Spatial Structure of Seagrass Suggests That Size-Dependent Plant Traits Have a Strong Influence on the Distribution and Maintenance of Tropical Multispecies Meadows

Background

Seagrass species in the tropics occur in multispecies meadows. How these meadows are maintained through species co-existence and what their ecological drivers may be has been an overarching question in seagrass biogeography. In this study, we quantify the spatial structure of four co-existing species and infer potential ecological processes from these structures.

Methods and Results

Species presence/absence data were collected using underwater towed and dropped video cameras in Pulau Tinggi, Malaysia. The geostatistical method, utilizing semivariograms, was used to describe the spatial structure of Halophila spp, Halodule uninervis, Syringodium isoetifolium and Cymodocea serrulata. Species had spatial patterns that were oriented in the along-shore and across-shore directions, nested with larger species in meadow interiors, and consisted of multiple structures that indicate the influence of 2–3 underlying processes. The Linear Model of Coregionalization (LMC) was used to estimate the amount of variance contributing to the presence of a species at specific spatial scales. These distances were <2.5 m (micro-scale), 2.5–50 m (fine-scale) and >50 m (broad-scale) in the along-shore; and <2.5 m (micro-scale), 2.5–140 m (fine-scale) and >140 m (broad-scale) in the across-shore. The LMC suggests that smaller species (Halophila spp and H. uninervis) were most influenced by broad-scale processes such as hydrodynamics and water depth whereas large, localised species (S. isoetifolium and C. serrulata) were more influenced by finer-scale processes such as sediment burial, seagrass colonization and growth, and physical disturbance.

Conclusion

In this study, we provide evidence that spatial structure is distinct even when species occur in well-mixed multispecies meadows, and we suggest that size-dependent plant traits have a strong influence on the distribution and maintenance of tropical marine plant communities. This study offers a contrast from previous spatial models of seagrasses which have largely focused on monospecific temperate meadows.     

Seagrass landscapes: a terrestrial approach to the marine subtidal environment

Subtidal seagrass habitats are prime candidates for the application of principles derived from landscape ecology. Although seagrass systems are relatively simple compared to their terrestrial counterparts in terms of species diversity and structural complexity, seagrasses do display variation in spatial patterns over a variety of scales. The presence of a moving water layer and its influence on faunal dispersal may be a distinguishing feature impacting ecological processes in the subtidal zone. Studying seagrass-dominated landscapes may provide a novel approach to investigating questions regarding self-similarity of spatial patterns, and offers a new perspective for analysing habitat change in a variety of marine environments.     

Grassland plant functional groups exhibit distinct time-lags in response to historical landscape change

Recent studies have shown significant impacts of past landscapes on present distributions of species, and discussed the existence of an extinction debt. Understanding of the processes building an extinction debt is fundamentally important for explaining present and future diversity patterns of species in fragmented landscapes. Few empirical studies, however, have examined the responses of different plant functional groups (PFGs) to historical landscape changes. We aimed to reveal PFG-based differences in species’ persistence by focusing on their vegetative, reproductive, and dispersal traits. We examined whether the present distributions of PFGs of grassland species in the edges of remnant woodlands established on former semi-natural grasslands are related to the past surrounding landscapes at different time periods and spatial scales. The effects of past landscapes varied significantly among the PFGs. Richness of short, early flowering forbs and tall, late-flowering, wind-dispersed forbs showed significant positive relationships with the surrounding habitat proportions more than 50 years ago (the 1950s) and at wide spatial scales (more than 1 km²). Richness of tall, late-flowering forbs with unassisted and other types of dispersal mechanisms showed positive relationships with the surrounding habitat proportions in recent times (the 1970s) and at smaller spatial scales (0.25 km²). Our results suggested that plant growth form, flowering season and dispersal ability—with additional information on seed bank persistence—can be good indicators for identifying species’ specific sensitivity to surrounding habitat loss. Trait-based approaches can be useful for understanding present and future distributions of grassland species with different persistence strategies in human-modified landscapes.     

Animal movement in dynamic landscapes: interaction between behavioural strategies and resource distributions

Most ecological and evolutionary processes are thought to critically depend on dispersal and individual movement but there is little empirical information on the movement strategies used by animals to find resources. In particular, it is unclear whether behavioural variation exists at all scales, or whether behavioural decisions are primarily made at small spatial scales and thus broad-scale patterns of movement simply reflect underlying resource distributions. We evaluated animal movement responses to variable resource distributions using the grey teal (Anas gracilis) in agricultural and desert landscapes in Australia as a model system. Birds in the two landscapes differed in the fractal dimension of their movement paths, with teal in the desert landscape moving less tortuously overall than their counterparts in the agricultural landscape. However, the most striking result was the high levels of individual variability in movement strategies, with different animals exhibiting different responses to the same resources. Teal in the agricultural basin moved with both high and low tortuosity, while teal in the desert basin primarily moved using low levels of tortuosity. These results call into question the idea that broad-scale movement patterns simply reflect underlying resource distributions, and suggest that movement responses in some animals may be behaviourally complex regardless of the spatial scale over which movement occurs.     

Distribution and dispersal of desert mistletoe is scale-dependent, hierarchically nested

Spatial patterns are important to many ecological processes, and scale is a critical component of both patterns and processes. I examined the pattern and scale of the spatial distribution of infection of host plants by the desert mistletoe, Phoradendron californicum, in a landscape that spans several square kilometers. I also studied the relationship between mistletoe infection and seed dispersal. I found elevated seed rain in areas with a high prevalence of mistletoes and I found that a greater proportion of trees receive seeds than are infected, suggesting that mistletoes will be aggregated in space. Using nested analysis of variance and variograms, I found that mistletoe infections were distributed in hierarchical patches. Mistletoes were aggregated within trees and mistletoe prevalence was correlated at scales of <1500 m, and at scales >4000 m. Patterns at the largest scales were correlated with elevation: sites at higher elevations showed reduced mistletoe infection compared to those at lower elevations. I propose that at small scales, mistletoe distributions are primarily the result of aggregation of seed-dispersing birds, and that the elevational effect could reflect the recent colonization of higher elevations by the mistletoes' mesquite hosts or the limits of the mistletoes' physiological tolerance to freezing-induced cavitation.     

Coherence and discontinuity in the scaling of species' distribution patterns

The spatial distribution of a species can be characterized at many different spatial scales, from fine-scale measures of local population density to coarse-scale geographical-range structure. Previous studies have shown a degree of correlation in species' distribution patterns across narrow ranges of scales, making it possible to predict fine-scale properties from coarser-scale distributions. To test the limits of such extrapolation, we have compiled distributional information on 16 species of British plants, at scales ranging across six orders of magnitude in linear resolution (1 m to 100 km). As expected, the correlation between patterns at different spatial scales tends to degrade as the scales become more widely separated. There is, however, an abrupt breakdown in cross-scale correlations across intermediate (ca. 0.5 km) scales, suggesting that local and regional patterns are influenced by essentially non-overlapping sets of processes. The scaling discontinuity may also reflect characteristic scales of human land use in Britain, suggesting a novel method for analysing the 'footprint' of humanity on a landscape.     

The tempo and mode of gopher mound production in a tallgrass prairie remnant

To gain better insight into how small-scale disturbances might affect ecological processes, such as the maintenance of plant species diversity, we conducted a two-year study characterizing spatio-temporal patterns of gopher mound production on a tallgrass prairie remnant located at Anderson Prairie, Iowa. USA. The locations of all newly produced gopher mounds were mapped on two 80 × 80 m permanent plots. We used these data to characterize spatio-temporal patterns of mound production across a range of scales. We found that mound production was highly clustered at scales of < 8 m over short periods of time (< 2 weeks), but shifted in location over a 3–4 weeks time period, resulting in a clustered pattern at scales of < 20 m over longer time periods (up to the 2 yr of the study). We also found that patterns of mound production at intermediate spatial scales (> 20 m) remained fairly static over time, although they differed significantly from site to site. The results of this study suggest that small-scale patterns of variability in mound production may increase habitat variability over very short spatial scales, possibly providing a mechanism that can enhance the development and maintenance of species diversity.     

基于零模型的宁夏荒漠草原优势种群点格局分析

植物种群空间分布格局是多种生态过程综合作用的结果。明确植物优势种群个体的空间分布格局与形成机制有助于认识种群生态适应对策与群落多样性维持机制。以宁夏荒漠草原优势种群蒙古冰草、短花针茅、牛枝子和牛心朴子为研究对象,采用完全空间随机零模型分析其种群空间分布格局特征,并通过异质泊松零模型与泊松聚块零模型探讨生境异质性、扩散限制等因子在其空间分布格局形成过程中的作用。结果显示:(1)完全空间随机零模型下,4个物种在4 m尺度范围内表现为聚集分布,随尺度增大,逐渐过渡到随机分布和均匀分布。(2)在排除生境异质性的异质泊松零模型下,蒙古冰草种群在整个研究尺度上表现为随机分布;牛枝子、短花针茅和牛心朴子种群仅分别在0—0.2、0.1—0.4 m与0—0.2 m尺度范围内发生偏离,表现为均匀分布与聚集分布,其他尺度均为随机分布。(3)在排除扩散限制的泊松聚块零模型下,所研究种群均表现为随机分布。综上,荒漠草原优势种群在小尺度范围内主要表现为聚集分布;生境异质性与扩散限制均是驱动其空间分布格局形成的重要因子,相对而言,小尺度空间范围内扩散限制的作用更为显著。     

Habitat selection behaviour links local and regional scales in aquatic systems

The role of habitat selection behaviour in the assembly of natural communities is an increasingly important theme in ecology. At the same time, ecologists and conservation biologists are keenly interested in scale and how processes at scales from local to regional interact to determine species distributions and patterns of biodiversity. How important is habitat selection in generating observed patterns of distribution and diversity at multiple spatial scales? In theory, habitat selection in response to interacting species can generate both positive and negative covariances among species distributions and create the potential to link processes of community assembly across multiple scales. Here I demonstrate that habitat selection by treefrogs in response to the distribution of fish predators functions at both the regional scale among localities and the local scale among patches within localities, implicating habitat selection as a critical link between local communities and the regional dynamics of metacommunities in complex landscapes.     

Mountain metacommunities: climate and spatial connectivity shape ant diversity in a complex landscape

Understanding what drives biodiversity patterns across scales is a central goal of ecology. Both environmental gradients and spatial landscape structure have been found to be important factors influencing species distributions and community composition, and partly reflect the balance of underlying deterministic and stochastic community processes. In some systems, environmental gradients and spatial connectivity are intertwined in that steep environmental gradients serve as boundaries on species movements and impose environment‐derived complex spatial structure to metacommunities. Mountainous landscapes are prime examples of this, and recent theory has linked principles of geomorphology, environmental gradients, and spatial structure to make predictions for resulting community patterns. In this context, we examine variation in taxonomic and phylogenetic ant diversity patterns along a geographic transect spanning > 5000 m in elevational range in the Hengduan mountains of southern China. We found that environmental gradients dominate variation in both alpha and beta diversity in this landscape, with alpha diversity strongly declining with elevation and beta diversity driven by elevational differences. However, within an elevational band spatial connectivity predicts beta diversity better than geographic distance. Our findings deviate from theoretical predictions in several ways, notably alpha diversity is monotonically declining and within‐band beta diversity is invariant with increasing elevation. The discrepancies between theory and observation may be explained by differences in the Hengduan landscape from idealized fluvial landscapes, such as a lack of a mid‐elevation peak in connectivity, as well as evolutionary limits on the source pool of species available to populate metacommunities at different elevations. The latter is supported by variation in phylogenetic community structure with elevation. Our results demonstrate the power of conceptual, statistical, and theoretical frameworks that integrate the roles of environment and spatial structure in metacommunities, but that additional work is needed to bridge the gap between abstract theory and real systems.     

From diversity indices to community assembly processes: a test with simulated data

Ecological theory suggests that spatial distribution of biodiversity is strongly driven by community assembly processes. Thus the study of diversity patterns combined with null model testing has become increasingly common to infer assembly processes from observed distributions of diversity indices. However, results in both empirical and simulation studies are inconsistent. The aim of our study is to determine with simulated data which facets of biodiversity, if any, may unravel the processes driving its spatial patterns, and to provide practical considerations about the combination of diversity indices that would produce significant and congruent signals when using null models. The study is based on simulated species’ assemblages that emerge under various landscape structures in a spatially explicit individual‐based model with contrasting, predefined assembly processes. We focus on four assembly processes (species‐sorting, mass effect, neutral dynamics and competition colonization trade‐off) and investigate the emerging species’ distributions with varied diversity indices (alpha, beta and gamma) measured at different spatial scales and for different diversity facets (taxonomic, functional and phylogenetic). We find that 1) the four assembly processes result in distinct spatial distributions of species under any landscape structure, 2) a broad range of diversity indices allows distinguishing between communities driven by different assembly processes, 3) null models provide congruent results only for a small fraction of diversity indices and 4) only a combination of these diversity indices allows identifying the correct assembly processes. Our study supports the inference of assembly processes from patterns of diversity only when different types of indices are combined. It highlights the need to combine phylogenetic, functional and taxonomic diversity indices at multiple spatial scales to effectively infer underlying assembly processes from diversity patterns by illustrating how combination of different indices might help disentangling the complex question of coexistence.     

Effects of habitat heterogeneity at multiple spatial scales on fish community assembly

Habitat variability at multiple spatial scales may affect community structure within a given habitat patch, even within seemingly homogenous landscapes. In this context, we tested the importance of habitat variables at two spatial scales (patch and landscape) in driving fish community assembly using experimental artificial reefs constructed across a gradient of seagrass cover in a coastal bay of The Bahamas. We found that species richness and benthic fish abundance increased over time, but eventually reached an asymptote. The correlation between habitat variables and community structure strengthened over time, suggesting deterministic processes were detectable in community assembly. Abundance of benthic fishes, as well as overall community structure, were predicted by both patch- and landscape-scale variables, with the cover of seagrass at the landscape-scale emerging as the most important explanatory variable. Results of this study indicate that landscape features can drive differences in community assembly even within a general habitat type (i.e., within seagrass beds). A primary implication of this finding is that human activities driving changes in seagrass cover may cause significant shifts in faunal community structure well before complete losses of seagrass habitat.     

Spatial Ecological Hierarchies: Coexistence on Heterogeneous Landscapes via Scale Niche Diversification

Spatially heterogeneous environments are generally characterized by nested landscape patterns with resource aggregations on several scales. Empirical studies indicate that such nested landscape patterns impose selection constraints on the perceptive scales of animals, but the underlying selection mechanisms are unclear. We investigated the selection dynamics of perceptive scale within a spatial resource utilization model, where the environment is characterized by its resource distribution and species differ in their perceptive scales and resource preemption capabilities. Using three model landscapes with various resource distributions, we found that the optimal perceptive scale is determined by scale-specific attributes of the landscape pattern and that the number of coexisting species increases with the number of characteristic scales. Based on the results of this model, we argue that resource aggregations on different scales act as distinct resources and that animal species of particular perceptive scales are superior in utilizing resource aggregations of comparable spatial extent. Due to the allometric relationship between body size and perceptive scale, such fitness difference might result in discontinuous body mass distributions.     

Species interactions weakly modify climate‐induced tree co‐occurrence patterns

Aims

Species distributions are hypothesized to be underlain by a complex association of processes that span multiple spatial scales including biotic interactions, dispersal limitation, fine‐scale resource gradients and climate. Species disequilibrium with climate may reflect the effects of non‐climatic processes on species distributions, yet distribution models have rarely directly considered non‐climatic processes. Here, we use a Joint Species Distribution Model (JSDM) to investigate the influence of non‐climatic factors on species co‐occurrence patterns and to directly quantify the relative influences of climate and alternative processes that may generate correlated responses in species distributions, such as species interactions, on tree co‐occurrence patterns.

Location

US Rocky Mountains.

Methods

We apply a Bayesian JSDM to simultaneously model the co‐occurrence patterns of ten dominant tree species across the Rocky Mountains, and evaluate climatic and residual correlations from the fitted model to determine the relative contribution of each component to observed co‐occurrence patterns. We also evaluate predictions generated from the fitted model relative to a single‐species modelling approach.

Results

For most species, correlation due to climate covariates exceeded residual correlation, indicating an overriding influence of broad‐scale climate on co‐occurrence patterns. Accounting for covariance among species did not significantly improve predictions relative to a single‐species approach, providing limited evidence for a strong independent influence of species interactions on distribution patterns.

Conclusions

Overall, our findings indicate that climate is an important driver of regional biodiversity patterns and that interactions between dominant tree species contribute little to explain species co‐occurrence patterns among Rocky Mountain trees.     

Asynchronous local dynamics contributes to stability of a seagrass bed in Tokyo Bay

It is known that asynchronous temporal variations in local populations can contribute to the stability of metapopulations. However, studies evaluating the hierarchical organization of multiple spatial scales are rare for continuous marine landscapes, especially for marine vegetation such as seagrass beds. In this study, long‐term observation (26 yr) of temporal changes and nested spatial analyses were combined for an extensive seagrass meadow in Tokyo Bay, Japan, using remote sensing and geographic information system technologies. We examined how the dynamics at the whole‐bed scale (~1 km²) are related to those at a local scale (0.04 km²), and investigated the relationship between the seagrass dynamics and long‐term changes in environmental conditions using data on oceanography, water quality, and sediment dynamics. The seagrass bed size fluctuated between a maximum of 1.28 km² (in 1987) and a minimum of 0.39 km² (in 2001), with an average of 0.90 km². The temporal variation in seagrass bed size at the whole‐bed scale correlated with sand movement within the seagrass bed related to changes in the position of a sandbar. Seagrass bed size fluctuated asynchronously at a local scale. Multivariate analyses recognized clusters of local areas showing similar patterns of fluctuation. Temporal patterns in the various clusters responded differently to changes in environmental factors, e.g. the position of the sandbar was highly correlated with seagrass bed size in shallow habitats but not in deeper areas. The magnitudes of the temporal variations for the local clusters were greater than that of the entire bed, suggesting that asynchronous fluctuation in different areas of the bed plays an important role in the overall stability of the seagrass bed. The results of the present study also highlight the importance of physical processes in regulating the temporal dynamics of seagrass beds in shallow sedimentary landscapes.     

Production of diaspores at the landscape level regulates local colonization: an experiment with a spore‐dispersed moss

Effective dispersal is crucial to species inhabiting transient substrates in order for them to be able to persist in a landscape. Bryophytes, pteridophytes, lichens and fungi all have wind‐dispersed small diaspores and can be efficiently dispersed if their diaspores reach air masses above canopy height. However, empirical data on dispersal over landscape scales are scarce. We investigated how the colonization of an acrocarpous clay‐inhabiting pioneer moss, Discelium nudum, varied between sites that differed in connectivity to potential dispersal sources at spatial scales from 1 to 20 km in a region in northern Sweden. We recorded the colonization on ?25 introduced clay heaps at each of 14 experimental sites some months after the dispersal period. The colonization rate ranged from 0–82% and had a statistically significant relationship with a proxy for potential habitats (amount of clay‐dominated soil) in a buffer of 20 km radius surrounding the experimental sites (and also weakly with the amount of substrate in a 10 km buffer). There were no significant relationships between colonization rate and connectivity at smaller scales (1 and 5 km). We made a rough estimate of the number of spores available for dispersal in a landscape, given the amount of clay‐dominated soil, by recording the number of Discelium nudum colonies in two 25 × 25 km landscapes. The estimated available spore numbers in the different 20 km buffers were of the same order of magnitude as the deposition densities at the experimental sites calculated from the colonization rates. The results suggest that the spores of species with scattered occurrences and small diaspores (25 μm) in open landscapes can be deposited over extensive areas, at rates high enough to drive colonization patterns. This also implies that regional connectivity may be more important than local connectivity for these kinds of species.     

Hierarchical scales of spatial variation in the smaller surface and near-surface macrobenthos of a subtropical intertidal seagrass system in Moreton Bay,Queensland

Core samples were taken along a 4 km stretch of intertidal seagrass on North Stradbroke Island, eastern Australia, at nested scales of 1 m (stations), 150 m (sites), and 2 km (localities) to investigate the extent to which abundance, diversity, and assemblage composition of the dominant smaller members (<10 mm) of the intertidal seagrass macrobenthos vary spatially and over what scales. Gastropods and polychaetes dominated both the 91 species present and, together with decapods, also the numbers of individuals. Abundance was low (mean < 2000 individuals m⁻²) but species diversity was high (overall Simpson’s index of diversity 0.91), with 44% of species occurring only as one or two individuals, and with only two species contributing >10% to the total numbers (the microgastropod Calopia imitata and crab Enigmaplax littoralis, both little known, rarely recorded endemics). On average, a species only occurred at 6% of stations and only four occurred at >25%. Assemblages at the three localities did not vary significantly in gross ecological features (levels of species richness, faunal abundance and species diversity per component site) (ANOVA P ≫ 0.05), but did vary markedly in their composition at all spatial scales (PERMANOVA P < 0.05). Variance partitioning showed that components of total variance were least at the largest spatial scale (locality 15.9%) and greatest at the smallest scale (station 59.3%). The commoner individual species all showed random distributions at small spatial scales but clumped distributions at large spatial scales.     

Habitat and scale shape the demographic fate of the keystone sea urchin Paracentrotus lividus in Mediterranean macrophyte communities

Demographic processes exert different degrees of control as individuals grow, and in species that span several habitats and spatial scales, this can influence our ability to predict their population at a particular life-history stage given the previous life stage. In particular, when keystone species are involved, this relative coupling between demographic stages can have significant implications for the functioning of ecosystems. We examined benthic and pelagic abundances of the sea urchin Paracentrotus lividus in order to: 1) understand the main life-history bottlenecks by observing the degree of coupling between demographic stages; and 2) explore the processes driving these linkages. P. lividus is the dominant invertebrate herbivore in the Mediterranean Sea, and has been repeatedly observed to overgraze shallow beds of the seagrass Posidonia oceanica and rocky macroalgal communities. We used a hierarchical sampling design at different spatial scales (100 s, 10 s and <1 km) and habitats (seagrass and rocky macroalgae) to describe the spatial patterns in the abundance of different demographic stages (larvae, settlers, recruits and adults). Our results indicate that large-scale factors (potentially currents, nutrients, temperature, etc.) determine larval availability and settlement in the pelagic stages of urchin life history. In rocky macroalgal habitats, benthic processes (like predation) acting at large or medium scales drive adult abundances. In contrast, adult numbers in seagrass meadows are most likely influenced by factors like local migration (from adjoining rocky habitats) functioning at much smaller scales. The complexity of spatial and habitat-dependent processes shaping urchin populations demands a multiplicity of approaches when addressing habitat conservation actions, yet such actions are currently mostly aimed at managing predation processes and fish numbers. We argue that a more holistic ecosystem management also needs to incorporate the landscape and habitat-quality level processes (eutrophication, fragmentation, etc.) that together regulate the populations of this keystone herbivore.     

Bumblebees experience landscapes at different spatial scales: possible implications for coexistence

Coexistence in bumblebee communities has largely been investigated at local spatial scales. However, local resource partitioning does not fully explain the species diversity of bumblebee communities. Theoretical studies provide new evidence that partitioning of space can promote species coexistence, when species interact with their environment at different spatial scales. If bumblebee species possess specific foraging ranges, different spatial resource utilisation patterns might operate as an additional mechanism of coexistence in bumblebee communities. We investigated the effects of the landscape-wide availability of different resources (mass flowering crops and semi-natural habitats) on the local densities of four bumblebee species at 12 spatial scales (landscape sectors with 250–3,000 m radius) to indirectly identify the spatial scales at which the bumblebees perceive their environment. The densities of all bumblebee species were enhanced in landscapes with high proportions of mass flowering crops (mainly oilseed rape). We found the strongest effects for Bombus terrestris agg. and Bombus lapidarius at large spatial scales, implying foraging distances of 3,000 and 2,750 m, respectively. The densities of Bombus pascuorum were most strongly influenced at a medium spatial scale (1,000 m), and of Bombus pratorum (with marginal significance) at a small spatial scale (250 m). The estimated foraging ranges tended to be related to body and colony sizes, indicating that larger species travel over larger distances than smaller species, presumably enabling them to build up larger colonies through a better exploitation of food resources. We conclude that coexistence in bumblebee communities could potentially be mediated by species-specific differences in the spatial resource utilisation patterns, which should be considered in conservation schemes.     

Aggregated spatial distributions of species in a subtropical karst forest,southwestern China

Aims Spatial distribution patterns of species reflect not only the ecological processes but also the habitat features that are related to species distribution. In karst topography, species distribution patterns provide more specific information about their environments. The objectives of this study are as follows: (i) to analyse and explain the spatial distribution patterns of conspecific trees in an old-growth subtropical karst forest; (ii) to investigate pattern changes at different spatial scales; (iii) to test the spatial pattern similarity (or dissimilarity) between trees at different abundances, diameter at breast height classes, canopy layers and different functional groups (shade tolerance and seed dispersal mode); (iv) to examine whether habitat heterogeneity has an important effect on the species spatial distribution.Methods The spatial distributions of woody species with ≥20 individuals in a 1-ha subtropical karst forest plot at Maolan in southwestern China were quantified using the relative neighbourhood density Ω based on the average density of conspecific species in a circular neighbourhood around each species.Important findings Aggregated distribution is the dominant pattern in the karst forest, but the ratio of aggregated species in total species number decreases with an increase in spatial scale. Less abundant species are more aggregated than most abundant species. Aggregation is weaker in larger diameter classes, which is consistent with the prediction of self-thinning. Seed dispersal mode influences spatial patterns, with species dispersed by animals being less aggregated than those dispersed by wind and gravity. Other species functional traits (e.g. shade tolerance) also influence the species spatial distributions. Moreover, differences among species habitat associations, e.g. with rocky outcrops, play a significant role in species spatial distributions. These results indicate that habitat heterogeneity, seed dispersal limitation and self-thinning primarily contribute to the species spatial distributions in this subtropical karst forest.