Showing papers by "Thorsten Wiegand published in 2015"

Adopting a spatially explicit perspective to study the mysterious fairy circles of Namibia

Abstract: The mysterious ‘fairy circles’ are vegetation-free discs that cover vast areas along the pro-Namib Desert. Despite 30 yr of research their origin remains unknown. Here we adopt a novel approach that focuses on analysis of the spatial patterns of fairy circles obtained from representative 25-ha aerial images of north-west Namibia. We use spatial point pattern analysis to quantify different features of their spatial structures and then critically inspect existing hypotheses with respect to their ability to generate the observed circle patterns. Our working hypothesis is that fairy circles are a self-organized vegetation pattern. Finally, we test if an existing partial-differential-equation model, that was designed to describe vegetation pattern formation, is able to reproduce the characteristic features of the observed fairy circle patterns. The model is based on key-processes in arid areas such as plant competition for water and local resource-biomass feedbacks. The fairy circles showed at all three study areas the same regular spatial distribution patterns, characterized by Voronoi cells with mostly six corners, negative correlations in their size up to a distance of 13 m, and remarkable homogeneity over large spatial scales. These results cast doubts on abiotic gas-leakage along geological lines or social insects as causal agents of their origin. However, our mathematical model was able to generate spatial patterns that agreed quantitatively in all of these features with the observed patterns. This supports the hypothesis that fairy circles are self-organized vegetation patterns that emerge from positive biomass-water feedbacks involving water transport by extended root systems and soil-water diffusion. Future research should search for mechanisms that explain how the different hypotheses can generate the patterns observed here and test the ability of self-organization to match the birth- and death dynamics of fairy circles and their regional patterns in the density and size with respect to environmental gradients.

Moving beyond abundance distributions: neutral theory and spatial patterns in a tropical forest

Abstract: Assessing the relative importance of different processes that determine the spatial distribution of species and the dynamics in highly diverse plant communities remains a challenging question in ecology. Previous modelling approaches often focused on single aggregated forest diversity patterns that convey limited information on the underlying dynamic processes. Here, we use recent advances in inference for stochastic simulation models to evaluate the ability of a spatially explicit and spatially continuous neutral model to quantitatively predict six spatial and non-spatial patterns observed at the 50 ha tropical forest plot on Barro Colorado Island, Panama. The patterns capture different aspects of forest dynamics and biodiversity structure, such as annual mortality rate, species richness, species abundance distribution, beta-diversity and the species–area relationship (SAR). The model correctly predicted each pattern independently and up to five patterns simultaneously. However, the model was unable to match the SAR and beta-diversity simultaneously. Our study moves previous theory towards a dynamic spatial theory of biodiversity and demonstrates the value of spatial data to identify ecological processes. This opens up new avenues to evaluate the consequences of additional process for community assembly and dynamics.

Mechanisms underlying local functional and phylogenetic beta diversity in two temperate forests.

Abstract: Although trait information has been widely used to explore underlying mechanisms of forest community structure, most studies have focused on local patterns of phylogenetic or functional alpha diversity. Investigations of functional beta diversity, on the other hand, have not been conducted at local scales in a spatially explicit way. In this study, we provide a powerful methodology based on recent advances in spatial point pattern analysis using fully mapped data of large and small trees in two large temperate forest plots. This approach allowed us to assess the relative importance of different ecological processes and mechanisms for explaining patterns of local phylogenetic and functional beta diversity. For both forests and size classes, we found a clear hierarchy of scales: habitat filtering accounted for patterns of phylogenetic and functional beta diversity at larger distances (150-250 m), dispersal limitation accounted for the observed decline in beta diversity at distances below 150 m, and species interactions explained small departures from functional and phylogenetic beta diversity at the immediate plant-neighborhood scale (below 20 m). Thus, both habitat filtering and dispersal limitation influenced the observed patterns in phylogenetic and functional beta diversity at local scales. This result contrasts with a previous study from the same forests, where dispersal limitation alone approximated the observed species beta diversity for distances up to 250 m. In addition, species interactions were relatively unimportant for predicting phylogenetic and functional beta diversity. Our analysis suggests that phylogenetic and functional beta diversity can provide insights into the mechanisms of local community assembly that are missed by studies focusing exclusively on species beta diversity.

The structure of tropical forests and sphere packings.

Abstract: The search for simple principles underlying the complex architecture of ecological communities such as forests still challenges ecological theorists. We use tree diameter distributions—fundamental for deriving other forest attributes—to describe the structure of tropical forests. Here we argue that tree diameter distributions of natural tropical forests can be explained by stochastic packing of tree crowns representing a forest crown packing system: a method usually used in physics or chemistry. We demonstrate that tree diameter distributions emerge accurately from a surprisingly simple set of principles that include site-specific tree allometries, random placement of trees, competition for space, and mortality. The simple static model also successfully predicted the canopy structure, revealing that most trees in our two studied forests grow up to 30–50 m in height and that the highest packing density of about 60% is reached between the 25- and 40-m height layer. Our approach is an important step toward identifying a minimal set of processes responsible for generating the spatial structure of tropical forests.

Unravelling conflicting density- and distance-dependent effects on plant reproduction using a spatially explicit approach

Abstract: Summary Density- and distance-dependent (DDD) mechanisms are important determinants of plant reproductive success (PRS). Different components of sequential PRS can operate either in the same or in different directions and thus reinforce or neutralize each other, and they may also operate at different spatial scales. Thus, spatially explicit approaches are needed to detect such complex DDD effects across multiple PRS components and spatial scales. To reveal DDD effects of different components of early PRS of the Iberian pear (Pyrus bourgaeana) sampled over three consecutive years, we used marked point pattern analysis. Our special interest is to identify conflicting processes that regulate populations at different spatial scales, for example whether DDD on fruit initiation and on fruit development acted in opposite directions. To evaluate the significance of observed mark correlation functions based on empirical data (e.g. fruiting success), we compared them to expectations given by spatially explicit null models. Diverse DDD processes affected several aspects of PRS in a variable extent over the three seasons. First, fruit initiation was higher for individuals with more neighbours at small distances (i.e. up to 40 m). However, P. bourgaeana fruit development decreased with increasing number of nearby neighbours, but these effects cancelled for overall fruit set that did not show DDD effects. Secondly, the absolute number of fruits produced (crop sizes) by trees showed positive density dependence in 2011 and 2012 but not in 2013. Finally, the total number of seeds produced did not show DDD effects, indicating that conflicting demographic processes can disrupt the initial spatial pattern of tree investment in reproduction. Synthesis. Understanding complex spatial effects of DDD processes requires dissection of component processes to attain the complete picture since contrasting DDD processes may be hidden behind a single cumulative measure of reproductive success. The combination of novel and classic mark correlation functions used here constitutes a powerful spatially explicit tool that can be broadly applied to unravel conflicting mechanisms of DDD regulating the persistence of sessile organisms at a range of spatial scales. Our findings help to explain why some authors failed to find expected DDD of PRS and highlight the importance of detailed multiyear field studies on plant reproductive success.

Neighborhood diversity of large trees shows independent species patterns in a mixed dipterocarp forest in Sri Lanka

Abstract: Interactions among neighboring individuals influence plant performance and should create spatial patterns in local community structure. In order to assess the role of large trees in generating spatial patterns in local species richness, we used the individual species-area relationship (ISAR) to evaluate the species richness of trees of different size classes (and dead trees) in circular neighborhoods with varying radius around large trees of different focal species. To reveal signals of species interactions, we compared the ISAR function of the individuals of focal species with that of randomly selected nearby locations. We expected that large trees should strongly affect the community structure of smaller trees in their neighborhood, but that these effects should fade away with increasing size class. Unexpectedly, we found that only few focal species showed signals of species interactions with trees of the different size classes and that this was less likely for less abundant focal species. However, the few and relatively weak departures from independence were consistent with expectations of the effect of competition for space and the dispersal syndrome on spatial patterns. A noisy signal of competition for space found for large trees built up gradually with increasing life stage; it was not yet present for large saplings but detectable for intermediates. Additionally, focal species with animal-dispersed seeds showed higher species richness in their neighborhood than those with gravity- and gyration-dispersed seeds. Our analysis across the entire ontogeny from recruits to large trees supports the hypothesis that stochastic effects dilute deterministic species interactions in highly diverse communities. Stochastic dilution is a consequence of the stochastic geometry of biodiversity in species-rich communities where the identities of the nearest neighbors of a given plant are largely unpredictable. While the outcome of local species interactions is governed for each plant by deterministic fitness and niche differences, the large variability of competitors causes also a large variability in the outcomes of interactions and does not allow for strong directed responses at the species level. Collectively, our results highlight the critical effect of the stochastic geometry of biodiversity in structuring local spatial patterns of tropical forest diversity.

Linking trait similarity to interspecific spatial associations in a moist tropical forest

Abstract: Aim Community assembly theory predicts that niche differentiation promotes the spatial clustering of functionally dissimilar species, whereas habitat filtering has the converse effect We used these predictions to assess the relative effects of habitat filtering and niche differentiation on recruit community assembly over spatial (5- and 30-m neighbourhoods) and temporal (20-yr) scales in the Forest Dynamics Plot at Barro Colorado Island Location Barro Colorado Island, Panama Methods We integrated data on the spatial patterns of ≥1 cm DBH (diameter at 13 m above ground) recruits with data on seven functional traits for 64 species First, we quantified the interspecific association patterns of all species pairs i and j using the K-function Kij(r) and the nearest-neighbour distribution function Dij(r) Second, for those pairs with significant spatial associations, we calculated an index of interspecific spatial association using the results of these two summary statistics Finally, we examined the relationship between interspecific spatial association and trait similarity using simple and partial Mantel tests Results In all censuses, almost one-half of species pairs had no spatial associations, but for pairs that were significantly spatially associated, positive relationships between trait similarity and spatial association occurred in 5-m and 30-m neighbourhoods, whereas significant negative relationships only appeared in 5-m neighbourhoods This suggests that habitat filtering was more important than niche differentiation in assembling recruit communities at 5- and 30-m scales Habitat filtering mainly acted upon traits related to topographic habitat preferences and dispersal mode, whereas spatial association was inversely related to similarity in terms of wood specific gravity and shade tolerance Conclusions Our findings suggest that both stochastic and deterministic processes operate in species-rich ecological communities, but the role of habitat filtering and niche differentiation as determinants of community assembly vary over spatial and temporal scales Species co-occurrence was driven by habitat filtering at small and large scales, but also by a combination of niche differentiation and weaker-competitor exclusion effects at small scales Temporal variations in the importance of habitat filtering and niche differentiation could be related to the occurrence of disturbances such as tree falls Our results emphasize the role of trait-based processes in plant community assembly

Clarifying misunderstandings regarding vegetation self‐organisation and spatial patterns of fairy circles in Namibia: a response to recent termite hypotheses

Abstract: S T E P H A N G E T Z I N, 1 K E R S T I N W I E G A N D, 2 T H O R S T E N W I E G A N D, 1 H E Z I Y I Z H A Q, 3,4 J O S T V O N H A R D E N B E R G 5 and E H U D M E R O N 3,6 1Department of Ecological Modelling, Helmholtz Centre for Environmental Research – UFZ, Leipzig, Germany, 2Department of Ecosystem Modelling, Faculty of Forest Sciences and Forest Ecology, University of Goettingen, Goettingen, Germany, 3Department of Solar Energy and Environmental Physics, Blaustein Institutes for Desert Research, Ben-Gurion University of the Negev, Beer Sheva, Israel, 4The Dead Sea and Arava Science Center, Tamar Regional Council, Israel, 5ISAC-CNR, Torino, Italy and 6Physics Department, Ben-Gurion University of the Negev, Beer Sheva, Israel

Individual Species-Area Relationship of Woody Plant Communities in a Heterogeneous Subtropical Monsoon Rainforest

Abstract: The spatial structure of species richness is often characterized by the species-area relationship (SAR). However, the SAR approach rarely considers the spatial variability of individual plants that arises from species interactions and species' habitat associations. Here, we explored how the interactions of individual plants of target species influence SAR patterns at a range of neighborhood distances. We analyzed the data of 113,988 woody plants of 110 species from the Fushan Forest Dynamics Plot (25 ha), northern Taiwan, which is a subtropical rainforest heavily influenced by typhoons. We classified 34 dominant species into 3 species types (i.e., accumulator, repeller, or no effect) by testing how the individual species-area relationship (i.e., statistics describing how neighborhood species richness changes around individuals) of target species departs (i.e., positively, negatively, or with no obvious trend) from a null model that accounts for habitat association. Deviation from the null model suggests that the net effect of species' interactions increases (accumulate) or decreases (repel) neighborhood species richness. We found that (i) accumulators were dominant at small interaction distances ( 30 m); (iii) repellers were rarely detected; and (iv) large-sized and abundant species tended to be accumulators. The findings suggest that positive species interactions have the potential to accumulate neighborhood species richness, particularly through size- and density-dependent mechanisms. We hypothesized that the frequently disturbed environment of this subtropical rainforest (e.g., typhoon-driven natural disturbances such as landslides, soil erosion, flooding, and windthrow) might create the spatial heterogeneity of species richness and promote positive species interactions.