One of the central problems in ecology is how to scale from small‐scale observations and experiments to large‐scale patterns and processes. One approach to such upscaling is to use dynamic simulation ...models, but their application to large scales relevant for management is limited by computational costs, and their outputs are difficult to analyse without a systematic strategy. Our general objective is to propose such a strategy. The idea is to approximate the dynamics of detailed simulation models through a set of states, external drivers, and transition matrices, and then use Markov chain and network analysis of the resulting transition matrices to gain insights into the dynamics of the underlying detailed model. We used the individual‐based model COIRON, which simulates the dynamics of semiarid grass steppes in Patagonia (Argentina) under alternative grazing management, as example. Our specific objectives are to identify pathways of degradation and rehabilitation, as well as critical grazing thresholds and early‐warning vegetation states to guide sustainable grazing management in these steppes. Our results indicate nonlinear effects of stocking rate and grazing season on steppe dynamics. Markov chain analysis suggests benefits of seasonal over continuous grazing at intermediate stocking rates, and network analysis of recovery and degradation trajectories shows that intermediate stocking rates maximize differences between grazing seasons. Finally, our analysis identified specific vegetation states as early warning signals that indicate a high risk of irreversible vegetation changes. Patagonian grass steppes should ideally be managed with multi‐paddock grazing at moderate stocking rates around 0.5 sheep ha−1. The transition matrices summarize the relevant key features of the detailed model for larger scales, and applying Markov and network theory provides a systematic strategy to analyse its dynamics to respond to biological questions, both are often difficult to obtain by direct analysis of the detailed model.
The spatial placement of recruits around adult conspecifics represents the accumulated outcome of several pattern-forming processes and mechanisms such as primary and secondary seed dispersal, ...habitat associations or Janzen–Connell effects. Studying the adult–recruit relationship should therefore allow the derivation of specific hypotheses on the processes shaping population and community dynamics. We analysed adult–recruit associations for 65 tree species taken from six censuses of the 50 ha neotropical forest plot on Barro Colorado Island (BCI), Panama. We used point pattern analysis to test, at a range of neighbourhood scales, for spatial independence between recruits and adults, to assess the strength and type of departure from independence, and its relationship with species properties. Positive associations expected to prevail due to dispersal limitation occurred only in 16% of all cases; instead a majority of species showed spatial independence (≈73%). Independence described the placement of recruits around conspecific adults in good approximation, although we found weak and noisy signals of species properties related to seed dispersal. We hypothesize that spatial mechanisms with strong stochastic components such as animal seed dispersal overpower the pattern-forming effects of dispersal limitation, density dependence and habitat association, or that some of the pattern-forming processes cancel out each other.
The analysis of the spatial phylogenetic and phenotypic structure of plant communities can provide insight into the underlying processes and interactions governing their assembly, and how these may ...change during plant ontogeny. We used point pattern analysis to find out if saplings and adult plants are surrounded by phylogenetically and phenotypically more similar or more dissimilar neighbours than expected by chance, and whether these associations change from the sapling to the adult stage. To this end, we combined information on the phylogenetic structure and eight phenotypic traits of 15 woody plant species in two Mediterranean mixed forests of southeastern Spain. At the community level, we found that the sapling bank at both sites did not show phylogenetic or phenotypic spatial patterns, but adults showed phylogenetic clustering (i.e. heterospecific neighbours were more similar than expected). At the species level, we found frequently repulsive patterns in the sapling bank of less abundant species (i.e. heterospecific sapling or adult neighbours were more dissimilar than expected) in both, phylogenetic and phenotypic analyses. For the adult stage, we found phylogenetic attraction (i.e. more similar neighbours) in just one species and phenotypic clustering in four species. The processes driving the assembly of the communities of saplings and adults leave detectable signals in the spatial phylogenetic and phenotypic structure of our two forest communities. Our findings reinforce the existence of ontogenetic shifts in the mechanisms involved in plant community assembly. Facilitation between phylogenetically distant and phenotypically divergent species favours the recruitment of less abundant species. However, processes acting later in the ontogeny ameliorate the competition between close relatives and determine the spatial structure of adult plants. Nevertheless, the role of phenotype in shaping the interactions between adult plants was context‐ and trait‐dependent. The use of spatial point pattern analysis allowed a nuanced interpretation of the phylogenetic and phenotypic structures of plant communities.
Studying the spatial pattern and interspecific associations of plant species may provide valuable insights into processes and mechanisms that maintain species coexistence. Point pattern analysis was ...used to analyze the spatial distribution patterns of twenty dominant tree species, their interspecific spatial associations and changes across life stages in a 20-ha permanent plot of seasonal tropical rainforest in Xishuangbanna, China, to test mechanisms maintaining species coexistence. Torus-translation tests were used to quantify positive or negative associations of the species to topographic habitats. The results showed: (1) fourteen of the twenty tree species were negatively (or positively) associated with one or two of the topographic variables, which evidences that the niche contributes to the spatial pattern of these species. (2) Most saplings of the study species showed a significantly clumped distribution at small scales (0-10 m) which was lost at larger scales (10-30 m). (3) The degree of spatial clumping deceases from saplings, to poles, to adults indicates that density-dependent mortality of the offspring is ubiquitous in species. (4) It is notable that a high number of positive small-scale interactions were found among the twenty species. For saplings, 42.6% of all combinations of species pairs showed positive associations at neighborhood scales up to five meters, but only 38.4% were negative. For poles and adults, positive associations at these distances still made up 45.5% and 29.5%, respectively. In conclusion, there is considerable evidence for the presence of positive interactions among the tree species, which suggests that species herd protection may occur in our plot. In addition, niche assembly and limited dispersal (likely) contribute to the spatial patterns of tree species in the tropical seasonal rain forest in Xishuangbanna, China.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Abstract
Network analysis is an important tool to analyze the structure of complex systems such as tropical forests. Here, we infer spatial proximity networks in tropical forests by using network ...science. First, we focus on tree neighborhoods to derive spatial tree networks from forest inventory data. In a second step, we construct species networks to describe the potential for interactions between species. We find remarkably similar tree and species networks among tropical forests in Panama, Sri Lanka and Taiwan. Across these sites only 32 to 51% of all possible connections between species pairs were realized in the species networks. The species networks show the common small-world property and constant node degree distributions not yet described and explained by network science. Our application of network analysis to forest ecology provides a new approach in biodiversity research to quantify spatial neighborhood structures for better understanding interactions between tree species. Our analyses show that details of tree positions and sizes have no important influence on the detected network structures. This suggests existence of simple principles underlying the complex interactions in tropical forests.
Abstract Model testing is a central step of spatial point pattern analysis, which allows ecologists to judge if their data agree with ecological hypotheses. We present a simple and elegant solution ...of a challenging problem: the construction of a goodness‐of‐fit envelope test with prescribed significance level α . Our new Analytical Global Envelope ( AGE ) test is not restricted to the narrow frame of complete spatial randomness testing and its envelopes can be determined by mathematical calculations. This allows us to investigate the influence of key settings of the AGE test on the width of the envelope strip. To circumvent some assumptions of the simulation‐free AGE test we present a corresponding Simulation‐Based Global Envelope ( SBGE ) test. The envelope strip of the AGE and the SBGE test encircles the range of a summary function such as the pair correlation function under the null model, and it has the desired property that the null hypothesis can be rejected with significance level α if the empirical summary function wanders outside the envelopes. The AGE test can be applied under the mild conditions that the values of the summary functions under the null model are (approximately) normally distributed and are (approximately) independent for different distance bins r j . The SBGE test requires only the independence assumption. The width of the strip of the AGE envelopes scales for a broad range of point processes with 1/ n , where n is the number of points. This casts doubt about attempts of goodness‐of‐fit testing with low n (say <100). The AGE and SBGE test operate with wider envelope strips than the classical “pointwise” test. Therefore, the pointwise test has to be considered as too liberal. Furthermore, we show that the width of the AGE / SBGE strip increases approximately with ln( b ), where b is the number of distance bins. For example, the AGE / SBGE envelopes are for b = 20 more than 50% wider than the corresponding pointwise envelopes. Our study opens up new avenues to the test problem in point pattern statistics and the new AGE and SBGE tests can be widely applied in ecology to improve the practice in null model testing.
Because orchids are dependent on mycorrhizal fungi for germination and establishment of seedlings, differences in the mycorrhizal communities associating with orchids can be expected to mediate the ...abundance, spatial distribution and coexistence of terrestrial orchids in natural communities.
We assessed the small-scale spatial distribution of seven orchid species co-occurring in 25 × 25 m plots in two Mediterranean grasslands. In order to characterize the mycorrhizal community associating with each orchid species, 454 pyrosequencing was used. The extent of spatial clustering was assessed using techniques of spatial point pattern analysis.
The community of mycorrhizal fungi consisted mainly of members of the Tulasnellaceae, Thelephoraceae and Ceratobasidiaceae, although sporadically members of the Sebacinaceae, Russulaceae and Cortinariaceae were observed. Pronounced differences in mycorrhizal communities were observed between species, whereas strong clustering and significant segregation characterized the spatial distribution of orchid species. However, spatial segregation was not significantly related to phylogenetic dissimilarity of fungal communities.
Our results indicate that co-occurring orchid species have distinctive mycorrhizal communities and show strong spatial segregation, suggesting that mycorrhizal fungi are important factors driving niche partitioning in terrestrial orchids and may therefore contribute to orchid coexistence.
Environmental filtering and dispersal limitation can both maintain diversity in plant communities by aggregating conspecifics, but parsing the contribution of each process has proven difficult ...empirically. Here, we assess the contribution of filtering and dispersal limitation to the spatial aggregation patterns of 456 tree species in a hyperdiverse Amazonian forest and find distinct functional trait correlates of interspecific variation in these processes. Spatial point process model analysis revealed that both mechanisms are important drivers of intraspecific aggregation for the majority of species. Leaf drought tolerance was correlated with species topographic distributions in this aseasonal rainforest, showing that future increases in drought severity could significantly impact community structure. In addition, seed mass was associated with the spatial scale and density of dispersal-related aggregation. Taken together, these results suggest environmental filtering and dispersal limitation act in concert to influence the spatial and functional structure of diverse forest communities.
Context
Land-use change is one of the main threats to biodiversity on the global scale. Legacy effects of historical land-use changes may affect population dynamics of long-lived species, but they ...are difficult to evaluate through observational studies alone. We present here an interdisciplinary modelling approach as an alternative to address this problem in landscape ecology.
Objectives
Assess effects of agricultural abandonment and anthropisation on the population dynamics of long-lived species. Specifically, we evaluated: (a) how changes in movement patterns caused by land-use change might impact population dynamics; (b) time-lag responses of demographic variables in relation to land-use changes.
Methods
We applied an individual-based and spatial-explicit simulation model of the spur-tighed tortoise (
Testudo graeca
), an endangered species, to sequences of real-world landscape changes representing agricultural abandonment and anthropisation at the local scale. We analysed different demographic variables and compared an “impact scenario” (i.e., historical land-use changes) with a “control scenario” (no land-use changes).
Results
While agricultural abandonment did not lead to relevant changes in demographic variables, anthropisation negatively affected the reproductive rate, population density and the extinction probability with time-lag responses of 20, 30 and 130 years, respectively, and caused an extinction debt of 22%.
Conclusions
We provide an understanding of how changes in animal movement driven by land-use changes can translate into lagged impacts on demography and, ultimately, on population viability. Implementation of proactive mitigation management are needed to promote landscape connectivity, especially for long-lived species for which first signatures of an extinction debt may arise only after decades.
While the successional dynamics and large-scale structure of Douglas-fir forest in the Pacific Northwest region is well studied, the fine-scale spatial characteristics at the stand level are still ...poorly understood. Here we investigated the fine-scale spatial structure of forest on Vancouver Island, in order to understand how the three dominant species, Douglas-fir, western hemlock, and western redcedar, coexist and partition space along a chronosequence comprised of immature, mature, and old-growth stands. We quantified the changes in spatial distribution and association of the species along the chronosequence using the scale-dependent point pattern analyses pair-correlation function g(r) and Ripley's L-function. Evidence on intra- and inter-specific competition was also inferred from correlations between nearest-neighbor distances and tree size. Our results show that 1) the aggregation of Douglas-fir in old-growth was primarily caused by variation in local site characteristics, 2) only surviving hemlock were more regular than their pre-mortality patterns, a result consistent with strong intra-specific competition, 3) inter-specific competition declined rapidly with stand age due to spatial resource partitioning, and 4) tree death was spatially randomly distributed among larger overstory trees. The study highlights the importance of spatial heterogeneity for the long-term coexistence of shade-intolerant pioneer Douglas-fir and shade-tolerant western hemlock and western redcedar.