Measuring the abruptness of patchy ecotones – A simulation-based comparison of landscape pattern statistics

The use of statistics of landscape pattern to infer ecological process at ecotones requires knowledge of the specific sensitivities of statistics to ecotone characteristics. In this study, sets of patch-based and boundary-based statistics were evaluated to assess their suitability as measures of abruptness on simulated ecotone landscapes. We generated 50 realizations each for 25 groups of ecotones that varied systematically in their degree of abruptness and patchiness. Factorial ANOVA was used to evaluate the sensitivity of statistics to the known differences among the simulated groups. Suitability of each index for measuring abruptness was evaluated using the ANOVA results. The statistics were then ranked in order of their suitability as abruptness statistics based on their sensitivity to abruptness, the consistency of the relationship, and their lack of sensitivity to patchiness. The two best statistics for quantifying abruptness were those we developed based on lattice delineation methods, and are called cumulative boundary elements and boundary element dispersion. The results of this research provide support for studies of ecotone process that rely on the interpretation of patch or boundary statistics.     

Spatiotemporal complexity of patchy invasion in a predator-prey system with the Allee effect

Invasion of an exotic species initiated by its local introduction is considered subject to predator-prey interactions and the Allee effect when the prey growth becomes negative for small values of the prey density. Mathematically, the system dynamics is described by two nonlinear diffusion-reaction equations in two spatial dimensions. Regimes of invasion are studied by means of extensive numerical simulations. We show that, in this system, along with well-known scenarios of species spread via propagation of continuous population fronts, there exists an essentially different invasion regime which we call a patchy invasion. In this regime, the species spreads over space via irregular motion and interaction of separate population patches without formation of any continuous front, the population density between the patches being nearly zero. We show that this type of the system dynamics corresponds to spatiotemporal chaos and calculate the dominant Lyapunov exponent. We then show that, surprisingly, in the regime of patchy invasion the spatially average prey density appears to be below the survival threshold. We also show that a variation of parameters can destroy this regime and either restore the usual invasion scenario via propagation of continuous fronts or brings the species to extinction; thus, the patchy spread can be qualified as the invasion at the edge of extinction. Finally, we discuss the implications of this phenomenon for invasive species management and control.     

Validation of the sigmoid wave curve fitting algorithm on a forest-tundra ecotone in the Northwest Territories,Canada

Ecotones are gradual transitions between two adjacent ecological systems. Ecotones are characterized by their spatial properties which are reflected in an ecotone width and location and the temporal variation across time during succession or environmental change on both a local and a global scale. If only one main environmental factor drives this gradual change the shape of the ecotone is evident as a sigmoid wave. In this paper we explore a two-dimensional sigmoid wave curve fitting algorithm that describes an ecotone and validate it on classified remote sensing data of a forest-tundra ecotone in the Northwest Territories of Canada. The estimated location and width of the forest-tundra ecotone are validated with digital land cover data. We conclude that the algorithm was able to delineate the forest-tundra ecotone based upon the classified remote sensing image and is robust for various algorithm parameter settings.     

Spatial patterns of plant richness across treeline ecotones in the Pyrenees reveal different locations for richness and tree cover boundaries

Aim  To forecast the responses of alpine flora to the expected upward shift of treeline ecotones due to climatic warming, we investigated species richness patterns of vascular plants at small spatial scales across elevational transects.
Location  Richness patterns were assessed at local scales along the elevational gradient in two undisturbed treeline ecotones and one disturbed treeline ecotone in the Spanish Pyrenees.
Methods  We placed a rectangular plot (0.3–0.4 ha) in each treeline ecotone. We estimated and described the spatial patterns of plant richness using the point method and Moran's I correlograms. We delineated boundaries based on plant richness and tree cover using moving split windows and wavelet analysis. Then, to determine if floristic and tree cover boundaries were spatially related, overlap statistics were used.
Results  Plant richness increased above the forest limit and was negatively related to tree cover in the undisturbed sites. The mean size of richness patches in one of these sites was 10–15 m. Moving split windows and wavelets detected the sharpest changes in plant richness above the forest limit at both undisturbed sites. Most tree cover and plant richness boundaries were not spatially related.
Main conclusions  The upslope decrease of tree cover may explain the increase of plant richness across alpine treeline ecotones. However, the detection of abrupt richness boundaries well above the forest limit indicates the importance of local environmental heterogeneity to explain the patterns of plant richness at smaller scales. We found highly diverse microsites dominated by alpine species above the forest limit, which should be monitored to describe their response to the predicted upward shift of forests.     

Floristic composition across a climatic gradient in a neotropical lowland forest

Abstract. This study deals with the floristic composition of lowland tropical forest in the watershed of the Panama Canal. The floristic composition of large trees in 54 forest plots was analysed with respect to environmental factors, including precipitation, geologic parent material, stand age, topography, and soils. The plots contain 824 species of trees with a diameter at breast height ≥10 cm and represent a regional flora with exceptional β‐diversity. Plot data indicate that the Panamanian forest is strongly spatially structured at the landscape scale with floristic similarity decreasing rapidly as a function of inter‐plot geographic distance, especially for distances <5 km. The ordinations and patterns of endemism across the study area indicate broad floristic associations well correlated with Holdridge life zones. The results indicate the positive aspects of life zone classification at regional scales, while simultaneously highlighting its inadequacy for finer scales of analysis and resource management. Multivariate gradient analysis techniques (Non‐metric Multidimensional Distance Scaling and Detrended Correspondence Analysis) show clear patterns of floristic variability correlated with regional precipitation trends, surficial geology, and local soil attributes. Geologic and edaphic conditions, such as acidic soils or excessively drained limestone substrates, appear to override the effects of precipitation and modify forest composition. We conclude that the Panamanian forest shows clear patterns of spatial organization along environmental gradients, predominantly precipitation. The rapid decline in floristic similarity with distance between stands also suggests a role for dispersal limitation and stochastic events.     

Estimating the consequences of land-use changes on butterfly diversity in a marginal agricultural landscape in Sweden

Abandonment of farming with the resultant increase in forest cover is one of the major threats to semi-natural grasslands in marginal agricultural areas. In Sweden, the loss of semi-natural grassland is a serious nature conservation problem since it is one of the most species-rich habitats. In this study, the consequences of grassland abandonment and afforestation on butterfly diversity and butterfly dispersal costs are estimated and used to compare three different future land-use scenarios for a marginal agricultural landscape in Sweden. Based on previous butterfly surveys on grasslands in the area, a relationship between land-use type and butterfly diversity was established. By comparing land-use maps of different scenarios, the number of suitable habitat patches and total suitable habitat patch area with low, medium and high butterfly diversity could be estimated. To obtain an indication of possible fragmentation effects, a least-cost analysis was used to compare travel costs of the butterflies between suitable habitat patches for the different scenarios. The results show that different land-use scenarios affect butterfly diversity and travel costs differently. In the extreme case scenario of cessation of full-time farming and a reduction in part-time farming, nearly all valuable butterfly habitats will vanish, since the most species-rich habitats lie in the periphery of the settlement and are expected to be abandoned and afforested first. If, on the other hand, grassland management is less reduced the effect of abandonment on butterflies depends very much on which areas continue to be managed. To preserve the most important grasslands for butterflies an active management strategy for the whole study area would be needed. While it seems relatively easy to identify the areas most important to conserve from a butterfly diversity perspective, it will be more difficult to find an optimal spatial solution that also minimises dispersal costs for butterflies.     

Assignment and evaluation of ground beetle (Coleoptera: Carabidae) assemblages to sites on different scales in a grassland landscape

Carabid beetles and environmental parameters were investigated in 52 grassland sites with three replicate pitfall traps in each site and in the valley of the River Eider in Schleswig–Holstein (northern Germany) with 61 pitfall traps. Environmental parameters included water content of soil, sand content, organic matter content and pH. Ground beetle assemblages were derived by detrended correspondence analyses (DCA) and characterised by the specific environmental conditions as means for each assemblage. On the regional scale including all investigated sites of Schleswig–Holstein, five assemblages were differentiated. On the local scale including the investigated sites in the valley of the River Eider, three assemblages were found corresponding well with those found on the regional scale. Environmental conditions at the sites of the five assemblages were correlated with land use data, soil types, and water level stages provided by three maps of a geographic information system (GIS). The GIS maps were combined to develop smaller areas with land use, soil type and water level stage information. The characteristic environmental conditions were assigned to each area to derive the spatial distribution of the five ground beetle assemblages. Spatial prediction was correct for 65% of investigated sites. The potential area of each assemblage was estimated for the valley. The different grassland areas were evaluated as potential habitats for ground beetle species comparing total species richness with the regional species richness of each assemblage. The comparison shows that species richness in the evaluated assemblages is relatively low compared to the regional potential.     

Top-down control in a patchy environment: revisiting the stabilizing role of food-dependent predator dispersal

In this paper, we revisit the stabilizing role that predator dispersal and aggregation have in the top-down regulation of predator-prey systems in a heterogeneous environment. We consider an environment consisting of sites interconnected by dispersal, and propose a novel mechanism of stabilization for the case with a non-sigmoid functional response of predators. We assume that the carrying capacity of the prey is infinitely large in each site, and show that successful top-down regulation of this otherwise globally unstable system is made possible through an interplay between the unevenness of prey fitness across the sites and the rapid food-dependent migration of predators. We argue that this mechanism of stabilization is different from those previously reported in the literature: in particular, it requires a high degree of synchronicity in local oscillations of species densities across the sites. Prey outbreaks take place synchronously, but the unevenness of prey growth rates across the sites results in a pronounced difference in the species densities, and so the predator quickly disperses to the sites with the highest prey abundances. For this reason, the consumption of prey mostly takes place in the sites with high densities of prey, which assures an efficient suppression of outbreaks. Furthermore, when the total size of prey population is low, the distribution of both species among the sites becomes more even, and this prevents overconsumption of the prey by the predator. Finally, we put forward the hypothesis that this mechanism, when considered in a tri-trophic plankton community in the water column, can explain the stability of the nutrient-rich low-chlorophyll open ocean regions.     

Identification of the key ecological factors influencing vegetation degradation in semi-arid agro-pastoral ecotone considering spatial scales

When assessing re-vegetation project performance and optimizing land management, identification of the key ecological factors inducing vegetation degradation has crucial implications. Rainfall, temperature, elevation, slope, aspect, land use type, and human disturbance are ecological factors affecting the status of vegetation index. However, at different spatial scales, the key factors may vary. Using Helin County, Inner-Mongolia, China as the study site and combining remote sensing image interpretation, field surveying, and mathematical methods, this study assesses key ecological factors affecting vegetation degradation under different spatial scales in a semi-arid agro-pastoral ecotone. It indicates that the key factors are different at various spatial scales. Elevation, rainfall, and temperature are identified as crucial for all spatial extents. Elevation, rainfall and human disturbance are key factors for small-scale quadrats of 300 m × 300 m and 600 m × 600 m, temperature and land use type are key factors for a medium-scale quadrat of 1 km × 1 km, and rainfall, temperature, and land use are key factors for large-scale quadrats of 2 km × 2 km and 5 km × 5 km. For this region, human disturbance is not the key factor for vegetation degradation across spatial scales. It is necessary to consider spatial scale for the identification of key factors determining vegetation characteristics. The eco-restoration programs at various spatial scales should identify key influencing factors according their scales so as to take effective measurements. The new understanding obtained in this study may help to explore the forces which driving vegetation degradation in the degraded regions in the world.     

Estimating niche width using stable isotopes in the face of habitat variability: a modelling case study in the marine environment

Distributions of stable isotopes have been used to infer an organism's trophic niche width, the 'isotopic niche', and examine resource partitioning. Spatial variation in the isotopic composition of prey may however confound the interpretation of isotopic signatures especially when foragers exploit resources across numerous locations. In this study the isotopic compositions from marine assemblages are modelled to determine the role of variation in the signature of prey items and the effect of dietary breadth and foraging strategies on predator signatures. Outputs from the models reveal that isotopic niche widths can be greater for populations of dietary specialists rather than for generalists, which contravenes what is generally accepted in the literature. When a range of different mixing models are applied to determine if the conversion from δ to p-space can be used to improve model accuracy, predator signature variation is increased rather than model precision. Furthermore the mixing models applied failed to correctly identify dietary specialists and/or to accurately estimate diet contributions that may identify resource partitioning. The results presented illustrate the need to collect sufficiently large sample sizes, in excess of what is collected under most current studies, across the complete distribution of a species and its prey, before attempts to use stable isotopes to make inferences about niche width can be made.     

Thermal analysis of marginal conditions to facilitate cryopreservation by vitrification using a semi-empirical approach

This study aims at the thermal analysis of marginal conditions leading to cryopreservation by vitrification, which appears to be the only alternative for indefinite preservation of large-size tissues and organs. The term “marginal conditions” here refers to cooling rates in close range with the so-called critical cooling rate, above which crystallization is avoided. The analysis of thermal effects associated with partial crystallization during vitrification is associated with the coupled phenomena of heat transfer and kinetics of crystallization. This study takes a practical, semi-empirical approach, where heat transfer is analyzed based on its underlying theoretical principles, while the thermal effects associated with partial crystallization are taken into account by means of empirical correlations. This study presents a computation framework to solve the coupled problem, while presenting a proof-of-concept for DP6 as a representative cryoprotective agent. The thermal effects associated with crystallization at various relevant cooling rates are measured in this study by means of differential scanning calorimetry. Results of this study demonstrate that, due to the thermal effects associated with partial crystallization, the cooling rate at the center of a large organ may lag behind the cooling rate in its surroundings under some scenarios, but may also exceed the surroundings cooling rate in other scenarios, leading to counter-intuitive effects associated with partial crystallization.     

Toward a landscape approach in seagrass beds: using macroalgal accumulation to address questions of scale

An experimental investigation of drift macroalgal accumulation in seagrass beds was conducted to determine if the relationship between passively dispersed plant structure and the spatial arrangement of rooted macrophytes differed when examined across two spatial scales. Experiments were performed from December 1992 to April 1993 at four different sites in Tampa Bay, Florida, utilizing artificial seagrass units (ASUs) of uniform shoot length and density but with different areal dimensions [1 m² (S) versus 4 m² (L)]. Drift macroalgae were also collected from 1 m×1 m plots of natural seagrass at each of the experimental sites from November 1990 to May 1992 to determine the relationship between macroalgal abundance and structural characteristics of natural seagrass. Disproportionately higher amounts of macroalgae were captured in L compared to S plots suggesting that macroalgal accumulation does not scale up directly with the areal dimensions of ASU patches. Higher amounts of algae recovered in L plots is in accordance with patterns expected if algae accumulate in zones of attenuated water flow. Neither seagrass shoot density nor blade length could adequately describe the patterns of algal accumulation. These combined results suggest that explanations for trapping/retention of passively dispersed particles should extend beyond traditional measures of vegetation complexity.     

A river-based approach in reconstructing connectivity among protected areas: Insights and challenges from the Balkan region

Global efforts to halt biodiversity loss mandate the establishment of protected areas. In the face of habitat loss and climate uncertainty, large-scale networks of protected areas connected by corridors are needed to increase the dispersal and persistence potential of biota. For example, the recent European Biodiversity Strategy for 2030 defines clear targets for identifying, establishing and integrating ecological corridors, as part of a Trans-European Nature Network. Here, we examined whether rivers could serve as such corridors, facilitating landscape connectivity (i.e. extent to which landscape facilitates or impedes species movement, exchange of genes, natal dispersal and metapopulation dynamics) among protected areas hosted within different countries in the Balkan Peninsula, southeastern Europe. To quantitatively address this question, we calculated the proportion of the river network enclosed within a protected area per country to detect patterns of protection coverage and explored the degree to which spatial connections between 1878 protected areas are supported by the river network. Acknowledging that dams hinder instream continuity, we further quantified potential loss of connections between protected areas caused by the existence of already implemented or planned dam projects upon critical river habitats of endangered fish species. Our results highlight that Balkan rivers have great potential in providing the spatial connections needed to establish landscape connectivity between most of protected areas in the region. Still, heedless hydropower growth and dissimilarities in river protection between neighboring countries remain key challenges for the evolution of a relative framework. Transnational cooperation and systematic planning of infrastructure development could be the only efficient steps towards supporting the establishment of a river-based network to reconstruct connectivity between protected areas and meet biodiversity goals.     

Estimating epidemiological parameters for bovine tuberculosis in British cattle using a Bayesian partial-likelihood approach

Fitting models with Bayesian likelihood-based parameter inference is becoming increasingly important in infectious disease epidemiology. Detailed datasets present the opportunity to identify subsets of these data that capture important characteristics of the underlying epidemiology. One such dataset describes the epidemic of bovine tuberculosis (bTB) in British cattle, which is also an important exemplar of a disease with a wildlife reservoir (the Eurasian badger). Here, we evaluate a set of nested dynamic models of bTB transmission, including individual- and herd-level transmission heterogeneity and assuming minimal prior knowledge of the transmission and diagnostic test parameters. We performed a likelihood-based bootstrapping operation on the model to infer parameters based only on the recorded numbers of cattle testing positive for bTB at the start of each herd outbreak considering high- and low-risk areas separately. Models without herd heterogeneity are preferred in both areas though there is some evidence for super-spreading cattle. Similar to previous studies, we found low test sensitivities and high within-herd basic reproduction numbers (R₀), suggesting that there may be many unobserved infections in cattle, even though the current testing regime is sufficient to control within-herd epidemics in most cases. Compared with other, more data-heavy approaches, the summary data used in our approach are easily collected, making our approach attractive for other systems.     

Ranking bioprocess variables using global sensitivity analysis: a case study in centrifugation

The selection of appropriate operating conditions for bioprocessing is complex due to the large number of interacting stages and variables. Bioprocesses also operate under tight regulation and therefore tools to assist bioprocess design are of significant utility. Conventional approaches for the analysis of variable sensitivities are inadequate. We propose the use of global sensitivity analysis to determine the level of importance of each variable and their interactions. Once key variables have been determined, the designer may focus on the most significant subset. Two case studies are used to demonstrate the applicability of the approach. Each is based on centrifugation and determines the impact of flow-rate, feed viscosity, density difference and particle size, while performance is assessed by supernatant clarification. Significant differences in sensitivities were found between the two studies due to the different feed material properties. Variable sensitivities were found to be system-specific and provide insight for potential operating strategies.     

Efficiency of using spatial analysis in first-generation coastal Douglas-fir progeny tests in the US Pacific Northwest

Single-trial and across-trial spatial analyses using autoregressive error structures were conducted for growth traits based on 1,146 data sets from 275 Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] progeny trials in 45 first-generation breeding zones in the US Pacific Northwest. The breeding zones encompassed a wide range of latitude, longitude, and elevation. Efficiency of using spatial analysis in reducing variation due to site heterogeneity, estimating genetic parameters, and increasing prediction accuracy was compared among different experimental designs, traits, assessment ages, and tree spacings. More than 97% of the data sets showed significant model improvement with spatial analysis, and height showed more improvement than diameter or volume. Spatial analysis on average removed 14~34% of residual variance due to spatial heterogeneity, which resulted in an up to 20% increase in accuracy of breeding value prediction. The coefficient of variation decreased substantially due to spatial adjustment. Rank correlation between predicted gains before and after spatial analysis was about 0.96, and spatial analysis had little effect on the average predicted gain of the top 20% of parents. We did not observe substantial geographic trends in improvements due to spatial adjustment. Across-site spatial analysis had almost no effect on genotype-by-environment interaction but tended to increase among-trial heterogeneity of residual variance. Two different methods for across-trial spatial analysis were compared and discussed.     

Identifying habitat types in a disturbed area of the forest-steppe ecotone of Patagonia

We surveyed an old ranch (22,000 ha) in the eastern catchment of lake Nahuel Huapi (Argentina), representing both landscape and historical use patterns of the forest-steppe ecotone in the Andes foothills. The objective was to describe plant-environment relations and to develop a procedure to classify habitat types for conservation aims. Floristic and landscape variables were recorded in 72 upland sites. Vegetation types were classified with Two-way indicator Species Analysis and environmental gradients detected with Detrended Canonical Correspondence Analysis. Vegetation types were tested for environmental homogeneity with Discriminant Analysis. A key to classify habitat types was built out from the classification functions obtained by DA analysis. Floristic classification resulted in 7 steppe types and 4 shrubland/woodland types. Vegetation samples were placed along two main gradients, one related to topography and the other to the type of substrate. Steppe types occur in all positions of the topographic gradient forming a temperature-soil moisture continuum, but only (except one of them) on accumulated substrates. Islands of woody types are confined to relatively warm positions of the topographic gradient and to deflated, rocky substrates. This distribution pattern would be caused by frost sensibility of forest species, and by the different strategies of water capture of woody and herbaceous plants. Three pairs of plant assemblages were not discriminated, allowing classification of 8 habitat types out of the 11 communities. Undiscriminated communities (3 pairs) and misclassified samples (29%) were discussed as the result of disturbances.     

Designing ecological corridors in a fragmented landscape: A fuzzy approach to circuit connectivity analysis

Landscape connectivity analysis is a major tool in supporting biodiversity conservation. Several methodologies have been developed to tackle it by following two main paths. The first path exploits graph approaches and models focal nodes’ connections on a resistance/conductance matrix depending on focal species’ movement potential. The second path considers geometrical pattern analyses based on the calculation of structural landscape metrics. These approaches separately investigate functional and structural features of the landscape, and may come short of a total definition if used separately. Here we propose a new scalable, modular, participative and open-source procedure based on Fuzzy logic to combine the functional and structural aspects of connectivity. We applied this method on the highly fragmented landscape of the Po Plain, focusing on its rare and endangered plain springs named fontanili. We identified an expert panel and involved it in the assignation of permeability values of land use classes with respect to the capacity of movement of animal species typical of fontanili. We concurrently performed a quantitative evaluation of the landscape fragmentation with a moving window. We found that the functional and structural evaluations were poorly correlated in the area under study (Pearson's r = −0.35, p < 0.001). We thus integrated these two non-overlapping analyses of the landscape by Fuzzy logic using thresholds and combination weights obtained from questionnaires proposed to the expert panel. The resulting index, termed Fuzzy Functionality Index (FFI), improved the level of information associated with landscape classification. By merging functional and structural aspects of the landscape, the FFI allowed us to discriminate different functional values of equally permeable parcels and vice versa. We demonstrate that FFI may act as a conductance measure in a circuit theory approach, highlighting ecological corridors between focal points of species’ distribution. We present FFI as an effective predictive index to inspect complex and non-linear landscape dynamics.     

Human activity facilitates altitudinal expansion of exotic plants along a road in montane grassland,South Africa

Question: Do anthropogenic activities facilitate the distribution of exotic plants along steep altitudinal gradients? Location: Sani Pass road, Grassland biome, South Africa. Methods: On both sides of this road, presence and abundance of exotic plants was recorded in four 25‐m long road‐verge plots and in parallel 25 m x 2 m adjacent land plots, nested at five altitudinal levels: 1500, 1800, 2100, 2400 and 2700 m a.s.l. Exotic community structure was analyzed using Canonical Correspondence Analysis while a two‐level nested Generalized Linear Model was fitted for richness and cover of exotics. We tested the upper altitudinal limits for all exotics along this road for spatial clustering around four potential propagule sources using a t‐test. Results: Community structure, richness and abundance of exotics were negatively correlated with altitude. Greatest invasion by exotics was recorded for adjacent land at the 1500 m level. Of the 45 exotics, 16 were found at higher altitudes than expected and observations were spatially clustered around potential propagule sources. Conclusions: Spatial clustering of upper altitudinal limits around human inhabited areas suggests that exotics originate from these areas, while exceeding expected altitudinal limits suggests that distribution ranges of exotics are presently underestimated. Exotics are generally characterised by a high propagule pressure and/or persistent seedbanks, thus future tarring of the Sani Pass may result in an increase of exotic species richness and abundance. This would initially result from construction‐related soil disturbance and subsequently from increased traffic, water run‐off, and altered fire frequency. We suggest examples of management actions to prevent this.