Naoki Kachi

The mechanism of general flowering in Dipterocarpaceae in the Malay Peninsula

The mechanism of general flowering in Dipterocarpaceae in the Malay Peninsula is revealed through field survey and meteorological data analyses. The regions of general flowering coincide with those which experienced a low night-time temperature (LNT) c. 2 mo before flowering. This supports the hypo- thesis that low air temperature induces the development of floral buds of diptero- carps. LNT was found to be caused by radiative cooling during dry spells in winter when the northern subtropical ridge (STR) occasionally migrates southwards with a dry air mass into the equatorial region. LNT events usually occur in La Nifia episodes, not in El Nifio episodes as believed previously. This is because the south- ward migration of the STR is associated with the intensification of local meridional Hadley Circulation in the western Pacific, which is strengthened in a La Nifia episode. Results suggest that El Nifio-like climate change in increased atmospheric carbon dioxide concentrations may be critical for the tropical rain forest biome in south-east Asia.

Temporal and spatial patterns of mass flowerings on the Malay Peninsula

We propose a hypothesis to explain the temporal and spatial patterns of mass flowerings in dipterocarp tree species on the Malay Peninsula. The literature on these mass flowerings reveals that during 1980-2002 at least 11 flowerings occurred at irregular intervals of 1-6 yr in a lowland rain forest. Five of them were typical mass flowerings-a high density of flowering trees and the characteristic sequential flowering of Shorea species. The 11 flowerings were classified into two flowering times: spring and autumn. There is evidence that low temperature and drought triggered the flowerings. Therefore, the seasonality of mass flowerings is characterized by the annual patterns of rainfall and low temperature. In addition, changes in El Niño-Southern Oscillation (ENSO) may play important roles in determining the supra-annual occurrence of mass flowerings. Flowering surveys on the Malay Peninsula implied that regions with spring or autumn mass flowerings corresponded geographically to those regions that had one cool season (December-February) or two (December-February and June-August), respectively. This finding anticipates the seasonal pattern and geographical distribution of mass flowerings on the Malay Peninsula.

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.

Development of a Local Size Hierarchy Causes Regular Spacing of Trees in an Even-aged Abies Forest: Analyses Using Spatial Autocorrelation and the Mark Correlation Function

BACKGROUND AND AIMS During the development of an even-aged plant population, the spatial distribution of individuals often changes from a clumped pattern to a random or regular one. The development of local size hierarchies in an Abies forest was analysed for a period of 47 years following a large disturbance in 1959. METHODS In 1980 all trees in an 8 x 8 m plot were mapped and their height growth after the disturbance was estimated. Their mortality and growth were then recorded at 1- to 4-year intervals between 1980 and 2006. Spatial distribution patterns of trees were analysed by the pair correlation function. Spatial correlations between tree heights were analysed with a spatial autocorrelation function and the mark correlation function. The mark correlation function was able to detect a local size hierarchy that could not be detected by the spatial autocorrelation function alone. KEY RESULTS The small-scale spatial distribution pattern of trees changed from clumped to slightly regular during the 47 years. Mortality occurred in a density-dependent manner, which resulted in regular spacing between trees after 1980. The spatial autocorrelation and mark correlation functions revealed the existence of tree patches consisting of large trees at the initial stage. Development of a local size hierarchy was detected within the first decade after the disturbance, although the spatial autocorrelation was not negative. Local size hierarchies that developed persisted until 2006, and the spatial autocorrelation became negative at later stages (after about 40 years). CONCLUSIONS This is the first study to detect local size hierarchies as a prelude to regular spacing using the mark correlation function. The results confirm that use of the mark correlation function together with the spatial autocorrelation function is an effective tool to analyse the development of a local size hierarchy of trees in a forest.

Optimal resource allocation to seeds and vegetative propagules under density-dependent regulation in Syneilesis palmata (Compositae)

Syneilesis palmata reproduces by both seeds and vegetative propagules (short rhizomes). The latter result in the production of new plants that are larger in size and hence have a higher survival probability and a higher growth rate than seeds. A previous study predicted that the optimal reproductive strategy, in terms of maximizing population growth rate (a fitness measure under no density regulations), was pure vegetative reproduction. However, high resource investment to vegetative propagules can cause local crowding resulting in reduced demographic performances of the plants, because the vegetative propagules of Syneilesis are produced close to one another. We examined, in this situation, the impact of allocating a certain proportion of reproductive resource to seeds with relatively greater capacity for dispersal. We simulated dynamics of hypothetical Syneilesis populations with various reproductive resource allocation balances (from pure seed to pure vegetative reproduction), using a density-dependent matrix model. In the model, it was assumed that plants from vegetative propagules experienced density-dependent reduction in their survival probabilities, but this was not the case for plants originating from seeds. Each allocation strategy was evaluated based on an equilibrium population density, a fitness measure under density-dependent regulations. The optimal reproductive strategy predicted was pure vegetative reproduction. Unrealistic conditions were required for seed reproduction to be favoured, such as the production of seeds one hundred times the normal number per unit resource investment. However, the conditions were fairly relaxed compared with those required in the model where no density effects were incorporated. This indicates that escape from local crowding is likely to be one of the roles of seed production in Syneilesis.

Differential salt tolerance of two Artemisia species growing in contrasting coastal habitats.

To investigate factors determining the differences in their salt tolerance, growth and germination, experiments were conducted on two plant species belonging to genus Artemisia: Artemisia fukudo Makino, a biennial salt marsh plant and Artemisia stelleriana Bess, a perennial coastal hind dune plant. Growth experiments revealed that salinity (100 and 300 m m NaCl) inhibited the relative growth rate (RGR) in A. stelleriana significantly but not in A. fukudo. These specific differences in salt tolerance were mainly attributed to differential responses of net assimilation rate (NAR). That is, the reduction in RGR in A. stelleriana was mainly due to the reduction in NAR, whereas no significant reduction in NAR was observed in A. fukudo. The reduction in RGR in A. stelleriana in the salt treatment was also attributable to a reduced leaf area ratio (LAR). Specific leaf area (SLA) in the two species decreased in the 300 m m treatment. The decrease in SLA in A. fukudo was, however, compensated for partly by an increase in leaf weight ratio (LWR). Germination experiments also showed that A. fukudo has a higher salt tolerance than does A. stelleriana. These results are consistent with the differences in the salinity conditions between the native habitats of the two species.

Effects of temporal heterogeneity of water supply on the growth of Perilla frutescens depend on plant density

BACKGROUND AND AIMS Plants respond to the spatial and temporal heterogeneity of a resource supply. However, their responses will depend on intraspecific competition for resource acquisition. Although plants are subject to various intensities of intraspecific competition, most studies of resource heterogeneity have been carried out under a single density so that the effects of intraspecific competition on plant responses to resource heterogeneity are largely unknown. METHODS A growth experiment was performed to investigate plant responses to the temporal heterogeneity of water supply and nutrient levels under multiple plant densities. The annual plant Perilla frutescens was grown using different combinations of frequency of water supply, nutrient level and density, while providing the same total amount of water under all conditions. The effects of the treatments on biomass, allocation to roots and intensity of competition were analysed after 48 d. KEY RESULTS Biomass and allocation to roots were larger under homogeneous than under heterogeneous water supply, and the effects of water heterogeneity were greater at high density than at low density. The effects of water heterogeneity were greater at high nutrient level than at low level for biomass, while the effects were greater at low nutrient level than high level for allocation to roots. Competition was severer under homogeneous than under heterogeneous water supply. CONCLUSIONS Competition for water probably makes plants more sensitive to the water heterogeneity. In addition, the intensity of intraspecific competition can be affected by the temporal patterns of water supply. Because both resource heterogeneity and intraspecific competition affect resource acquisition and growth of plants, their interactive effects should be evaluated more carefully under future studies.

Availability and Temporal Heterogeneity of Water Supply Affect the Vertical Distribution and Mortality of a Belowground Herbivore and Consequently Plant Growth

We examined how the volume and temporal heterogeneity of water supply changed the vertical distribution and mortality of a belowground herbivore, and consequently affected plant biomass. Plantago lanceolata (Plantaginaceae) seedlings were grown at one per pot under different combinations of water volume (large or small volume) and heterogeneity (homogeneous water conditions, watered every day; heterogeneous conditions, watered every 4 days) in the presence or absence of a larva of the belowground herbivorous insect, Anomala cuprea (Coleoptera: Scarabaeidae). The larva was confined in different vertical distributions to top feeding zone (top treatment), middle feeding zone (middle treatment), or bottom feeding zone (bottom treatment); alternatively no larva was introduced (control treatment) or larval movement was not confined (free treatment). Three-way interaction between water volume, heterogeneity, and the herbivore significantly affected plant biomass. With a large water volume, plant biomass was lower in free treatment than in control treatment regardless of heterogeneity. Plant biomass in free treatment was as low as in top treatment. With a small water volume and in free treatment, plant biomass was low (similar to that under top treatment) under homogeneous water conditions but high under heterogeneous ones (similar to that under middle or bottom treatment). Therefore, there was little effect of belowground herbivory on plant growth under heterogeneous water conditions. In other watering regimes, herbivores would be distributed in the shallow soil and reduced root biomass. Herbivore mortality was high with homogeneous application of a large volume or heterogeneous application of a small water volume. Under the large water volume, plant biomass was high in pots in which the herbivore had died. Thus, the combinations of water volume and heterogeneity affected plant growth via the change of a belowground herbivore.

Chemical defences of fruits and mast-fruiting of dipterocarps

Mast-fruiting is the intermittent and synchronous production of large fruits by a population of plants at long intervals (Herrera et al. 1998, Kelly 1994). Several hypotheses have been proposed concerning the adaptive advantages of mast-fruiting (Janzen 1971, 1974; Kelly 1994), and some field observations have provided evidence for these hypotheses (Norton & Kelly 1988, Shibata et al. 1998, Sork 1993). The predator-satiation hypothesis is one well-known explanation for reproductive synchrony in plants and animals (Janzen 1971, 1974; Kelly 1994). This hypothesis claims that mast fruiting at irregular intervals of several years is an effective means of satiating vertebrate fruit predators: low seed production can only support low densities of predators during the periods between mast-fruiting events, but more fruits are produced than predators can consume in masting years (Janzen 1971, Kelly 1994). Thus, it may be said that mast-fruiting is a defence strategy of plants against post-dispersal vertebrate fruit predators.

The ecological significance of clonal growth in the understory tree, Pawpaw ( Asimina triloba )

Abstract We evaluated three possible functions of clonal growth related to genet persistence in the root-suckering understory tree pawpaw, Asimina triloba (L.) Dunal: (1) risk spreading through multiplication of stems, (2) enhanced establishment and survival of new stems, and (3) horizontal expansion growth of patches. The number, diameter growth, and spatial distribution of annual stem recruits were examined over three years in a natural population of pawpaw. The rate of stem recruitment was consistently higher than stem mortality. We found no difference in stem turnover rate for patches of different size, indicating that stem production is more than high enough to avoid patch extinction. Although newly formed stems were considerably smaller than previously established stems, they grew and survived as well as established stems. We found no evidence for clonal growth contributing to extensive horizontal expansion of patches. Our results suggest that ensuring survivorship of new stems is the main ecological role of clonal growth in pawpaw.