Dimorphic flowers modify the visitation order of pollinators from male to female flowers.

Sexual dimorphism is a pervasive form of variation within species. Understanding how and why sexual dimorphism evolves would contribute to elucidating the mechanisms underlying the diversification of traits. In flowering plants, pollinators are considered a driver of sexual dimorphism when they affect female and male plant fitness in distinct ways. Here, we found that flowers appear to manipulate the behavior of pollinators using sexually dimorphic traits in the dioecious tree Eurya japonica. In this plant, female flowers present a higher-quality reward for pollinators, whereas male flowers have a more conspicuous appearance. Plants benefit by inducing pollinators to carry pollen from male to female flowers, and their sexual dimorphism might thus facilitate pollen movement through pollinator behavior. In two-choice experiments, pollinators frequently moved from male to female flowers, whereas computer simulation suggested that sexually dimorphic traits would evolve if pollinators changed behavior depending on the traits of the flowers they had just visited. These results suggest that the floral traits affecting the visiting order of pollinators have evolved in plants. Using E. japonica, we theoretically show that the induction of sequential behavior in pollinators might be crucial to the evolution of sexual dimorphism in flowers, and our experiments support these findings.

The N-terminal and third transmembrane domain of PsCor413im1 are essential for targeting to chloroplast envelope membrane.

Cold-regulated (COR) genes, located downstream of the C-repeat binding factors (CBFs) in cold signaling pathways, play a central role in plant response to cold stress. In our previous studies, a Cor413 chloroplast envelope membrane protein, PsCor413im1, was identified from the cold-tolerant plant Phlox subulata. Its overexpression enhanced cold tolerance and altered AtCor15 expression in Arabidopsis. In the present study, the function of PsCor413im1 was further investigated. Transmission electron microscope observation showed that the chloroplast envelope membrane of cold-treated transgenic Arabidopsis seedlings was more stable than that of cold-treated wild-type seedlings. Subcellular localization of green fluorescent protein as a marker revealed that the N-terminal and putative third transmembrane domain (TMD) of PsCor413im1 were essential for its targeting of the chloroplast envelope membrane. Furthermore, overexpression of PsCor413im1 fragments containing N-terminal and third TMD also altered the expression of AtCor15 genes in Arabidopsis. Overall, our results suggest that PsCor413im1 may stabilize the chloroplast envelope membrane under cold stress, and its N-terminal and third TMD are important for its targeting capability and function.

Osmophores of Caryocar brasiliense (Caryocaraceae): a particular structure of the androecium that releases an unusual scent.

Caryocar brasiliense is a flagship species of the Brazilian Cerrado. It produces flowers with a strong peculiar scent, which are pollinated by bats and occasionally moths with short mouthparts. However, the cues responsible for attracting these nocturnal pollinators remain unknown. We aimed to identify osmophores of C. brasiliense, describe the ultrastructure of the cells involved in the synthesis and release of floral odour, and identify the constituents of the floral bouquet. We performed field observations and histochemical and ultrastructural analyses of flowers focusing on the androecium. Gas chromatography-mass spectrometry was used to analyse the scents emitted. Filament epidermal cells were found to possess an unusual shape and be responsible for the main production and release of odour. These cells, called foraminous cells, are elongate and possess pores where their cell walls are abruptly thin. The cuticle is practically absent over the pores, which facilitates odour emission. The foraminous cells have conspicuous nuclei and organelle-rich cytoplasm where oil droplets can be seen prior to anthesis. The features of these cells remain similar during anthesis, but many vesicles fuse with the plasma membrane and the number of oil droplets in the cytosol decreases. Twenty-nine components were found in the scent, especially fatty acid derivatives and N- and S-bearing compounds. Our analyses revealed that the androecium of C. brasiliense has a particular structure that acts as an osmophore. The scent from the androecium resembles that of the entire flower, which is an unprecedented finding for a plant with single flowers as the pollination unit.

Inside the horn of plenty: Leaf-mining micromoth manipulates its host plant to obtain unending food provisioning.

Leaves represent the main resource for herbivorous insects and their performances are mainly a function of leaf nutritional quality. Two feeding strategies are known to optimize the exploitation of leaf resources: leaf-miners that selectively feed on tissues of high nutritional quality and gall-inducers that induce the development of a new tissue showing an enhanced nutritional value. Some leaf-miners are known to also manipulate their nutritional environment, but do not affect plant development. Cases of callus proliferation in leaf-mines have been reported, however, the direct role of the insect in the formation of additional plant cells and the nutritional function of this tissue have never been established. Using an experimental approach, we show that leaf-mining larvae of micromoth, Borboryctis euryae (Lepidoptera: Gracillariidae), that grow on Eurya japonica (Pentaphylacaceae), actively induce callus proliferation within their leaf-mine at the fourth instar. We experimentally demonstrated that, at this developmental stage, the larva feeds exclusively on this newly formed tissue and feeding of the tissue is essential for completing larval stage. Phenological census revealed considerable expansion and variation of fourth instar duration caused by the continuous production of callus. We propose here the "cornucopia" hypothesis which states that the newly produced callus induced by the leaf-mining larvae provides virtually unending nourishment, which in turn allows flexible larval development time. This represents the first example of a leaf-miner manipulating plant development to its benefit, like a gall-inducer. We propose to name this life style "mine-galler".

Climatic niche comparison among ploidal levels in the classic autopolyploid system, Galax urceolata.

PREMISE OF THE STUDY: Autopolyploidy, or whole-genome duplication within a species, leads to closely related cytotypes in one geographic location. One hypothesized mechanism by which autopolyploids become established is climatic niche divergence from their diploid progenitor. Here we tested this hypothesis in diploid, triploid, and tetraploid Galax urceolata (Diapensiaceae) and predicted the effects of climate change on the relative distributions of these cytotypes. METHODS: We investigated whether climatic niche divergence has shaped the current distributions of Galax urceolata cytotypes in eastern North America using climatic niche modeling, multivariate analyses of environmental space, and geographic range analyses. We then projected the models of the three cytotypes onto an ensemble of future climate maps to determine how the distributions might be altered over time. KEY RESULTS: All cytotypes are geographically sympatric; however, climatic niche contraction and a slight niche shift of the tetraploids was observed relative to that of the diploids. Climate projections for all diploid and tetraploid cytotypes showed substantial range contraction without much, or any, range shift, suggesting that Galax urceolata will likely go extinct in nature as mountain refugia become warmer. CONCLUSIONS: Galax urceolata tetraploids occupy a slightly wetter habitat than that of their diploid progenitors. While we cannot take into account future adaptation, our models suggest extensive decreases in range distributions for both diploid and tetraploid G. urceolata based on climate change projections. Galax urceolata may therefore be under extreme threat due to loss of suitable habitat, and conservation efforts will be needed.

Clines in traits compared over two decades in a plant hybrid zone.

Background and Aims: Clines in traits across hybrid zones reflect a balance between natural selection and gene flow. Changes over time in average values for traits, and especially the shapes of their clines, are rarely investigated in plants, but could result from evolution in an unstable hybrid zone. Differences in clines between floral and vegetative traits could indicate different strengths of divergent selection. Methods: Five floral and two vegetative traits were measured in 12 populations along an elevational gradient spanning a natural hybrid zone between Ipomopsis aggregata and Ipomopsis tenuituba. We compared clines in the floral traits with those measured 25 years ago. Observed changes in mean trait values were compared with predictions based on prior estimates of natural selection. We also compared the steepness and position of clines between the floral and vegetative traits. Key Results: Corolla length has increased over five generations to an extent that matches predictions from measurements of phenotypic selection and heritability. The shape of its cline, and that of other traits, has not changed detectably. Clines varied across traits, but not all floral traits showed steeper clines than did vegetative traits. Both suites of morphological traits had steeper clines than did neutral molecular markers. Conclusions: The increase in corolla length provides a rare example of a match between predicted and observed evolution of a plant trait in natural populations. The clinal properties are consistent with the hypothesis that habitat-mediated divergent selection on vegetative traits and pollinator-mediated selection on floral traits both maintain species differences across the hybrid zone.

Consequences of secondary nectar robbing for male components of plant reproduction.

PREMISE OF THE STUDY: Organisms engage in multiple species interactions simultaneously. While pollination studies generally focus on plants and pollinators exclusively, secondary robbing, a behavior that requires other species (primary robbers) to first create access holes in corollas, is common. It has been shown that secondary robbing can reduce plants' female fitness; however, we lack knowledge about its impact on male plant fitness. METHODS: We experimentally simulated primary and secondary robbing in the monocarpic perennial Ipomopsis aggregata (Polemoniaceae), quantifying indirect effects on pollinator-mediated pollen (dye) donation. We also assessed whether continual nectar removal via the floral opening has similar effects on hummingbird-pollinator behavior as continual secondary robbing through robber holes. KEY RESULTS: We found no significant indirect effects of secondary robbing on a component of Ipomopsis male fitness. Although robbing did reduce pollen (dye) donation due to avoidance of robbed plants by pollinating hummingbirds, pollen donation did not differ between the two robbing treatments. The effects of secondary robbing on hummingbird behavior resembled effects of chronic nectar removal by pollinators. Our results indicate that hummingbird pollinators may use a combination of cues, including cues given by the presence or absence of nectar, to make foraging decisions. CONCLUSIONS: Combined with prior research, this study suggests that secondary robbing is less costly to a component of male fitness than to female fitness in Ipomopsis, broadening our knowledge of the overall costs of mutualism exploitation to total plant fitness.

A novel cold-regulated gene from Phlox subulata, PsCor413im1, enhances low temperature tolerance in Arabidopsis.

Low temperature stress adversely affects plant growth, development, and crop productivity. Analysis of the function of genes in the response of plants to low temperature stress is essential for understanding the mechanism of chilling and freezing tolerance. In this study, PsCor413im1, a novel cold-regulated gene isolated from Phlox subulata, was transferred to Arabidopsis to investigate its function under low temperature stress. Real-time quantitative PCR analysis revealed that PsCor413im1 expression was induced by cold and abscisic acid. Subcellular localization revealed that PsCor413im1-GFP fusion protein was localized to the periphery of the chloroplast, consistent with the localization of chloroplast inner membrane protein AtCor413im1, indicating that PsCor413im1 is a chloroplast membrane protein. Furthermore, the N-terminal of PsCor413im1 was determined to be necessary for its localization. Compared to the wild-type plants, transgenic plants showed higher germination and survival rates under cold and freezing stress. Moreover, the expression of AtCor15 in transgenic plants was higher than that in the wild-type plants under cold stress. Taken together, our results suggest that the overexpression of PsCor413im1 enhances low temperature tolerance in Arabidopsis.

Caryocar coriaceum (Caryocaraceae) diaspore removal and dispersal distance on the margin and in the interior of a Cerrado area in Northeastern Brazil / Remoción y distancia de dispersión de las diásporas de Caryocar coriaceum (Caryocaraceae) en el margen e interior de un área de Cerrado, Noreste de Brasil

Abstract:The seed dispersal process is a crucial stage in plant regeneration and maintenance of forest biological diversity. While the number of removed seeds is quantitative, the distance to which a seed is removed from its origin is qualitative, because it affects the probability that a seed will germinate and recruit to the next life stage. However, the creation of forest margins can negatively affect the seed dispersal process, especially for largediaspore plant species. In this study, the diaspore removal and dispersal distance of Caryocar coriaceum, a tree with large diaspores that is in danger of extinction, were analyzed. The study was conducted for two consecutive years in a protected forest in Northeastern Brazil. Each year, 1 200 diaspores with a nylon wire and a satin tape yellow were used and equally distributed in 120 experimental stations established on the forest margin and in the interior. During the first year of the study, no differences in diaspore removal and dispersal distance were found among the investigated environments. However, for the second year of the study, the number of removed diaspores differed significantly; nevertheless, the dispersal distance was not different between the forest margin and the interior. The low diaspore removal percentages suggest that species recruitment may be compromised because the diaspore accumulation close to the relatives enables higher fungi and insect attack. In addition, most of the few removed diaspores were found at short distances from their sources (up to 5 m), which can lead to low genetic variability. Virtually no diaspore was found buried by hoarding rodents, and no diaspore was found preyed upon by these animals. Evidence found in this study suggests the local loss of species dispersers, which can compromise the maintenance of forest biological diversity. Rev. Biol. Trop. 64 (3): 1117-1127. Epub 2016 September 01.

ResumenEl proceso de dispersión de semillas es una etapa crucial en la regeneración de las plantas y el mantenimiento de la diversidad biológica de los bosques. Mientras que el número de semillas removidas es cuantitativo, la distancia a la que se elimina una semilla desde su origen es cualitativa, porque afecta la probabilidad de que una semilla germine y sea reclutada para la próxima etapa del ciclo de vida. Sin embargo, la creación de márgenes de los bosques puede afectar negativamente el proceso de dispersión, especialmente para especies de plantas con diásporas grandes. En este estudio, analizamos la remoción diásporas y la distancia de dispersión de C. coriaceum, un árbol de diásporas grandes que está en peligro de extinción. El estudio se llevó a cabo durante dos años consecutivos en un bosque protegido en el Noreste de Brasil. Cada año, un total de 1 200 diásporas fueron igualmente distribuidas, con un hilo de nylon y una cinta amarilla de satén, en 120 estaciones experimentales establecidas en el margen y en el interior del bosque. En el primer año del estudio, no se encontraron diferencias en la remocíon de las diásporas y la distancia de dispersión entre los ambientes estudiados. Sin embargo, en el segundo año el número de diásporas retiradas difería significativamente, pero la distancia de dispersión no fue diferente entre el margen de bosques y el interior. El bajo porcentaje de remoción encontrado sugiere que el reclutamiento de especies puede estar comprometido, ya que la acumulación de diásporas próximas unas de otras es propicio para un mayor ataque de hongos e insectos. Además, la mayoría de las pocas diásporas removidas fueron encontradas a pequeñas distancias de su origen (hasta 5 m), lo que puede generar una baja variabilidad genética. Prácticamente ninguna diáspora fue encontrada enterrada por los roedores recolectores y ninguna estaba depredada por estos animales. La evidencia encontrada en este estudio indica la pérdida local de dispersores de las especies, lo que puede poner en peligro el mantenimiento de la diversidad biológica de los bosques.

Caryocar coriaceum (Caryocaraceae) diaspore removal and dispersal distance on the margin and in the interior of a Cerrado area in Northeastern Brazil.

The seed dispersal process is a crucial stage in plant regeneration and maintenance of forest biological diversity. While the number of removed seeds is quantitative, the distance to which a seed is removed from its origin is qualitative, because it affects the probability that a seed will germinate and recruit to the next life stage. However, the creation of forest margins can negatively affect the seed dispersal process, especially for large-diaspore plant species. In this study, the diaspore removal and dispersal distance of Caryocar coriaceum, a tree with large diaspores that is in danger of extinction, were analyzed. The study was conducted for two consecutive years in a protected forest in Northeastern Brazil. Each year, 1 200 diaspores with a nylon wire and a satin tape yellow were used and equally distributed in 120 experimental stations established on the forest margin and in the interior. During the first year of the study, no differences in diaspore removal and dispersal distance were found among the investigated environments. However, for the second year of the study, the number of removed diaspores differed significantly; nevertheless, the dispersal distance was not different between the forest margin and the interior. The low diaspore removal percentages suggest that species recruitment may be compromised because the diaspore accumulation close to the relatives enables higher fungi and insect attack. In addition, most of the few removed diaspores were found at short distances from their sources (up to 5 m), which can lead to low genetic variability. Virtually no diaspore was found buried by hoarding rodents, and no diaspore was found preyed upon by these animals. Evidence found in this study suggests the local loss of species dispersers, which can compromise the maintenance of forest biological diversity.

Floral acoustics: conspicuous echoes of a dish-shaped leaf attract bat pollinators.

The visual splendor of many diurnal flowers serves to attract visually guided pollinators such as bees and birds, but it remains to be seen whether bat-pollinated flowers have evolved analogous echo-acoustic signals to lure their echolocating pollinators. Here, we demonstrate how an unusual dish-shaped leaf displayed above the inflorescences of the vine Marcgravia evenia attracts bat pollinators. Specifically, this leaf's echoes fulfilled requirements for an effective beacon, that is, they were strong, multidirectional, and had a recognizable invariant echo signature. In behavioral experiments, presence of the leaves halved foraging time for flower-visiting bats.

[Effects of light intensity on seed germination and seedling early growth of Shorea wantianshuea].

With shading experiment, this paper studied the seed germination and seedling early growth of Shorea wantianshuea, a representative species in Xishuangbanna seasonal rain forest under different light intensity and forest habitat. The results showed that both 100% light and deepest shading did not facilitate the germination of S. wantianshuea seed, while medium shading accelerated the seed germination. Gap center was more appropriate to the seed germination than gap edge and under-canopy. The seeds in gap center had the highest germination rate and rapid germination. Under partially shading, the seedling height, diameter of basal stem, and leaf area per seedling were in their peak, while the root-shoot ratio decreased with increasing shading and achieved its maximum under 100% light. The specific leaf area increased with increasing shading in a certain range, and peaked under 4.2% light. Gap center was beneficial to the early seedling growth, with all growth parameters except root-shoot ratio peaked there. The effects of environmental factors, mainly light intensity and quality, on the seed germination and seedling growth of S. wantianshuea were discussed.

Effects of flowering tree density on the mating system and gene flow in Shorea leprosula (Dipterocarpaceae) in Peninsular Malaysia.

Pristine tropical rainforests in Southeast Asia have rich species diversity and are important habitats for many plant species. However, the extent of these forests has declined in recent decades and they have become fragmented due to human activities. These developments may reduce the genetic diversity of species within them and, consequently, the species' ability to adapt to environmental changes. Our objective in the study presented here was to clarify the effect of tree density on the genetic diversity and gene flow patterns of Shorea leprosula Miq. populations in Peninsular Malaysia. For this purpose, we related genetic diversity and pollen flow parameters of seedling populations in study plots to the density of mature trees in their vicinity. The results show that gene diversity and allelic richness were not significantly correlated to the mature tree density. However, the number of rare alleles among the seedlings and the selfing rates of the mother trees were negatively correlated with the density of the adult trees. Furthermore, in a population with high mature tree density pollination distances were frequently <200 m, but in populations with low adult tree density the distances were longer. These findings suggest that the density of flowering trees affects selfing rates, gene flow and, thus, the genetic diversity of S. leprosula populations. We also found an individual S. leprosula tree with a unique reproductive system, probably apomictic, mating system.

Changes in photosynthesis and leaf characteristics with tree height in five dipterocarp species in a tropical rain forest.

Variations in leaf photosynthetic, morphological and biochemical properties with increasing plant height from seedlings to emergent trees were investigated in five dipterocarp species in a Malaysian tropical rain forest. Canopy openness increased significantly with tree height. Photosynthetic properties, such as photosynthetic capacity at light saturation, light compensation point, maximum rate of carboxylation and maximum rate of photosynthetic electron transport, all increased significantly with tree height. Leaf morphological and biochemical traits, such as leaf mass per area, palisade layer thickness, nitrogen concentration per unit area, chlorophyll concentration per unit dry mass and chlorophyll to nitrogen ratio, also changed significantly with tree height. Leaf properties had simple and significant relationships with tree height, with few intra- and interspecies differences. Our results therefore suggest that the photosynthetic capacity of dipterocarp trees depends on tree height, and that the trees adapt to the light environment by adjusting their leaf morphological and biochemical properties. These results should aid in developing models that can accurately estimate carbon dioxide flux and biomass production in tropical rain forests.

Physiological and ecological significance of sunflecks for dipterocarp seedlings.

Irradiance is highly dynamic in many plant canopies. Photosynthesis during sunflecks provides 10-90% of daily carbon gain. The survivorship of tree seedlings in the deeply shaded understorey of tropical rain forests is limited by their ability to maintain a positive carbon balance. Dipterocarp seedlings from the SE Asian rain forest were used as a model system to test novel aspects of the physiological and ecological significance of sunflecks. First, understorey seedlings experienced leaf temperatures up to 38 degrees C in association with sunflecks. Under controlled environment conditions, the inhibition of carbon gain at 38 degrees C, compared with 28 degrees C, was significantly greater during a sequence of sunflecks (-59%), than under uniform irradiance (-40%), providing the same total photosynthetic photon flux density (PPFD). Second, the relative enhancement effects of elevated [CO2] were greater under sunflecks (growth +60%, carbon gain +89%), compared with uniform irradiance (growth +25%, carbon gain +59%), supplying the same daily PPFD. Third, seedling growth rates in the forest understorey were 4-fold greater under a dynamic irradiance treatment characterized by long flecks, compared with a regime of short flecks. Therefore, stresses associated with dynamic irradiance may constrain photosynthetic carbon gain. Additionally, seedling photosynthesis and growth may be more responsive to interactions with abiotic factors, including future changes in climate, than previously estimated. The sensitivity of seedling growth to varying patterns of dynamic irradiance, and the increased likelihood of species-specific responses through interactions with environmental factors, indicates the potential for sunflecks to influence regeneration processes, and hence forest structure and composition.

Variation in pollen dispersal between years with different pollination conditions in a tropical emergent tree.

We examined differences in pollen dispersal efficiency between 2 years in terms of both spatial dispersal range and genetic relatedness of pollen in a tropical emergent tree, Dipterocarpus tempehes. The species was pollinated by the giant honeybee (Apis dorsata) in a year of intensive community-level mass-flowering or general flowering (1996), but by several species of moths in a year of less-intensive general flowering (1998). We carried out paternity analysis based on six DNA microsatellite markers on a total of 277 mature trees forming four spatially distinct subpopulations in a 70 ha area, and 147 and 188 2-year-old seedlings originating from seeds produced in 1996 and 1998 (cohorts 96 and 98, respectively). Outcrossing rates (0.93 and 0.96 for cohorts 96 and 98, respectively) did not differ between years. Mean dispersal distances (222 and 192 m) were not significantly different between the 2 years but marginally more biased to long distance in 1996. The mean relatedness among cross-pollinated seedlings sharing the same mothers in cohort 96 was lower than that in cohort 98. This can be attributed to the two facts that the proportion of intersubpopulations pollen flow among cross-pollination events was marginally higher in cohort 96 (44%) than in cohort 98 (33%), and that mature trees within the same subpopulations are genetically more related to each other than those between different subpopulations. We conclude that D. tempehes maintained effective pollen dispersal in terms of outcrossing rate and pollen dispersal distance in spite of the large difference in foraging characteristics between two types of pollinators. In terms of pollen relatedness, however, a slight difference was suggested between years in the level of biparental inbreeding.

Interspecific variation of photosynthesis and leaf characteristics in canopy trees of five species of Dipterocarpaceae in a tropical rain forest.

Photosynthetic rate, nitrogen concentration and morphological properties of canopy leaves were studied in 18 trees, comprising five dipterocarp species, in a tropical rain forest in Sarawak, Malaysia. Photosynthetic rate at light saturation (Pmax) differed significantly across species, varying from 7 to 18 micro mol m(-2) s(-1). Leaf nitrogen concentration and morphological properties, such as leaf blade and palisade layer thickness, leaf mass per area (LMA) and surface area of mesophyll cells per unit leaf area (Ames/A), also varied significantly across species. Among the relationships with leaf characteristics, Pmax had the strongest correlation with leaf mesophyll parameters, such as palisade cell layer thickness (r2 = 0.76, P < 0.001) and Ames/A (r2 = 0.73, P < 0.001). Leaf nitrogen concentration and Pmax per unit area also had a significant but weaker correlation (r2 = 0.46, P < 0.01), whereas Pmax had no correlation, or only weakly significant correlations, with leaf blade thickness and LMA. Shorea beccariana Burck, which had the highest P(max) of the species studied, also had the thickest palisade layer, with up to five or more layers. We conclude that interspecific variation in photosynthetic capacity in tropical rain forest canopies is influenced more by leaf mesophyll structure than by leaf thickness, LMA or leaf nitrogen concentration.

Spatial structure and genetic diversity of two tropical tree species with contrasting breeding systems and different ploidy levels.

Analyses of the spatial distribution pattern, spatial genetic structure and of genetic diversity were carried out in two tropical tree species with contrasting breeding systems and different ploidy levels using a 50-ha demographic plot in a lowland dipterocarp forest in Peninsular Malaysia. Shorea leprosula is a diploid and predominantly outcrossed species, whereas S. ovalis ssp. sericea is an autotetraploid species with apomictic mode of reproduction. Genetic diversity parameters estimated for S. leprosula using microsatellite were consistently higher than using allozyme. In comparisons with S. leprosula and other tropical tree species, S. ovalis ssp. sericea also displayed relatively high levels of genetic diversity. This might be explained by the lower pressure of genetic drift due to tetrasomic inheritance, and for autotetraploids each locus can accommodate up to four different alleles and this allows maintenance of more alleles at individual loci. The observed high levels of genetic diversity in S. ovalis ssp. sericea can also be due to a random retention of more heterogeneous individuals in the past, and the apomictic mode of reproduction might be an evolutionary strategy, which allows the species to maintain high levels of genetic diversity. The spatial distribution pattern analyses of both species showed significant levels of aggregation at small and medium but random distribution at the big diameter-class. The decrease in magnitude of spatial aggregation from small- to large-diameter classes might be due to compensatory mortality during recruitment and survival under competitive thinning process. Spatial genetic structure analyses for both species revealed significant spatial genetic structure for short distances in all the three diameter-classes. The magnitude of spatial genetic structure in both species was observed to be decreasing from smaller- to larger-diameter classes. The high spatial genetic structuring observed in S. ovalis ssp. sericea at the small-diameter class is due primarily to limited seed dispersal and apomictic mode of reproduction. The similar observation in S. leprosula, however, can be explained by limited seed and pollen dispersal, which supports further the fact that the species is pollinated by weak fliers, mainly of Thrips and Megalurothrips in the lowland dipterocarp forest.

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.