Colin R. Maycock

Reproduction of dipterocarps during low intensity masting events in a Bornean rain forest

Abstract Question: The evolution of general flowering and mast fruiting of dipterocarps in tropical rain forest has been explained by different mechanisms. We studied whether the abundance of flowering conspecifics influences the recruitment success of tropical forest trees in Borneo. Location: Sepilok Forest Reserve, Sabah, Malaysia. Method: We examined the recruitment success of 17 species of Dipterocarpaceae in a 640-ha sample area over two flowering events in 2001–2002 using seed traps and quadrats. Seed predation experiments were used to investigate if post-dispersal seed survival was density-dependent. Results: We found a negative relationship between the percentage of flowers abscised per individual and the number of flowering conspecifics and a positive relationship between the percent of flowering trees that produced viable seeds and the number of flowering conspecifics. However, we found no evidence of decreasing pre- and post-dispersal predation with increasing numbers of flowering conspecifics. High levels of flower abscission, and pre- and post-dispersal seed mortality, resulted in near-complete recruitment failure of most species, with only three species successfully recruiting in only one year. One of these, Parashorea tomentella, satiated seed predators over a large area, while Hopea beccariana and Shorea multiflora only recruited in small isolated clumps. Seed predation experiments suggest that post-dispersal seed survival was positively density-dependent in the short-term. Conclusions: Increased density of flowering conspecifics may contribute to increased likelihood of successful cross-pollination during low intensity flowering events and, in some cases, to enhanced probability of short-term seed and seedling survival. Both processes may contribute to the evolution of mast fruiting and general flowering in Southeast Asian lowland dipterocarp forests.

Ecological implications of a flower size/number trade-off in tropical forest trees

Background In angiosperms, flower size commonly scales negatively with number. The ecological consequences of this trade-off for tropical trees remain poorly resolved, despite their potential importance for tropical forest conservation. We investigated the flower size number trade-off and its implications for fecundity in a sample of tree species from the Dipterocarpaceae on Borneo. Methodology/Principal Findings We combined experimental exclusion of pollinators in 11 species, with direct and indirect estimates of contemporary pollen dispersal in two study species and published estimates of pollen dispersal in a further three species to explore the relationship between flower size, pollinator size and mean pollen dispersal distance. Maximum flower production was two orders of magnitude greater in small-flowered than large-flowered species of Dipterocarpaceae. In contrast, fruit production was unrelated to flower size and did not differ significantly among species. Small-flowered species had both smaller-sized pollinators and lower mean pollination success than large-flowered species. Average pollen dispersal distances were lower and frequency of mating between related individuals was higher in a smaller-flowered species than a larger-flowered confamilial. Our synthesis of pollen dispersal estimates across five species of dipterocarp suggests that pollen dispersal scales positively with flower size. Conclusions and Their Significance Trade-offs embedded in the relationship between flower size and pollination success contribute to a reduction in the variance of fecundity among species. It is therefore plausible that these processes could delay competitive exclusion and contribute to maintenance of species coexistence in this ecologically and economically important family of tropical trees. These results have practical implications for tree species conservation and restoration. Seed collection from small-flowered species may be especially vulnerable to cryptic genetic erosion. Our findings also highlight the potential for differential vulnerability of tropical tree species to the deleterious consequences of forest fragmentation.

Mass fruiting in Borneo: a missed opportunity.

![Figure][1] Dipterocarp tree seedlings. Many endangered Indonesian trees rarely produce seeds. CREDIT: CAMPBELL WEBB Large-scale restoration of tropical forest is increasingly recognized as a credible option for climate change mitigation and biodiversity conservation ([ 1 ][2]–[ 3 ][3

Predicting dispersal of auto‐gyrating fruit in tropical trees: a case study from the Dipterocarpaceae

Seed dispersal governs the distribution of plant propagules in the landscape and hence forms the template on which density-dependent processes act. Dispersal is therefore a vital component of many species coexistence and forest dynamics models and is of applied value in understanding forest regeneration. Research on the processes that facilitate forest regeneration and restoration is given further weight in the context of widespread loss and degradation of tropical forests, and provides impetus to improve estimates of seed dispersal for tropical forest trees. South-East Asian lowland rainforests, which have been subject to severe degradation, are dominated by trees of the Dipterocarpaceae family which constitute over 40% of forest biomass. Dipterocarp dispersal is generally considered to be poor given their large, gyration-dispersed fruits. However, there is wide variability in fruit size and morphology which we hypothesize mechanistically underpins dispersal potential through the lift provided to seeds mediated by the wings. We explored experimentally how the ratio of fruit wing area to mass (“inverse wing loading,” IWL) explains variation in seed dispersal kernels among 13 dipterocarp species by releasing fruit from a canopy tower. Horizontal seed dispersal distances increased with IWL, especially at high wind speeds. Seed dispersal of all species was predominantly local, with 90% of seed dispersing <10 m, although maximum dispersal distances varied widely among species. We present a generic seed dispersal model for dipterocarps based on attributes of seed morphology and provide modeled seed dispersal kernels for all dipterocarp species with IWLs of 1–50, representing 75% of species in Borneo.

Long-term carbon sink in Borneo's forests halted by drought and vulnerable to edge effects

Less than half of anthropogenic carbon dioxide emissions remain in the atmosphere. While carbon balance models imply large carbon uptake in tropical forests, direct on-the-ground observations are still lacking in Southeast Asia. Here, using long-term plot monitoring records of up to half a century, we find that intact forests in Borneo gained 0.43 Mg C ha−1 per year (95% CI 0.14–0.72, mean period 1988–2010) in above-ground live biomass carbon. These results closely match those from African and Amazonian plot networks, suggesting that the world’s remaining intact tropical forests are now en masse out-of-equilibrium. Although both pan-tropical and long-term, the sink in remaining intact forests appears vulnerable to climate and land use changes. Across Borneo the 1997–1998 El Niño drought temporarily halted the carbon sink by increasing tree mortality, while fragmentation persistently offset the sink and turned many edge-affected forests into a carbon source to the atmosphere.The existence of a pan-tropical forest carbon sink remains uncertain due to the lack of data from Asia. Here, using direct on-the-ground observations, the authors confirm remaining intact forests in Borneo have provided a long-term carbon sink, but carbon net gains are vulnerable to drought and edge effects.

Genetic diversity affects seedling survival but not growth or seed germination in the Bornean endemic dipterocarp Parashorea tomentella

Background: Logging and habitat fragmentation of tropical rain forests may disrupt patterns of gene flow and genetic diversity. Consequently, inbreeding in tree populations may reduce fitness and increase extinction risks, especially among species that are predominantly outcrossing, dependent on biotic pollination and/or display limited seed dispersal such as species of the Dipterocarpaceae. Aims: To test the hypothesis that heterozygosity of individual progeny affects their likelihood of germination and the growth and survival of seedlings. Methods: Standardised measure of multilocus heterozygosity (sMLH) was estimated from seven microsatellite loci for individual progeny collected from 18 mother trees of the large dipterocarp Parashorea tomentella. The relationships among sMLH, germination and seedling growth and survival were determined for the progeny. Results: Seedling survival over 18 months increased with greater sMLH and fresh fruit weight. This result was expressed under all experimentally controlled combinations of light and nutrient availability in the nursery and in the shaded understorey of primary forest where survival overall was much lower than in the nursery. sMLH did not affect the probability of germination or seedling growth rate in any experimental treatment. Conclusions: These results provide evidence that reduced heterozygosity is associated with reduced seedling survival in a tropical forest tree species.

Using High Resolution Ecological Niche Models to Assess the Conservation Status of Dipterocarpus lamellatus and Dipterocarpus ochraceus in Sabah, Malaysia

Sabah has experienced a rapid decline in the extent of forest cover. The precise impact of habitat loss on the conservation status of the plants of Sabah is uncertain. In this study we use the niche modelling algorithm MAXENT to construct preliminary, revised and final ecological niche models for Dipterocarpus lamellatus and Dipterocarpus ochraceus and combined these models with data on current land-use to derive conservation assessments for each species. Preliminary models were based on herbarium data alone. Ground surveys were conducted to evaluate the performance of these preliminary models, and a revised niche model was generated from the combined herbarium and ground survey data. The final model was obtained by constraining the predictions of the revised models by filters. The range overlap between the preliminary and revised models was 0.47 for D. lamellatus and 0.39 for D. ochraceus, suggesting poor agreement between them. There was substantial variation in estimates of habitat loss for D. ochraceus, among the preliminary, revised and constrained models, and this has the potential to lead to incorrect threat assessments. From these estimates of habitat loss, the historic distribution and estimates of population size we determine that both species should be classified as Critically Endangered under IUCN Red List guidelines. Our results suggest that ground-truthing of ecological niche models is essential, especially if the models are being used for conservation decision making.

Are patterns of fine-scale spatial genetic structure consistent between sites within tropical tree species?

Documenting the scale and intensity of fine-scale spatial genetic structure (FSGS), and the processes that shape it, is relevant to the sustainable management of genetic resources in timber tree species, particularly where logging or fragmentation might disrupt gene flow. In this study we assessed patterns of FSGS in three species of Dipterocarpaceae (Parashorea tomentella, Shorea leprosula and Shorea parvifolia) across four different tropical rain forests in Malaysia using nuclear microsatellite markers. Topographic heterogeneity varied across the sites. We hypothesised that forests with high topographic heterogeneity would display increased FSGS among the adult populations driven by habitat associations. This hypothesis was not supported for S. leprosula and S. parvifolia which displayed little variation in the intensity and scale of FSGS between sites despite substantial variation in topographic heterogeneity. Conversely, the intensity of FSGS for P. tomentella was greater at a more topographically heterogeneous than a homogeneous site, and a significant difference in the overall pattern of FSGS was detected between sites for this species. These results suggest that local patterns of FSGS may in some species be shaped by habitat heterogeneity in addition to limited gene flow by pollen and seed dispersal. Site factors can therefore contribute to the development of FSGS. Confirming consistency in species’ FSGS amongst sites is an important step in managing timber tree genetic diversity as it provides confidence that species specific management recommendations based on species reproductive traits can be applied across a species’ range. Forest managers should take into account the interaction between reproductive traits and site characteristics, its consequences for maintaining forest genetic resources and how this might influence natural regeneration across species if management is to be sustainable.

Author Correction: Long-term carbon sink in Borneo’s forests halted by drought and vulnerable to edge effects

The original version of this Article contained an error in the third sentence of the abstract and incorrectly read “Here, using long-term plot monitoring records of up to half a century, we find that intact forests in Borneo gained 0.43 Mg C ha−1 year−1 (95% CI 0.14–0.72, mean period 1988–2010) above-ground live biomass”, rather than the correct “Here, using long-term plot monitoring records of up to half a century, we find that intact forests in Borneo gained 0.43 Mg C ha−1 year−1 (95% CI 0.14–0.72, mean period 1988–2010) in above-ground live biomass carbon”. This has now been corrected in both the PDF and HTML versions of the Article.