Tree seedlings in tropical rain forests are subject to both damage from natural enemies and intense interspecific competition. This leads to a trade-off in investment between defense and growth, and it is likely that tree species specialized to particular habitats tailor this balance to correspond with local resource availability. It has also been suggested that differential herbivore impacts among tree species may drive habitat segregation, favoring species adapted to particular resource conditions. In order to test these predictions, a reciprocal transplant experiment in Sabah, Malaysia, was established with seedlings of five species of Dipterocarpaceae. These were specialized to either alluvial (Hopea nervosa, Parashorea tomentella) or sandstone soils (Shorea multiflora, H. beccariana), or were locally absent (S. fallax). A total of 3000 seedlings were planted in paired gap and understory plots in five sites on alluvial and sandstone soils. Half of all seedlings were fertilized. Seedling growth and mortality were recorded in regular samples over 3.5 years, and rates of insect herbivore damage were estimated from censuses of foliar tissue loss on marked mature leaves and available young leaves. Greater herbivory rates on mature leaves had no measurable effects on seedling growth but were associated with a significantly increased likelihood of mortality during the following year. In contrast, new-leaf herbivory rates correlated with neither growth nor mortality. There were no indications of differential impacts of herbivory among the five species, nor between experimental treatments. Herbivory was not shown to influence segregation of species between soil types, although it may contribute toward differential survival among light habitats. Natural rates of damage were substantially lower than have been shown to influence tree seedling growth and mortality in previous manipulative studies.
There is accumulating evidence that similar suites of plant traits may affect leaf palatability and leaf litter decomposability. However, the possible association between leaf herbivory and litter decomposition rates across species in species-diverse natural ecosystems such as tropical rain forests remains unexplored, despite its importance in estimating the herbivory effects on carbon and nutrient cycling of ecosystems. We found no strong association between leaf herbivory and litter decomposition rates across 40 tree species in a Malaysian tropical rain forest, even though the leaf and litter traits were tightly correlated. This is because the leaf and litter traits related to herbivory and decomposition rates in the field were inconsistent. Leaf toughness accounted for only a small part of the variation in the herbivory rate, whereas a number of litter traits (the leaf mass per area, lignin to nitrogen ratio, and condensed tannin concentration) accurately predicted the decomposition rate across species. These results suggest that herbivory rate across species may not be strongly related to single leaf traits, probably because plant-herbivore interactions in tropical rain forests are highly diverse; on the other hand, plant-decomposer interactions are less specific and can be governed by litter chemicals. We also investigated two factors, phylogeny and tree functional types, that could affect the relationship between herbivory and decomposition across species. Phylogenetic relatedness among the species did not affect the relationship between herbivory and decomposition. In contrast, when the plants were segregated according to their leaf emergence pattern, we found a significant positive relationship between herbivory and decomposition rates for continuous-leafing species. In these species, the condensed tannin to N ratios in leaves and litter were related to herbivory and decomposition rates, respectively. However, we did not observe a similar trend for synchronous-leafing species. These results suggest that the relationship between herbivory and decomposition may be more greatly affected by functional types than by phylogenetic relatedness among species. In conclusion, our results suggest that well-defended leaves are not necessarily less decomposable litter in a tropical rain forest community, implying that herbivory may not generate positive feedback for carbon and nutrient cycling in this type of ecosystem.
Pairwise similarity coefficients are downward biased when samples only record presences and sampling is partial. A simple but forgotten index proposed by Stephen Forbes in 1907 can help solve this problem. His original equation requires knowing the number of species absent in both samples that could have been present. It is proposed that this count should simply be ignored and that the coefficient should be adjusted using a simple heuristic correction. Four analyses show that the corrected equation outperforms the Dice and Simpson indices, which are highly correlated with many others. In two-sample simulations, similarity is almost always closer to the assumed value when the species pool size and sampling intensity are varied, regardless of whether the underlying abundance distribution is uniform, log-normal, or geometric. The index is also much more robust when sampling is unequal. An analysis of bat samples from peninsular Malaysia buttresses these conclusions. The corrected coefficient also indicates that local assemblages of North American mammals are random subsamples of larger species pools by returning similarity of values of around 1, and it suggests a more consistent relationship between biome-scale comparisons and local-scale comparisons. Finally, it yields a better-dispersed pattern when the biome-scale inventories are ordinated. If these results are generalizable, then the new and old equation should see wide application, potentially taking the place of the two most commonly used alternatives (the interrelated Dice and Jaccard indices) whenever sampling is incomplete.
The data set contains images of leaf venation networks obtained from tree species in Malaysian Borneo. The data set contains 726 leaves from 295 species comprising 50 families, sampled from eight forest plots in Sabah. Image extents are approximately 1 × 1 cm, or 50 megapixels. All images contain a region of interest in which all veins have been hand traced. The complete data set includes over 30 billion pixels, of which more than 600 million have been validated by hand tracing. These images are suitable for morphological characterization of these species, as well as for training of machine-learning algorithms that segment biological networks from images. Data are made available under the Open Data Commons Attribution License. You are free to copy, distribute, and use the database; to produce works from the database; and to modify, transform, and build upon the database. You must attribute any public use of the database, or works produced from the database, in the manner specified in the license. For any use or redistribution of the database, or works produced from it, you must make clear to others the license of the database and keep intact any notices on the original database.
Tree architecture, growth, and mortality change with increasing tree size and associated light conditions. To date, few studies have quantified how size-dependent changes in growth and mortality rates co-vary with architectural traits, and how such size-dependent changes differ across species and possible light capture strategies. We applied a hierarchical Bayesian model to quantify size-dependent changes in demographic rates and correlated demographic rates and architectural traits for 145 co-occurring Malaysian rain-forest tree species covering a wide range of tree sizes. Demographic rates were estimated using relative growth rate in stem diameter (RGR) and mortality rate as a function of stem diameter. Architectural traits examined were adult stature measured as the 95-percentile of the maximum stem diameter (upper diameter), wood density, and three tree architectural variables: tree height, foliage height, and crown width. Correlations between demographic rates and architectural traits were examined for stem diameters ranging from 1 to 47 cm. As a result, RGR and mortality varied significantly with increasing stem diameter across species. At smaller stem diameters, RGR was higher for tall trees with wide crowns, large upper diameter, and low wood density. Increased mortality was associated with low wood density at small diameters, and associated with small upper diameter and wide crowns over a wide range of stem diameters. Positive correlations between RGR and mortality were found over the whole range of stem diameters, but they were significant only at small stem diameters. Associations between architectural traits and demographic rates were strongest at small stem diameters. In the dark understory of tropical rain forests, the limiting amount of light is likely to make the interspecific difference in the effects of functional traits on demography more clear. Demographic performance is therefore tightly linked with architectural traits such as adult stature, wood density, and capacity for horizontal crown expansion. The enhancement of a demographic trade-off due to interspecific variation in functional traits in the understory helps to explain species coexistence in diverse rain forests.
In Southeast Asia, biodiversity-rich forests are being extensively logged and converted to oil palm monocultures. Although the impacts of these changes on biodiversity are largely well documented, we know addition to samples we collected in 201 little about how these large-scale impacts affect freshwater trophic ecology. We used stable isotope analyses (SIA) to determine the impacts of land-use changes on the relative contribution of allochthonous and autochthonous basal resources in 19 stream food webs. We also applied compound-specific SIA and bulk-SIA to determine the trophic position of fish apex predators and meso-predators (invertivores and omnivores). There was no difference in the contribution of autochthonous resources in either consumer group (70-82%) among streams with different land-use type. There was no change in trophic position for meso-predators, but trophic position decreased significantly for apex predators in oil palm plantation streams compared to forest streams. This change in maximum food chain length was due to turnover in identity of the apex predator among land-use types. Disruption of aquatic trophic ecology, through reduction in food chain length and shift in basal resources, may cause significant changes in biodiversity as well as ecosystem functions and services. Understanding this change can help develop more focused priorities for mediating the negative impacts of human activities on freshwater ecosystems.
A survey is made of the overall pattern of distribution of freshwater fish within Malaya and also of the associations found in the three important habitat types. The lack of east-west differentiation indicates that geographical barriers are not important in determining the distributional pattern. A marked north-south differentiation probably reflects edaphic and climatic factors. Analyses based on constancy are given for the fish associations of blackwaters, ordinary streams in tree country, and riceland habitats. Each has a distinctive fish association with a relatively small number of frequent species. Blackwater associations can be divided into subassociations corresponding to different types of blackwaters. Tree country and riceland associations show some regional differentiation. The tree country association is remarkably diversified and appears to be a true association, not a fortuitous assemblage of species with similar tolerances. Adverse factors such as low oxygen and low pH probably restrict the associations found in blackwaters and ricelands. These contain both specialist and generally tolerant species, the latter becoming more important the more extreme the habitat. The riceland fauna is depauperate as compared with neighboring countries, and there are possibly vacant ecological niches.
Elucidating how dispersal and landscape connectivity influence metacommunity stability will shed light on natural processes structuring ecosystems and help prioritize conservation actions in an increasingly fragmented world. Much of the theoretical and mathematical development of the metacommunity concept has been based on simplified experimental systems or simulated data. We still have limited understanding of how variation in the habitat matrix and species-specific differences in dispersal ability contribute to metacommunity dynamics in heterogeneous landscapes. We model a metacommunity of rainforest mammals in Borneo, a tropical biodiversity hotspot, where protected areas are increasingly isolated by ongoing habitat disturbance and loss. We employ a combination of hierarchical models of local abundance, circuit-theory-based dispersal analysis, and metapopulation models. Our goal was to understand which landscape links were the most important to metapopulation persistence and metacommunity stability. Links were particularly important if they were short and connected two large patches. This was partly because only the very shortest links could be traversed by poorly dispersing species, including small herbivores such as chevrotains (Tragulus spp.) and porcupines. Links that join large patches into a "super-patch" may also promote island-mainland rather than Levins-type metapopulation dynamics for good dispersers, particularly large carnivores such as clouded leopards (Neofelis diardi) and sun bears (Helarctos malayanus), reducing metapopulation extinction risk and thereby enhancing metacommunity stability. Link importance to metacommunity stability was highly correlated between heterogeneous and homogeneous landscapes. But link importance to metapopulation capacity varied strongly across species, and the correlation between heterogeneous and homogeneous landscape matrix scenarios was low for poorly dispersing taxa. This suggests that the environmental conditions in the area between habitat patches, the landscape matrix, is important for assessing certain individual species but less so for understanding the stability of the entire metacommunity.
Apex predators can have substantial and complex ecological roles in ecosystems. However, differences in species-specific traits, population densities, and interspecific interactions are likely to determine the strength of apex predators' roles. Here we report complementary studies examining how interactions between predator per capita metabolic rate and population density influenced the biomass, population energy use, and ecological effects of apex predators on their large mammalian prey. We first investigated how large mammal prey resources and field metabolic rates of terrestrial apex predators, comprising large mammals and the Komodo dragon (Varanus komodoensis), influenced their biomass densities and population energy use requirements. We next evaluated whether Komodo dragons, like apex mammalian predators, exerted top-down regulation of their large mammal prey. Comparison of results from field studies demonstrates that Komodo dragons attain mean population biomass densities that are 5.75-231.82 times higher than that of apex mammalian predator species and their guilds in Africa, Asia, and North America. The high biomass of Komodo dragons resulted in 1.96-108.12 times greater population energy use than that of apex mammalian predators. Nevertheless, substantial temporal and spatial variation in Komodo dragon population energy use did not regulate the population growth rates of either of two large mammal prey species, rusa deer (Rusa timorensis) and wild pig (Sus scrofa). We suggest that multiple processes weaken the capacity of Komodo dragons to regulate large mammal prey populations. For example, a low per capita metabolic rate requiring an infrequent and inactive hunting strategy (including scavenging), would minimize lethal and nonlethal impacts on prey populations. We conclude that Komodo dragons differ in their predatory role from, including not being the ecological analogs of, apex mammalian predators.
The importance of lianas through time and their effect on tree reproduction are evaluated for the first time in a Southeast Asian Dipterocarp forest. We quantified flower and seed production by lianas and trees for 13 years, assessed liana loads in the crowns of all trees larger than 30 cm in diameter at breast height (1.3 m) in 2002 and 2014, and assessed levels of reproduction for the same trees during a strong general flowering event in 2014 for the 50-ha forest dynamics plot at the Pasoh Forest Reserve, Malaysia. General flowering refers to synchronous reproduction by hundreds of plant species at irregular, multiyear intervals and only occurs in Southeast Asian Dipterocarp forests. Overall, lianas were present in 50% of tree crowns and comprised 31% of flower production and 46% of seed production. Lianas reduced growth, survival, and reproduction by their host trees. Lianas were less frequent in canopy- emergent trees, Dipterocarps comprised a disproportionately large proportion of canopy emergents, and, as a consequence, lianas were less frequent in Dipterocarps than in trees from other plant families. Lianas infested the crowns of significantly fewer trees in 2014 (47.9%) than in 2002 (52.3%); however, the decrease was restricted to trees with the lightest liana loads and sample sizes and statistical power were enormous. Lianas comprised a stable proportion of flower production and a highly variable proportion of seed production from 2002 through 2013. We conclude lianas have a huge impact on trees in this forest and were a stable component of the forest between 2002 and 2014. The emergent habit and associated ability to avoid lianas might contribute to the success of the Dipterocarpaceae.
Scleractinian corals are colonial animals with a range of life-history strategies, making up diverse species assemblages that define coral reefs. We tagged and tracked ~30 colonies from each of 11 species during seven trips spanning 6 years (2009-2015) to measure their vital rates and competitive interactions on the reef crest at Trimodal Reef, Lizard Island, Australia. Pairs of species were chosen from five growth forms in which one species of the pair was locally rare (R) and the other common (C). The sampled growth forms were massive (Goniastrea pectinata [R] and G. retiformis [C]), digitate (Acropora humilis [R] and A. cf. digitifera [C]), corymbose (A. millepora [R] and A. nasuta [C]), tabular (A. cytherea [R] and A. hyacinthus [C]) and arborescent (A. robusta [R] and A. intermedia [C]). An extra corymbose species with intermediate abundance, A. spathulata was included when it became apparent that A. millepora was too rare on the reef crest, making the 11 species in total. The tagged colonies were visited each year in the weeks prior to spawning. During visits, two or more observers each took two or three photographs of each tagged colony from directly above and on the horizontal plane with a scale plate to track planar area. Dead or missing colonies were recorded and new colonies tagged to maintain ~30 colonies per species throughout the 6 years of the study. In addition to tracking tagged corals, 30 fragments were collected from neighboring untagged colonies of each species for counting numbers of eggs per polyp (fecundity); and fragments of untagged colonies were brought into the laboratory where spawned eggs were collected for biomass and energy measurements. We also conducted surveys at the study site to generate size structure data for each species in several of the years. Each tagged colony photograph was digitized by at least two people. Therefore, we could examine sources of error in planar area for both photographers and outliners. Competitive interactions were recorded for a subset of species by measuring the margins of tagged colony outlines interacting with neighboring corals. The study was abruptly ended by Tropical Cyclone Nathan (Category 4) that killed all but nine of the more than 300 tagged colonies in early 2015. Nonetheless, these data will be of use to other researchers interested in coral demography and coexistence, functional ecology, and parametrizing population, community, and ecosystem models. The data set is not copyright restricted, and users should cite this paper when using the data.
Classical ecological theory posits that species partition resources such that each species occupies a unique resource niche. In general, the availability of more resources allows more species to co-occur. Thus, a strong relationship between communities of consumers and their resources is expected. However, correlations may be influenced by other layers in the food web, or by the environment. Here we show, by studying the relationship between communities of consumers (land snails) and individual diets (from seed plants), that there is in fact no direct, or at most a weak but negative, relationship. However, we found that the diversity of the individual microbiome positively correlates with both consumer community diversity and individual diet diversity in three target species. Moreover, these correlations were affected by various environmental variables, such as anthropogenic activity, habitat island size, and a possibly important nutrient source, guano runoff from nearby caves. Our results suggest that the microbiome and the environment explain the absence of correlations between diet and consumer community diversity. Hence, we advocate that microbiome inventories are routinely added to any community dietary analysis, which our study shows can be done with relatively little extra effort. Our approach presents the tools to quickly obtain an overview of the relationships between consumers and their resources. We anticipate our approach to be useful for ecologists and environmentalists studying different communities in a local food web.
Information on tropical Asian vertebrates has traditionally been sparse, particularly when it comes to cryptic species inhabiting the dense forests of the region. Vertebrate populations are declining globally due to land-use change and hunting, the latter frequently referred as "defaunation." This is especially true in tropical Asia where there is extensive land-use change and high human densities. Robust monitoring requires that large volumes of vertebrate population data be made available for use by the scientific and applied communities. Camera traps have emerged as an effective, non-invasive, widespread, and common approach to surveying vertebrates in their natural habitats. However, camera-derived datasets remain scattered across a wide array of sources, including published scientific literature, gray literature, and unpublished works, making it challenging for researchers to harness the full potential of cameras for ecology, conservation, and management. In response, we collated and standardized observations from 239 camera trap studies conducted in tropical Asia. There were 278,260 independent records of 371 distinct species, comprising 232 mammals, 132 birds, and seven reptiles. The total trapping effort accumulated in this data paper consisted of 876,606 trap nights, distributed among Indonesia, Singapore, Malaysia, Bhutan, Thailand, Myanmar, Cambodia, Laos, Vietnam, Nepal, and far eastern India. The relatively standardized deployment methods in the region provide a consistent, reliable, and rich count data set relative to other large-scale pressence-only data sets, such as the Global Biodiversity Information Facility (GBIF) or citizen science repositories (e.g., iNaturalist), and is thus most similar to eBird. To facilitate the use of these data, we also provide mammalian species trait information and 13 environmental covariates calculated at three spatial scales around the camera survey centroids (within 10-, 20-, and 30-km buffers). We will update the dataset to include broader coverage of temperate Asia and add newer surveys and covariates as they become available. This dataset unlocks immense opportunities for single-species ecological or conservation studies as well as applied ecology, community ecology, and macroecology investigations. The data are fully available to the public for utilization and research. Please cite this data paper when utilizing the data.