understorey < canopy < subcanopy and emergent), and not in the exact order of stratum height; (ii) among generalist pollinators, the proportion of social foragers was maximum in intermediate forest stratum; and (iii) plants pollinated by specialist pollinators were found on the forest floor and in gaps.

Beetle Pollination in Tropical Rain Forests

Lists provided by Irvine and Armstrong (1990) and the Canopy Biology Program in Sarawak, or, CBPS (Momose et al. 1998c) show that 42 families of Coleoptera include pollinating beetles (see Table 9.1). Generalist-pollinated plants are almost always visited by beetles that feed on nectar or pollen. However, a close association with beetles as pollinators is revealed in mechanisms for excluding other kinds of flower visitors. Furthermore, the morphology of flowers and floral rewards are the relevant and conspicuous mechanisms.