A negative heterogeneity–diversity relationship found in experimental grassland communities

Recent meta-analyses and simulation studies have suggested that the relationship between soil resource heterogeneity and plant diversity (heterogeneity–diversity relationship; HDR) may be negative when heterogeneity occurs at small spatial scales. To explore different mechanisms that can explain a negative HDR, we conducted a mesocosm experiment combining a gradient of soil nutrient availability (low, medium, high) and scale of heterogeneity (homogeneous, large-scale heterogeneous, small-scale heterogeneous). The two heterogeneous treatments were created using chessboard combinations of low and high fertility patches, and had the same overall fertility as the homogeneous medium treatment. Soil patches were designed to be relatively larger (156 cm²) and smaller (39 cm²) than plant root extent. We found plant diversity was significantly lower in the small-scale heterogeneous treatment compared to the homogeneous treatment of the same fertility. Additionally, low fertility patches in the small-scale heterogeneous treatment had lower diversity than patches of the same size in the low fertility treatment. Shoot and root biomass were larger in the small-scale heterogeneous treatment than in the homogeneous treatment of the same fertility. Further, we found that soil resource heterogeneity may reduce diversity indirectly by increasing shoot biomass, thereby enhancing asymmetric competition for light resources. When soil resource heterogeneity occurs at small spatial scales it can lower plant diversity by increasing asymmetric competition belowground, since plants with large root systems can forage among patches and exploit soil resources. Additionally, small-scale soil heterogeneity may lower diversity indirectly, through increasing light competition, when nutrient uptake by competitive species increases shoot biomass production.     

Plant species coexistence at local scale in temperate swamp forest: test of habitat heterogeneity hypothesis

It has been suggested that a heterogeneous environment enhances species richness and allows for the coexistence of species. However, there is increasing evidence that environmental heterogeneity can have no effect or even a negative effect on plant species richness and plant coexistence at a local scale. We examined whether plant species richness increases with local heterogeneity in the water table depth, microtopography, pH and light availability in a swamp forest community at three local spatial scales (grain: 0.6, 1.2 and 11.4 m). We also used the variance partitioning approach to assess the relative contributions of niche-based and other spatial processes to species occurrence. We found that heterogeneity in microtopography and light availability positively correlated with species richness, in accordance with the habitat heterogeneity hypothesis. However, we recorded different heterogeneity-diversity relationships for particular functional species groups. An increase in the richness of bryophytes and woody plant species was generally related to habitat heterogeneity at all measured spatial scales, whereas a low impact on herbaceous species richness was recorded only at the 11.4 m scale. The distribution of herbaceous plants was primarily explained by other spatial processes, such as dispersal, in contrast to the occurrence of bryophytes, which was better explained by environmental factors. Our results suggest that both niche-based and other spatial processes are important determinants of the plant composition and species turnover at local spatial scales in swamp forests.     

Joint Effects of Habitat Heterogeneity and Species’ Life-History Traits on Population Dynamics in Spatially Structured Landscapes

Both habitat heterogeneity and species’ life-history traits play important roles in driving population dynamics, yet there is little scientific consensus around the combined effect of these two factors on populations in complex landscapes. Using a spatially explicit agent-based model, we explored how interactions between habitat spatial structure (defined here as the scale of spatial autocorrelation in habitat quality) and species life-history strategies (defined here by species environmental tolerance and movement capacity) affect population dynamics in spatially heterogeneous landscapes. We compared the responses of four hypothetical species with different life-history traits to four landscape scenarios differing in the scale of spatial autocorrelation in habitat quality. The results showed that the population size of all hypothetical species exhibited a substantial increase as the scale of spatial autocorrelation in habitat quality increased, yet the pattern of population increase was shaped by species’ movement capacity. The increasing scale of spatial autocorrelation in habitat quality promoted the resource share of individuals, but had little effect on the mean mortality rate of individuals. Species’ movement capacity also determined the proportion of individuals in high-quality cells as well as the proportion of individuals experiencing competition in response to increased spatial autocorrelation in habitat quality. Positive correlations between the resource share of individuals and the proportion of individuals experiencing competition indicate that large-scale spatial autocorrelation in habitat quality may mask the density-dependent effect on populations through increasing the resource share of individuals, especially for species with low mobility. These findings suggest that low-mobility species may be more sensitive to habitat spatial heterogeneity in spatially structured landscapes. In addition, localized movement in combination with spatial autocorrelation may increase the population size, despite increased density effects.     

Point process models,the dimensions of biodiversity and the importance of small-scale biotic interactions

Aims Recent mechanistic explanations for community assembly focus on the debates surrounding niche-based deterministic and dispersal-based stochastic models. This body of work has emphasized the importance of both habitat filtering and dispersal limitation, and many of these works have utilized the assumption of species spatial independence to simplify the complexity of the spatial modeling in natural communities when given dispersal limitation and/or habitat filtering. One potential drawback of this simplification is that it does not consider species interactions and how they may influence the spatial distribution of species, phylogenetic and functional diversity. Here, we assess the validity of the assumption of species spatial independence using data from a subtropical forest plot in southeastern China.Methods We use the four most commonly employed spatial statistical models—the homogeneous Poisson process representing pure random effect, the heterogeneous Poisson process for the effect of habitat heterogeneity, the homogenous Thomas process for sole dispersal limitation and the heterogeneous Thomas process for joint effect of habitat heterogeneity and dispersal limitation—to investigate the contribution of different mechanisms in shaping the species, phylogenetic and functional structures of communities.Important findings Our evidence from species, phylogenetic and functional diversity demonstrates that the habitat filtering and/or dispersal-based models perform well and the assumption of species spatial independence is relatively valid at larger scales (50×50 m). Conversely, at local scales (10×10 and 20×20 m), the models often fail to predict the species, phylogenetic and functional diversity, suggesting that the assumption of species spatial independence is invalid and that biotic interactions are increasingly important at these spatial scales.     

海南岛霸王岭热带低地雨林树木的空间格局

树木空间格局及其形成过程是物种共存及生物多样性维持机制研究的一个重要方面。该文以海南岛两个1 hm ²的典型热带低地雨林老龄林森林动态样地为基础, 通过4个点格局模型(均质Poisson过程、异质Poisson过程、均质Thomas过程和异质Thomas过程)模拟扩散限制和生境异质性作用对树木空间分布格局的影响, 并分析不同空间尺度下(< 2 m, 2-5 m, 5-10 m, 10-15 m, 15-20 m和20-25 m)不同作用的相对重要性。结果表明: 热带低地雨林的所有树木总体上呈现聚集分布的空间格局, 随着尺度的增大, 聚集强度逐渐减小。树种在模拟空间分布格局最优模型中的比例由高到低分别是: 均质Thomas过程, 均质Poisson过程、异质Thomas过程和异质Poisson过程。扩散限制作用是形成热带低地雨林树木空间分布格局最重要的生态过程, 其次是完全随机作用以及生境异质性和扩散限制的联合作用, 而生境异质性的作用最小。不同空间尺度上模拟各树种空间分布格局的最优模型比例差异显著, 扩散限制作用能够在多数空间尺度上模拟多个树种的空间分布格局, 其次为随机作用; 生境异质性和扩散限制的联合作用主要在小尺度(0-5 m)影响树种分布, 而生境异质性在较大尺度(15-25 m)上影响树种的空间分布格局。     

Neighborhood models of plant population dynamics 3. Models with spatial heterogeneity in the physical environment

Population dynamic models are developed for communities of annual plants in spatially heterogeneous environments. These models are constructed from submodels of the survivorship, fecundity, germination, and dispersal of individual plants. The submodels include the effects of spatially local interactions on plant performance and the spatial variation in performance caused by spatial heterogeneity in the physical environment. It is possible to estimate the submodels from data on experimental communities in either the field or greenhouse and so it is possible to empirically calibrate the population dynamic models developed. Each population dynamic model explicitly includes the spatial distribution of individuals in a plant community.Several two-species models for plants in patchy environments are studied to examine the community-level consequences of spatial heterogeneity in the physical environment. The results fall into two classes. First, community structure is in part determined by a relation between patch size and mean seed dispersal distance. Specifically, coexistence is, in some cases, possible only if patches are sufficiently larger than the mean dispersal distance. Second, community structure is also affected by relations between patch size and the maximum distance over which two plants interact (termed the neighborhood radius). In some cases, coexistence is possible only if patch size is sufficiently larger than the neighborhood radius. In others, the species coexist only if patch size is sufficiently smaller than the neighborhood radius. In still other cases, coexistence is possible only if patch sizes are within critical bounds, where the sizes of the critical bounds are in units of the neighborhood radius. All results involving relations between the neighborhood radius and patch size are direct consequences of the sedentary nature of plants and the fact that individual plants interact primarily with nearby plants.     

The scale-dependent importance of habitat factors and dispersal limitation in structuring Great Lakes shoreline plant communities

Niche-based and neutral models of community structure posit distinct mechanisms underlying patterns in community structure; correlation between species’ distributions and habitat factors points to niche assembly while spatial pattern independent of habitat suggests neutral assembly via dispersal limitation. The challenge is to disentangle the relative contributions when both processes are operating, and to determine the scales at which each is important. We sampled shoreline plant communities on an island in Lake Michigan, varying the extent and the grain of sampling, and used both distance-based correlation methods and variance partitioning to quantify the proportion of the variation in plant species composition that was attributable to habitat factors and to spatial configuration independent of habitat. Our results were highly scale dependent. We found no distance decay of plant community similarity at the island scale (1−33 km). All of the explained variation (32%) in species composition among samples at this scale was attributed to habitat factors. However, at a site intensively sampled at a smaller scale (5−1,200 m), similarity of species composition did decay with distance. Using a coarse sampling grain (transects), habitat factors explained 40% of the variation, but the purely spatial component explained a comparable 22%. Analyzing plots within transects revealed variation in species composition that was still jointly determined by habitat and spatial factors (18 and 11% of the variance, respectively). For both grain sizes, most of the habitat component was spatially structured, reflecting an abrupt alongshore transition from sandy dunes to cobble beach. Space per se explained more variation in species composition at a second site where the habitat transition was more gradual; here, habitat acted as a less selective filter, allowing the signal of dispersal limitation to be detected more readily. We conclude that both adaptation to specific habitat factors and habitat-independent spatial position indicative of dispersal limitation determine plant species composition in this system. Our results support the prediction that dispersal limitation—a potentially, but not necessarily, neutral driver—is relatively more important at smaller scales.     

An allometric model of home range formation explains the structuring of animal communities exploiting heterogeneous resources

Understanding and predicting the composition and spatial structure of communities is a central challenge in ecology. An important structural property of animal communities is the distribution of individual home ranges. Home range formation is controlled by resource heterogeneity, the physiology and behaviour of individual animals, and their intra‐ and interspecific interactions. However, a quantitative mechanistic understanding of how home range formation influences community composition is still lacking. To explore the link between home range formation and community composition in heterogeneous landscapes we combine allometric relationships for physiological properties with an algorithm that selects optimal home ranges given locomotion costs, resource depletion and competition in a spatially‐explicit individual‐based modelling framework. From a spatial distribution of resources and an input distribution of animal body mass, our model predicts the size and location of individual home ranges as well as the individual size distribution (ISD) in an animal community. For a broad range of body mass input distributions, including empirical body mass distributions of North American and Australian mammals, our model predictions agree with independent data on the body mass scaling of home range size and individual abundance in terrestrial mammals. Model predictions are also robust against variation in habitat productivity and landscape heterogeneity. The combination of allometric relationships for locomotion costs and resource needs with resource competition in an optimal foraging framework enables us to scale from individual properties to the structure of animal communities in heterogeneous landscapes. The proposed spatially‐explicit modelling concept not only allows for detailed investigation of landscape effects on animal communities, but also provides novel insights into the mechanisms by which resource competition in space shapes animal communities.     

植物群落中物种小尺度空间结构研究

植物群落中, 物种小尺度空间结构影响着种群或群落的动态及有关的生态学过程。植物主要是和它同种或异种的邻近个体相互作用,植物个体周围的局部环境和大空间尺度下群落的平均水平是完全不同的。群落中的许多过程都影响小尺度空间结构的形成和动态,同样,局部空间结构反过来影响着植物的生长、更新和凋亡等重要过程。鉴于目前对小尺度空间结构进行的大量研究以及其重要性,有必要对其研究进展进行适当的总结,以期明确将来进一步的研究方向。该文以此为出发点,首先介绍了植物群落中物种小尺度空间结构产生的6个原因:1)生境的空间异质性;2)植物繁殖体的传播;3)植物之间的相互作用;4)生物环境(动物和微生物)的作用;5)外界干扰的作用;6)多因子综合作用。然后阐述了小尺度空间结构意义及对生物多样性、植物种群遗传学和恢复生态学研究的影响。最后对目前物种小尺度空间结构研究存在的几点问题及将来的研究方向作以下归纳:1)大尺度植被动态的研究应该整合小尺度空间结构的信息;2)不论从生物学还是生态学上来讲, 植被小尺度空间结构的研究应该把植物作为中心,确定适当的尺度和采取合理的空间统计方法;3)充分重视小尺度空间结构在退化生态系统恢复中的应用意义;4)注重从小尺度的局部格局研究入手对群落总体特征进行整合;5)植物群落动态研究中,物种小尺度空间结构与平均场假说相结合的必要性。     

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。     

Influence of grazing and fire frequency on small-scale plant community structure and resource variability in native tallgrass prairie

In grasslands worldwide, grazing by ungulates and periodic fires are important forces affecting resource availability and plant community structure. It is not clear, however, whether changes in community structure are the direct effects of the disturbance (i.e. fire and grazing) or are mediated indirectly through changes in resource abundance and availability. In North American tallgrass prairies, fire and grazing often have disparate effects on plant resources and plant diversity, yet, little is known about the individual and interactive effects of fire and grazing on resource variability and how that variability relates to heterogeneity in plant community structure, particularly at small scales. We conducted a field study to determine the interactive effects of different long-term fire regimes (annual vs four-year fire frequency) and grazing by native ungulates ( Bos bison ) on small-scale plant community structure and resource variability (N and light) in native tallgrass prairie. Grazing enhanced light and nitrogen availability, but did not affect small-scale resource variability. In addition, grazing reduced the dominance of C₄ grasses which enhanced species richness, diversity and community heterogeneity. In contrast, annual fire increased community dominance and reduced species richness and diversity, particularly in the absence of grazing, but had no effect on community heterogeneity, resource availability and resource variability. Variability in the abundance of resources showed no relationship with community heterogeneity at the scale measured in this study, however we found a relationship between community dominance and heterogeneity. Therefore, we conclude that grazing generated small-scale community heterogeneity in this mesic grassland by directly affecting plant community dominance, rather than indirectly through changes in resource variability.     

Local and Regional Effects on Community Structure of Dung Beetles in a Mainland-Island Scenario

Understanding the ecological mechanisms driving beta diversity is a major goal of community ecology. Metacommunity theory brings new ways of thinking about the structure of local communities, including processes occurring at different spatial scales. In addition to new theories, new methods have been developed which allow the partitioning of individual and shared contributions of environmental and spatial effects, as well as identification of species and sites that have importance in the generation of beta diversity along ecological gradients. We analyzed the spatial distribution of dung beetle communities in areas of Atlantic Forest in a mainland-island scenario in southern Brazil, with the objective of identifying the mechanisms driving composition, abundance and biomass at three spatial scales (mainland-island, areas and sites). We sampled 20 sites across four large areas, two on the mainland and two on the island. The distribution of our sampling sites was hierarchical and areas are isolated. We used standardized protocols to assess environmental heterogeneity and sample dung beetles. We used spatial eigenfunctions analysis to generate the spatial patterns of sampling points. Environmental heterogeneity showed strong variation among sites and a mild increase with increasing spatial scale. The analysis of diversity partitioning showed an increase in beta diversity with increasing spatial scale. Variation partitioning based on environmental and spatial variables suggests that environmental heterogeneity is the most important driver of beta diversity at the local scale. The spatial effects were significant only at larger spatial scales. Our study presents a case where environmental heterogeneity seems to be the main factor structuring communities at smaller scales, while spatial effects are more important at larger scales. The increase in beta diversity that occurs at larger scales seems to be the result of limitation in species dispersal ability due to habitat fragmentation and the presence of geographical barriers.     

How the spatial scales of dispersal, competition, and environmental heterogeneity interact to affect coexistence

Spatial coexistence depends on a variety of biological and physical processes, and the relative scales of these processes may promote or suppress coexistence. We model plant competition in a spatially varying environment to show how shifting scales of dispersal, competition, and environmental heterogeneity affect coexistence. Spatial coexistence mechanisms are partitioned into three types: the storage effect, nonlinear competitive variance, and growth-density covariance. We first describe how the strength of each of these mechanisms depends on covariances between population densities and between population densities and the environment, and we then explain how changes in the scales of dispersal, competition, and environmental heterogeneity should affect these covariances. Our quantitative approach allows us to show how changes in the scales of biological and physical processes can shift the relative importance of different classes of spatial coexistence mechanisms and gives us a more complete understanding of how environmental heterogeneity can enable coexistence. For example, we show how environmental heterogeneity can promote coexistence even when competing species have identical responses to the environment.     

Scale-dependent interactions and community structure on cobble beaches

Recent theory suggests that scale-dependent interaction between facilitation and competition can generate spatial structure in ecological communities. The application of this hypothesis, however, has been limited to systems with little underlying heterogeneity. We evaluated this prediction in a plant community along an intertidal stress gradient on cobble beaches in Rhode Island, USA. Prior studies have shown that Spartina alterniflora facilitates a forb-dominated community higher in the intertidal by modifying the shoreline environment. We tested the hypothesis that, at a smaller scale, Spartina competitively excludes forb species, explaining their marked absence within the lower Spartina zone. Transplant experiments showed forb species grow significantly better in the Spartina zone when neighbours were removed. Removal of the Spartina canopy led to a massive emergence of annual forbs, showing that competition limits local occupation. These findings indicate that interaction of large-scale facilitation and small-scale competition drives plant zonation on cobble beaches. This study is the first to provide empirical evidence of scale-dependent interactions between facilitation and competition spatially structuring communities in heterogeneous environments.     

辽东山区次生林木本植物空间分布

森林木本植物的空间格局有助于揭示群落结构的形成机制与潜在的生态学过程,且对林分经营具有一定指导意义。在0—50 m尺度范围内综合分析了辽东山区4 hm2温带次生林样地多度10的树种空间格局。研究发现:(1)在完全随机零模型下,大部分树种呈现聚集格局,聚集格局树种的比例随尺度增加而降低;在32 m的较大尺度下,随尺度增加,随机和规则格局成为树种分布的主要形式;(2)在异质性泊松过程零模型下,55.9%的树种呈现随机格局,其余大部分树种在10 m的尺度下呈现聚集格局,且随尺度增加,规则格局成为主要形式;(3)在完全随机零模型下,树种属性(林层、径级和多度)显著地影响种群聚集度,而在异质性泊松过程零模型下,树种属性对种群聚集度不存在显著影响。综上,生境异质性、扩散限制和树种属性部分解释了辽东山区次生林木本植物空间分布格局,相对而言,生境异质性的效应更为突出。研究结果有助于揭示次生林群落生物多样性的维持机制。     

Abiotic and biotic resistance to grass invasion in serpentine annual plant communities

Biological invasions severely impact native plant communities, causing dramatic shifts in species composition and the restriction of native species to spatially isolated refuges. Competition from resident species and the interaction between resource limitation and competition have been overlooked as mechanisms of community resistance in refugia habitats. We examined the importance of these factors in determining the resistance of California serpentine plant communities to invasion by three common European grasses, Avena barbata, Bromus diandrus, and Hordeum murinum. We added seeds of each of these grasses to plots subjected to six levels of resource addition (N, P, Ca, H₂O, all resources together, and a no-addition control) and two levels of competition (with resident community present or removed). Resource limitation and competition had strong effects on the biomass and reproduction of the three invaders. The addition of all resources together combined with the removal of the resident community yielded individual plants that were fourfold to 20-fold larger and sixfold to 20-fold more fecund than plants from control plots. Competitor removal alone yielded invaders that were twofold to sevenfold larger and twofold to ninefold more fecund. N addition alone or in combination with other resources led to a twofold to ninefold increase in the biomass and fecundity of the invaders. No other resource alone significantly affected native or invader performance, suggesting that N was the key limiting resource during our experiment. We found a significant interaction between abiotic and biotic resistance for Bromus, which experienced increased competitive suppression in fertilized plots. The threefold increase in resident biomass with N addition was likely responsible for this result. Our results confirm that serpentine plant communities are severely N limited, which, in combination with competition from resident species, promotes the resistance of these systems to invasions. Our work suggests that better understanding the relative sensitivities of invaders and residents to the physical environment is critical to predicting how abiotic and biotic factors interact to determine community resistance.     

Spatial scaling and transition in pneumatophore arthropod communities

Although most ecological variables are scale-dependent, few studies cover a broad range of spatial scales. Here, we consider South African mangrove pneumatophore arthropod communities (mites, crustaceans and insects), across seven spatial scales (from 10  cm to 100  km). We plot spatial autocorrelation in individual species, evaluate if resource and habitat availability determine spatial patterning, and identify the scales of community transition. Spatial autocorrelation in most ecological variables decreased with increasing spatial scale, with notable exceptions for the larger scales. Negative abundance autocorrelation was stronger at 10  km than at 100  km for common species, while the opposite was true for rare species. Spatial autocorrelation in species richness decreased from 1  m (strong positive) to 10  km (strong negative), but was not significant at the 100  km scale. These patterns reflect the patchy distribution of pneumatophores within mangrove forests, that of the forests along the coast, and the poor dispersal abilities of most of the arthropods sampled, in a highly dynamic environment. Although resource and habitat availability exhibited a similar autocorrelation pattern to that of the community, the total mass of pneumatophores did not appear to be an important determinant of community structure. Variations in the abundance of common species, as well as the restricted distribution of rare species caused assemblage structure to change gradually with increasing distance from 10 cm to 100 km, but only marginally from 10 to 100  km. We highlight the need for cross-scale studies in bridging the gap between two key ecological concepts: potential ecological niche and realized geographic range.     

Protozoan community structure in a fractal soil environment.

Protozoan abundance was quantified, and 365 protozoan species were recorded, in 150 soil samples from an upland grassland in Scotland. Across the entire size range (2-200 pm) protozoan species richness varied by a factor of two, whereas abundance increased by a factor of 20 with decreasing body size. As the soil had fractal structure, the relatively flat species curve can be explained if spatial heterogeneity determines species number--for in a fractal environment, heterogeneity will be the same at all spatial scales. Community structure appeared to approach a temporary steady-state about six days after re-hydration of dried soil. A simple model based on combining the fractal character of increasing habitat area at smaller spatial scales, with the weight-specific energy requirements of protozoa, provided theoretical curves of abundance and biovolume on body size which provide a reasonable fit to real data. We suggest two possibilities--that the apparent competence of the theoretical model is fortuitous and the product of poorly understood dynamic elements of the trophic structure in the community; or that key elements of protozoan community structure in a fractal soil environment may be largely explained in terms of habitat space and energy requirements.     

Mechanisms of ecological succession: insights from plant functional strategies

Successions are a central issue of ecological theory. They are governed by changes in community assembly processes that can be tracked by species’ traits. While single‐trait‐based approaches have been mostly promoted to address community assembly, ecological strategies actually encompass tradeoffs between multiple traits that are relevant to succession theory. We analyzed plant ecological strategies along a 140‐year‐long succession primary succession of 52 vertical outcrop communities after roadwork. We performed a RLQ analysis to relate six functional traits, associated with resource acquisition, competition, colonization ability and phenology, to the age of the outcrops. We found the prominence of two main axes of specialization, one related to resource acquisition and the other to reproduction and regeneration. We further examined the community‐level variation in ecological strategies to assess the abiotic and biotic drivers of community assembly. Using trait‐based statistics of functional richness, regularity and divergence, we found that different processes drove the variation in ecological strategies along the axes of specialization. In late succession, functional convergence was detected for the traits related to resource acquisition as a signature of habitat filtering, while the coexistence of contrasted strategies was found for the traits related to reproduction and regeneration as a result of spatial micro‐heterogeneity. We observed a lack of niche differentiation along the succession, revealing a weak importance of biotic interactions for the regulation of community assembly in the outcrops. Overall, we highlight a prominent role of habitat filtering and spatial micro‐heterogeneity in driving the primary succession governed by water and nutrient limitation.     

The effect of static and dynamic spatially structured disturbances on a locally dispersing population

Previous models of locally dispersing populations have shown that in the presence of spatially structured fixed habitat heterogeneity, increasing local spatial autocorrelation in habitat generally has a beneficial effect on such populations, increasing equilibrium population density. It has also been shown that with large-scale disturbance events which simultaneously affect contiguous blocks of sites, increasing spatial autocorrelation in the disturbances has a harmful effect, decreasing equilibrium population density. Here, spatial population models are developed which include both of these spatially structured exogenous influences, to determine how they interact with each other and with the endogenously generated spatial structure produced by the population dynamics. The models show that when habitat is fragmented and disturbance occurs at large spatial scales, the population cannot persist no matter how large its birth rate, an effect not seen in previous simpler models of this type. The behavior of the model is also explored when the local autocorrelation of habitat heterogeneity and disturbance events are equal, i.e. the two effects occur at the same spatial scale. When this scale parameter is very small, habitat fragmentation prevents the population from persisting because sites attempting to reproduce will drop most of their offspring on unsuitable sites; when the parameter is very large, large-scale disturbance events drive the population to extinction. Population levels reach their maximum at intermediate values of the scale parameter, and the critical values in the model show that the population will persist most easily at these intermediate scales of spatial influences. The models are investigated via spatially explicit stochastic simulations, traditional (infinite-dispersal) and improved (local-dispersal) mean-field approximations, and pair approximations.