Mapping species density of trees,shrubs and vines in a tropical forest,using field measurements,satellite multiespectral imagery and spatial interpolation

We estimated the number of species in a tropical forest landscape in Quintana Roo, Mexico, based on the relationship between reflectance values of satellite imagery and field measurements of plant species density (mean number of species per plot). Total species density as well as that of tree, shrub and vine species were identified from 141 sampling quadrats (16543 individuals sampled). Spatial prediction of plant diversity was performed using universal kriging. This approach considered the linear relationship between plant species density and reflectance values of Thematic Mapper™, as well as the spatial dependence of the observations. We explored the linear relationships between spectral properties of TM bands and the species density of trees, shrubs and vines, using regression analysis. We employed Akaike Information Criterion (AIC) to select a set of candidate models. Based on Akaike weights, we calculated model-averaged parameters. Linear regression between number of species and reflectance values of TM bands yielded regression residuals. We used variogram analysis to analyze the spatial structure of these residuals. Results show that accounting for spatial autocorrelation in the residual variation improved model R² from 0.57 to 0.66 for number of all species, from 0.58 to 0.65 for number of tree species, from 0.26 to 0.41 for number of shrub species and from 0.13 to 0.17 for species density of vines. The empirical models we developed can be used to predict landscape-level species density in the Yucatan Peninsula, helping to guide and evaluate management and conservation strategies.     

Effects of spatial heterogeneity on butterfly species richness in Rocky Mountain National Park, CO, USA

We investigated butterfly responses to plot-level characteristics (plant species richness, vegetation height, and range in NDVI [normalized difference vegetation index]) and spatial heterogeneity in topography and landscape patterns (composition and configuration) at multiple spatial scales. Stratified random sampling was used to collect data on butterfly species richness from seventy-six 20 × 50 m plots. The plant species richness and average vegetation height data were collected from 76 modified-Whittaker plots overlaid on 76 butterfly plots. Spatial heterogeneity around sample plots was quantified by measuring topographic variables and landscape metrics at eight spatial extents (radii of 300, 600 to 2,400 m). The number of butterfly species recorded was strongly positively correlated with plant species richness, proportion of shrubland and mean patch size of shrubland. Patterns in butterfly species richness were negatively correlated with other variables including mean patch size, average vegetation height, elevation, and range in NDVI. The best predictive model selected using Akaike’s Information Criterion corrected for small sample size (AIC_c), explained 62% of the variation in butterfly species richness at the 2,100 m spatial extent. Average vegetation height and mean patch size were among the best predictors of butterfly species richness. The models that included plot-level information and topographic variables explained relatively less variation in butterfly species richness, and were improved significantly after including landscape metrics. Our results suggest that spatial heterogeneity greatly influences patterns in butterfly species richness, and that it should be explicitly considered in conservation and management actions.     

Assessing species density and abundance of tropical trees from remotely sensed data and geostatistics

Question: What relationships exist between remotely sensed measurements and field observations of species density and abundance of tree species? Can these relationships and spatial interpolation approaches be used to improve the accuracy of prediction of species density and abundance of tree species? Location: Quintana Roo, Yucatan peninsula, Mexico. Methods: Spatial prediction of species density and abundance of species for three functional groups was performed using regression kriging, which considers the linear relationship between dependent and explanatory variables, as well as the spatial dependence of the observations. These relationships were explored using regression analysis with species density and abundance of species of three functional groups as dependent variables, and reflectance values of spectral bands, computed NDVI (normalized difference vegetation index), standard deviation of NDVI and texture measurements of Landsat 7 Thematic Mapper (TM) imagery as explanatory variables. Akaike information criterion was employed to select a set of candidate models and calculate model‐averaged parameters. Variogram analysis was used to analyze the spatial structure of the residuals of the linear regressions. Results: Species density of trees was related to reflectance values of TM4, NDVI and spatial heterogeneity of land cover types, while the abundance of species in functional groups showed different patterns of association with remotely sensed data. Models that accounted for spatial autocorrelation improved the accuracy of estimates in all cases. Conclusions: Our approach can substantially increase the accuracy of the spatial estimates of species richness and abundance of tropical tree species and can help guide and evaluate tropical forest management and conservation.     

Interactions between landcover pattern and geospatial processing methods: Effects on landscape metrics and classification accuracy

Remote sensing data is routinely used in ecology to investigate the relationship between landscape pattern as characterised by land use and land cover maps, and ecological processes. Multiple factors related to the representation of geographic phenomenon have been shown to affect characterisation of landscape pattern resulting in spatial uncertainty. This study investigated the effect of the interaction between landscape spatial pattern and geospatial processing methods statistically; unlike most papers which consider the effect of each factor in isolation only. This is important since data used to calculate landscape metrics typically undergo a series of data abstraction processing tasks and are rarely performed in isolation. The geospatial processing methods tested were the aggregation method and the choice of pixel size used to aggregate data. These were compared to two components of landscape pattern, spatial heterogeneity and the proportion of landcover class area. The interactions and their effect on the final landcover map were described using landscape metrics to measure landscape pattern and classification accuracy (response variables). All landscape metrics and classification accuracy were shown to be affected by both landscape pattern and by processing methods. Large variability in the response of those variables and interactions between the explanatory variables were observed. However, even though interactions occurred, this only affected the magnitude of the difference in landscape metric values. Thus, provided that the same processing methods are used, landscapes should retain their ranking when their landscape metrics are compared. For example, highly fragmented landscapes will always have larger values for the landscape metric “number of patches” than less fragmented landscapes. But the magnitude of difference between the landscapes may change and therefore absolute values of landscape metrics may need to be interpreted with caution. The explanatory variables which had the largest effects were spatial heterogeneity and pixel size. These explanatory variables tended to result in large main effects and large interactions. The high variability in the response variables and the interaction of the explanatory variables indicate it would be difficult to make generalisations about the impact of processing on landscape pattern as only two processing methods were tested and it is likely that untested processing methods will potentially result in even greater spatial uncertainty.     

Distribution patterns of tree species in a Malaysian tropical rain forest

Abstract. Spatial patterns of tree species were studied in a 50-ha tropical rain forest plot in the Pasoh forest, Malaysia. This forest is characterized by a high diversity and very high number of rare species. Out of the 745 species occurring with > five individuals, 80.4 % had an aggregated distribution, 19.5% were randomly distributed and one species had a regular distribution. The spatial patterns of rare vs. common species, juvenile vs. adult trees, and coarse vs. fine scales were compared. Rare species are generally less aggregated than common ones and most of the randomly distributed species are rare. Spatial patterns shift from high clumping to looser intensity or random distribution when moving from juveniles to adults for the same species. No adult tree species display a regular pattern, however. Regular distributions were rarely found; this is probably due to intraspecific competition at a local scale. There is a negative correlation between per capita death rate and population density. This study suggests that the Pasoh forest and its high diversity are subjected to multiple controlling factors, e.g., topography, spacing effect, density-dependent processes and species rarity. The importance of any factor changes across spatial and temporal scales.     

Spatiotemporal patterns and dynamics of species richness and abundance of woody plant functional groups in a tropical forest landscape of Hainan Island, South China

Tropical forests are among the most species-diverse ecosystems on Earth. Their structures and ecological functions are complex to understand. Functional group is defined as a group of species that play similar roles in an ecosystem. The functional group approach has been regarded as an effective way of linking the compositions of complex ecosystems with their ecological functions. To understand the variation of functional groups in species-rich ecosystems after disturbance, the present study investigated the spatial pattern and temporal dynamics of woody plants in a typically fragmented natural forest landscape of Hainan Island in South China. The study area was classified into eight landscape types based on vegetation type, disturbance manner and the time of recovery. The woody plant species were aggregated into seven functional groups based on the growth form, successional status and plant size. The results gained from the present study showed that all functional groups, except for the emergent and canopy tree species, were present in all eight landscape types. Each landscape type had different numbers of dominant functional groups. There are similar species richness and stem abundance structure among functional groups between mid-successional clear cut lowland rainforest and old growth tropical coniferous forest. This similarity exists in selective logged lowland rainforest and old-growth lowland rainforest, as well as among landscape types of montane rainforest. The functional groups with the same successional status had similar patterns of species richness and stem abundance ratios among different landscape types. The variation patterns of functional groups along the successional stages in terms of species richness and stem abundance among the tropical lowland rainforest landscape types were more similar to each other than those in the tropical montane reinforest landscape types. This study provides further support for the competition-colonization tradeoff and successional niche theory as opposed to models of neutrality and ecological equivalence.     

Effects of urbanization intensity on forest structural-taxonomic attributes,landscape patterns and their associations in Changchun,Northeast China: Implications for urban green infrastructure planning

Understanding the associations between urbanization intensity and urban forest structural-taxonomic attributes is a central theme of urban ecology, biodiversity conservation and forest management for maximizing ecological services to design proper urban green infrastructure. By selecting a typical provincial capital city of Changchun as an example, the effects of urbanization intensity (low, medium and heavy urbanization as measured by impervious surface area, ISA) on landscape patterns and structural-taxonomic attributes of urban forests were investigated in this study. The results showed that the urban forest Patch Density (PD), Landscape Shape Index (LSI), Interspersion & Juxtaposition Index (IJI), Tree Density (TD), Canopy Density (CD), Species Richness (SR) and Species Diversity (H’ index) exhibited strictly monotonic increases with urbanization intensity, increasing by 162%, 60%, 44%, 37%, 50%, 85%, and 84% from low to heavy urbanization areas, respectively. In contrast, the forest Mean Euclidian Nearest Neighbor Distance (ENN-MN) and Health Condition grade (HC) monotonically decreased by 12% and 37%, respectively. Furthermore, regression analysis suggested that structural-taxonomic attributes were closely associated with forest-related landscape patterns, but urbanization intensity dramatically influenced these associations. Our findings highlight that the planning of urban green infrastructure, in particular, urban afforestation and associated management, should be different at different urbanization intensities. In low urbanization areas, low Total area (TA, forest area) requires construction of larger forest patches and the protection of large remnant trees, and TD and above ground biomass (AGB) could be increased by enhancing the Mean Fractal Dimension (FRAC-MN) and PD, respectively. In medium urbanization areas, the same increases in TD and AGB could be more effectively achieved by decreasing the IJI and Area-Weighted Mean Contiguity (CONTIG-MN). Moreover, in heavy urbanization areas, more attention should be paid to increasing forest patch aggregation and contiguity, and both tree diversity and evenness could be increased by enhancing the FRAC-MN index. Because these structural-taxonomic attributes are the basis of various forest ecological services, our findings indicate that regulation of some of these landscape metrics could improve urban forest services in Changchun.     

Primary and secondary vegetation patches as contributors to floristic diversity in a tropical deciduous forest landscape

The floristic diversity of Mexican tropical deciduous forests (TDF) is of critical importance given the high species richness (alpha diversity), species turnover (beta diversity), and the intense deforestation rates. Currently, most TDF landscapes are mosaics of agricultural land, secondary vegetation, and patches of relatively undisturbed primary vegetation. Here we illustrate how both primary forest remnants and secondary vegetation patches contribute to the floristic diversity of TDF in a landscape of volcanic origin in central Veracruz, Mexico. Our objectives were to assess sampling efficiency and inventory completeness, to compare mean and cumulative species richness between primary forest and secondary vegetation sites, and to analyze beta diversity between vegetation types. In an area of 12,300 m2 we recorded 105 families, 390 genera, and 682 species. Species inventories for both vegetation types were about 80% complete. Secondary vegetation is more alpha diverse than primary forest, both in terms of cumulative and mean species richness. We found a remarkably high beta diversity between vegetation types (75% of complementarity, 91.60% of mean dissimilarity). We also identified the species that contribute the most to similarity within vegetation types and to dissimilarity between vegetation types. Our results support the idea that assessing biodiversity on the landscape scale is an appropriate way to ascertain the impact of human activities. For this land mosaic, conservation of the flora would not be possible by focusing solely on primary forest remnants. We propose the implementation of a network of small conservation areas with a flexible structure, following the “archipelago reserve” model.     

利用GIS和RS确定长白山自然保护区森林景观分布的环境范围

在对遥感数据进行景观分类和对环境因子进行空间表达基础上,在地理信息系统的支持下,确定长白山自然保护区森林景观分布的环境(包括年均温、年降水量、坡度和坡向)范围．结果表明。从苔原、岳桦、云冷杉到阔叶红松林,最适海拔高度范围依次为1780-2212、1705-1956、1042-1625、823-1184m;最适温度范围分别为-4．75～-2．40℃、-3．42～-2．07℃、-1．49-1．39℃、0．71-2．37℃;最适降水范围分别为1034～1110、1014-1060、883-1017、824-925mm;长白山自然保护区的森林景观主要分布在平、缓坡地上,并与坡向关系密切。苔原在各个坡向上均有分布。且在各个坡向上分布面积的变化不大;岳样、云冷杉林、阔叶红松林、山杨白样林主要呈现北、西北向分布,其次为东北、西向分布;落叶松林主要为东北向分布。其次为东和北向分布;疏林主要为西向分布,其次为西南、西北和南向分布;风倒区主要为西、西南、西北向分布。     

Relationships between spatial configuration of tropical forest patches and woody plant diversity in northeastern Puerto Rico

The destruction and fragmentation of tropical forests are major sources of global biodiversity loss. A better understanding of anthropogenically altered landscapes and their relationships with species diversity and composition is needed in order to protect biodiversity in these environments. The spatial patterns of a landscape may control the ecological processes that shape species diversity and composition. However, there is little information about how plant diversity varies with the spatial configuration of forest patches especially in fragmented tropical habitats. The northeastern part of Puerto Rico provides the opportunity to study the relationships between species richness and composition of woody plants (shrubs and trees) and spatial variables [i.e., patch area and shape, patch isolation, connectivity, and distance to the Luquillo Experimental Forest (LEF)] in tropical forest patches that have regenerated from pasturelands. The spatial data were obtained from aerial color photographs from year 2000. Each photo interpretation was digitized into a GIS package, and 12 forest patches (24–34 years old) were selected within a study area of 28 km². The woody plant species composition of the patches was determined by a systematic floristic survey. The species diversity (Shannon index) and species richness of woody plants correlated positively with the area and the shape of the forest patch. Larger patches, and patches with more habitat edge or convolution, provided conditions for a higher diversity of woody plants. Moreover, the distance of the forest patches to the LEF, which is a source of propagules, correlated negatively with species richness. Plant species composition was also related to patch size and shape and distance to the LEF. These results indicate that there is a link between landscape structure and species diversity and composition and that patches that have similar area, shape, and distance to the LEF provide similar conditions for the existence of a particular plant community. In addition, forest patches that were closer together had more similarity in woody plant species composition than patches that were farther apart, suggesting that seed dispersal for some species is limited at the scale of 10 km.     

Tree species richness complementarity, disturbance and fragmentation in a Mexican tropical montane cloud forest

Tropical montane cloud forest exhibits great heterogeneity in speciescomposition and structure over short geographic distances. In central Veracruz,Mexico, the conservation priority of seven cloud forest fragments was assessedby considering differences in woody plant species richness and complementarityof species among sites, forest structure, tree mortality, and timber andfirewood extraction as indicators of anthropogenic disturbance. Densities oftrees 5 cm dbh (360–1700 trees/ha) weredifferent among the sites, but basal area (35.3–89.3m²/ha) did not differ among fragments. The number of dead trees rangedfrom 10–30 to 170–200 trees/ha. The fragments'species composition was different but complementary. The Morisita–Hornindex indicated low similarity between fragments. A non-parametric estimator ofspecies richness indicated that more sampling effort would be necessary tocomplete the inventory (15 additional trees and two understory shrub species).Unfortunately, most of the fragments are threatened with deforestation. The numberof cut trees was similar among sites (0–15 cut trees/0.1 ha).Sites with immediate need for conservation were close to settlements, with highnumbers of cut trees and no legal protection. The selected sites represent thevariety of situations that exist in the region. Given the high complementarityobserved between fragments, a regional conservation approach will be required topreserve the last repositories of part of the tremendous biodiversity of theonce continuous forest in this region.     

Effects of fire on landscape heterogeneity in Yellowstone National Park,Wyoming

Abstract. A map of burn severity resulting from the 1988 fires that occurred in Yellowstone National Park (YNP) was derived from Landsat Thematic Mapper (TM) imagery and used to assess the isolation of burned areas, the heterogeneity that resulted from fires burning under moderate and severe burning conditions, and the relationship between heterogeneity and fire size. The majority of severely burned areas were within close proximity (50 to 200 m) to unburned or lightly burned areas, suggesting that few burned sites are very far from potential sources of propagules for plant reestablishment. Fires that occurred under moderate burning conditions early during the 1988 fire season resulted in a lower proportion of crown fire than fires that occurred under severe burning conditions later in the season. Increased dominance and contagion of burn severity classes and a decrease in the edge: area ratio for later fires indicated a slightly more aggregated burn pattern compared to early fires. The proportion of burned area in different burn severity classes varied as a function of daily fire size. When daily area burned was relatively low, the proportion of burned area in each burn severity class varied widely. When daily burned area exceeded 1250 ha, the burned area contained about 50 % crown fire, 30 % severe surface burn, and 20 % light surface burn. Understanding the effect of fire on landscape heterogeneity is important because the kinds, amounts, and spatial distribution of burned and unburned areas may influence the reestablishment of plant species on burned sites.     

Fern richness,tree species surrogacy,and fragment complementarity in a Mexican tropical montane cloud forest

We related pteridophytes versus tree species composition to identify surrogate measures of diversity, and complementarity of seven cloud forest fragments. Forest structure, and fern and tree composition were determined in 70 (2 × 50 m) transects. Fern density (10,150–25,080 individuals/ha) differed among sites. We recorded 83 fern species in the transects. Nonparametric richness estimators indicated that more sampling effort was needed to complete fern inventories (14 more species). However, ferns recorded outside of the transects increased richness to 103 species (six more species than predicted). Twenty-eight species were unique and rare due to special habitat requirements (Diplazium expansum, Hymenophyllum hirsutum, Melpomene leptostoma, Terpsichore asplenifolia), or were at a geographical distribution edge (Diplazium plantaginifolium, Lycopodium thyoides, Pecluma consimilis, Polypodium puberulum). Correlations between fern richness and tree richness and density were not significant, but were significant between fern richness and fern density, between epiphytic fern density and tree richness and density. Tree richness is not a good surrogate for fern diversity. Only three species were recorded in all fragments (Polypodium lepidotrichum, P. longepinnulatum, P. plebeium); thus fragments pteridophytes compositions are highly complementary, but more similar for ferns than for trees. A regional conservation approach which includes many small reserves needs to focus supplementarity on patterns of tree and fern species richness.     

Nutrient enrichment increased species richness of leaf litter fungal assemblages in a tropical forest

Microbial communities play a major role in terrestrial ecosystem functioning, but the determinates of their diversity and functional interactions are not well known. In this study, we explored leaf litter fungal diversity in a diverse Panama lowland tropical forest in which a replicated factorial N, P, K and micronutrient fertilization experiment of 40 × 40 m plots had been ongoing for nine years. We extracted DNA from leaf litter samples and used fungal‐specific amplification and a 454 pyrosequencing approach to sequence two loci, the nuclear ribosomal internal transcribed spacer (ITS) region and the nuclear ribosomal large subunit (LSU) D1 region. Using a 95% sequence similarity threshold for ITS1 spacer recovered a total of 2523 OTUs, and the number of unique ITS1 OTUs per 0.5–1.0 g leaf litter sample ranged from 55 to 177. Ascomycota were the dominant phylum among the leaf litter fungi (71% of the OTUs), followed by Basidiomycota (26% of the OTUs). In contrast to our expectations based on temperate ecosystems, long‐term addition of nutrients increased, rather than decreased, species richness relative to controls. Effect of individual nutrients was more subtle and seen primarily as changes in community compositions especially at lower taxonomic levels, rather than as significant changes in species richness. For example, plots receiving P tended to show a greater similarity in community composition compared to the other nutrient treatments, the +PK, +NK and +NPK plots appeared to be more dominated by the Nectriaceae than other treatments, and indicator species for particular nutrient combinations were identified.     

Determinants of fine‐scale plant species richness in a deciduous forest of northeastern North America

Question: What are the determinants of fine‐scale plant species richness (SR)? Location: Île‐aux‐Grues, Québec, Canada. Methods: Elevation, soil organic matter, soil pH, irradiance, tree basal area (BA) and plant SR (herbs, shrubs, and trees) were determined in 100 contiguous 25 m² quadrats in a deciduous forest. Each variable was analysed for spatial autocorrelation using Moran's I. Path analysis was used to determine the effects of different variables on tree, shrub and herb SR in a hierarchical modelling approach. Results: Most of the variables, except tree BA, PPFD (photo‐synthetic photon flux density) and shrub SR, were positively autocorrelated at a scale of ca. 20 m or less. The path analyses explained ca. ll%‐40% of the variance in plant SR; however, the model for shrub SR was not significant. Tree SR was positively associated, but herb SR was negatively associated with tree BA. Tree SR had a positive influence on shrub SR, but herb SR remained unaffected by tree or shrub SR. Conclusion: The positive association of tree BA and tree SR suggests that the data from the study site correspond to the left (ascending) portion of the SR‐biomass relationship (un‐dersaturated SR). The negative effect of tree BA on herb SR is direct and not mediated through reduced PPFD. High tree BA might cause high resource use, induce high litter production and affect soil properties, all of which might significantly affect herb SR. Several factors not considered here might influence fine‐scale SR, such as interspecific interactions, fine‐scale disturbances and heterogeneity (both spatial and temporal) in resources and abiotic conditions. Within‐site variations of SR might be difficult to model with precision because of the relative importance of stochastic vs deterministic processes at this spatial scale.     

西双版纳热带雨林林窗空间分布格局及其特征数与林窗下植物多样性的相关性

林窗作为森林群落中一种重要的干扰方式, 对林下物种构成有着重要的影响。开展林窗空间格局及其特征指数与林下植物多样性关系研究对于探讨林窗对林下生物多样性的影响有重要意义, 有助于进一步了解群落动态, 在物种多样性保护方面也具有指导作用。本研究在西双版纳热带雨林地区随机选取3块大小为1 ha的热带雨林为研究样地, 采用轻小型六旋翼无人机搭载Sony ILCE-A7r可见光传感器, 分别获取各个样地的高清数字影像, 结合数字表面高程模型以及各个样地的地形数据用以确定各样区的林窗分布格局, 并进一步提取出各林窗的景观格局指数。结合地面样方基础调查数据, 对各样地各林窗下植物多样性情况进行统计, 旨在分析热带雨林林窗空间分布格局以及林窗下植物多样性对各林窗空间格局特征的响应情况。研究表明, 西双版纳州热带雨林林窗呈大而分散的空间分布, 林窗空间格局特征指数如林窗形状复杂性指数、林窗面积都与林下植物多样性呈显著正相关关系。在面积小的林窗下, 较之林窗形状复杂性因子, 林窗面积大小对林下植物多样性影响更显著; 在面积达到一定程度后, 相对于面积因子, 林窗形状复杂性指数对林下植物多样性影响更显著, 各样地林窗皆趋于向各自所处样地顶极群落发展。     

Tree species composition and diversity of tropical mountain cloud forest in the Yunnan, southwestern China

Species composition, physiognomy, and plant diversity of the less known cloud forests in Yunnan were studied based on data collected from 35 sample plots at seven sites. In floristic composition, the cloud forests are mainly comprised of Fagaceae, Ericaceae, Vacciniaceae, Aceraceae, Magnoliaceae, Theaceae, Aquifoliaceae, Illiciaceae, Lauraceae, and Rosaceae. Physiognomically, the forests are dominated by tree and shrub species. Lianas are rare in the forests. The plants with microphyllous or nanophyllous leaves comprise 44.32–63.46% of the total species, and plants with an entire leaf margin account for more than 50% of the tree and shrub species. There are few tree and shrub species with a drip tip leaf apex and papery leaves. Evergreen species make up more than 75% of the total tree and shrub species. In a 2,500 m² sampling area, the number of vascular species ranged between 57 and 110; Simpson’s diversity index ranged from 0.7719 to 0.9544, Shannon–Winner’s diversity index from 1.8251 to 3.2905, and Pielou’s evenness index from 0.5836 to 0.8982 for trees. The cloud forests in Yunnan are physiognomically similar to the tropical cloud forests in America and Southeast Asia. They very much resemble the mountain dwarf mossy forest in Hainan Island, southeastern China, and the Mountain ericaceous forests in the Malay Peninsula. The cloud forests in Yunnan are considered to be developed, as are the tropical upper montane cloud forests in Asia.     

Growth strategies differentiate the spatial patterns of 11 dipterocarp species coexisting in a Malaysian tropical rain forest

We examined relationships between mortality rate, relative growth rate (RGR), and spatial patterns of three growth stages (small, medium, and large trees) for 11 dipterocarp species in the Pasoh 50-ha plot. Mortality rates for these species tended to be positively correlated with RGRs, although the correlation was significant only at the small-tree stage. Seven species with high growth and mortality rates exhibited peaks in spatial aggregation at small distances (<100 m) in small trees, but this aggregation disappeared in medium and large trees. In contrast, the other four species with low growth and mortality rates aggregated at large distances (>200 m) throughout the three growth stages in all but one species. Negative associations between different growth stages were observed only for the high-mortality species, suggesting density-dependent mortality. The high-mortality species showed habitat associations with topography, soil type, and the forest regeneration phase after gap formation, whereas the three low-mortality species only had associations with the forest regeneration phase. A randomization procedure revealed that these habitat associations explained little of their spatial aggregation. Our results suggest that the growth strategy has a large effect on the structuring of the spatial distribution of tree species through mortality processes.