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1.
William D. Simonson Harriet D. Allen Erin Parham Eduardo de Basto e Santos Paul Hotham 《Landscape Ecology》2018,33(5):811-827
Context
Montados are dynamic agroforestry systems of southern Portugal, with high economic and ecological values. Changes in land use and cover have important implications for landscape-level biodiversity and its conservation.Objectives
Our objectives were to evaluate the biodiversity values and trends in a montado system in the Alentejo, Portugal so as to inform landscape level conservation approaches. In doing so, we aimed to develop a replicable and robust approach drawing together field observation, expert opinion, and remote sensing to produce predictions relevant to land management planning.Methods
Field sampling and subsequent analysis of data on the birds, butterflies and plants in eight distinct land covers allowed the identification of two principal habitat groupings of importance: ‘montado mosaic’ and ‘shrubland’. Morphological spatial pattern analysis was performed on Landsat-derived GIS habitat layers for 1984 and 2009, generating maps and statistics for change in the different landscape functional classes. In addition, we demonstrated how the modelling of ecotones between open and closed biomes can identify the preferred hunting grounds of the threatened Iberian lynx and black vulture, flagship species whose conservation provides benefits to the area’s wider biodiversity values.Results
Total and core area of montado mosaics and shrubland increased over the 25 year period, whilst the amount of habitat connectivity declined in the case of shrubland. Considerable local variation in these trends highlighted targetable areas for conservation action (e.g. through agri-environment spending).Conclusions
A rapid and robust approach was demonstrated, with potentially wider utility for biodiversity assessment and planning.2.
Context
We address the issue of adapting landscapes for improved insect biodiversity conservation in a changing climate by assessing the importance of additive (main) and synergistic (interaction) effects of land cover and land use with climate.Objectives
We test the hypotheses that ant richness (species and genus), abundance and diversity would vary according to land cover and land use intensity but that these effects would vary according to climate.Methods
We used a 1000 m elevation gradient in eastern Australia (as a proxy for a climate gradient) and sampled ant biodiversity along this gradient from sites with variable land cover and land use.Results
Main effects revealed: higher ant richness (species and genus) and diversity with greater native woody plant canopy cover; and lower species richness with higher cultivation and grazing intensity, bare ground and exotic plant groundcover. Interaction effects revealed: both the positive effects of native plant canopy cover on ant species richness and abundance, and the negative effects of exotic plant groundcover on species richness were greatest at sites with warmer and drier climates.Conclusions
Impacts of climate change on insect biodiversity may be mitigated to some degree through landscape adaptation by increasing woody native vegetation cover and by reducing land use intensity, the cover of exotic vegetation and of bare ground. Evidence of synergistic effects suggests that landscape adaptation may be most effective in areas which are currently warmer and drier, or are projected to become so as a result of climate change.3.
Context
Species are expected to shift their distributions in response to global environmental changes and additional protected areas are needed to encompass the corresponding changes in the distributions of their habitats. Conservation policies are likely to become obsolete unless they integrate the potential impacts of climate and land-use change on biodiversity.Objectives
We identify conservation priority areas for current and future projected distributions of Iberian bird species. We then investigate the extent to which global change informed priority areas are: (i) covered by existing protected area networks (national protected areas and Natura 2000); (ii) threatened by agricultural or urban land-use changes.Methods
We use outputs of species distributions models fitted with climatic data as inputs in spatial prioritization tools to identify conservation priority areas for 168 bird species. We use projections of land-use change to then discriminate between threatened and non-threatened priority areas.Results
19% of the priority areas for birds are covered by national protected areas and 23% are covered by Natura 2000 sites. The spatial mismatch between protected area networks and priority areas for birds is projected to increase with climate change. But there are opportunities to improve the protection of birds under climate change, as half of the priority areas are currently neither protected nor in conflict with urban or agricultural land-uses.Conclusions
We identify critical areas for bird conservation both under current and climate change conditions, and propose that they could guide the establishment of new conservation areas across the Iberian Peninsula complementing existing protected areas.4.
Romaike S. Middendorp Alvaro J. Pérez Armando Molina Eric F. Lambin 《Landscape Ecology》2016,31(7):1581-1599
Context
Natural regenerating forests are rapidly expanding in the tropics. Forest transitions have the potential to restore biodiversity. Spatial targeting of land use policies could improve the biodiversity benefits of reforesting landscapes.Objective
We explored the relative importance of landscape attributes in influencing the potential of tree cover increase to restore native woody plant biodiversity at the landscape scale.Methods
We developed land use scenarios that differed in spatial patterns of reforestation, using the Pangor watershed in the Ecuadorian Andes as a case study. We distinguished between reforestation through natural regeneration of woody vegetation in abandoned fallows and planted forests through managed plantations of exotic species on previously cultivated land. We simulated the restoration of woody plant biodiversity for each scenario using LANDIS-II, a process-based model of forest dynamics. A pair-case comparison of simulated woody plant biodiversity for each scenario was conducted against a random scenario.Results
Species richness in natural regenerating fallows was considerably higher when occurring in: (i) close proximity to remnant forests; (ii) areas with a high percentage of surrounding forest cover; and (iii) compositional heterogeneous landscapes. Reforestation at intermediate altitudes also positively affected restoration of woody plant species. Planted exotic pine forests negatively affected species restoration.Conclusions
Our research contributes to a better understanding of the recolonization processes of regenerating forests. We provide guidelines for reforestation policies that aim to conserve and restore woody plant biodiversity by accounting for landscape attributes.5.
Context
Although biodiversity in cities is essential to ensure the healthy functioning of ecosystems and biosecurity over time, biodiversity loss resulting from human interventions in land cover patterns is widespread in urban landscapes. In the Southern Hemisphere, climate change is likely to accelerate the process of landscape upheavals, and consequently biodiversity loss.Objectives & Methods
The aim of this research is to test the potentials of landscape pattern composition and configuration in safeguarding indigenous avifauna against the local impacts of climate change in urban landscapes, with reference to New Zealand. To build up a platform for landscape pattern interpretation, the literature was reviewed and semi-structured interviews with six subject-matter experts were conducted to provide information about the most important avifauna in the study area, key information on their ecological traits and niches, possible impacts of climate change on their primary habitats, and spatial requirements for ongoing species survival as the climate continues to change. A spatial analysis of land cover patterns was undertaken in Wellington, New Zealand using GIS and FRAGSTATS.Results
Although there are still opportunities for biodiversity conservation in the study area, the current land cover patterns are unlikely to safeguard the selected species against climate change impacts.Conclusions
Eight implications for avifauna persistence under climate change are discussed for the first time in relation to a New Zealand context. These implications can give rise to a higher level of informed decision-making on a wide range of practices for biodiversity conservation related to uncertainties associated with climate change.6.
7.
Kevin McGarigal Bradley W. Compton Ethan B. Plunkett William V. DeLuca Joanna Grand Eduard Ene Scott D. Jackson 《Landscape Ecology》2018,33(7):1029-1048
Context
Conservation planning is increasingly using “coarse filters” based on the idea of conserving “nature’s stage”. One such approach is based on ecosystems and the concept of ecological integrity, although myriad ways exist to measure ecological integrity.Objectives
To describe our ecosystem-based index of ecological integrity (IEI) and its derivative index of ecological impact (ecoImpact), and illustrate their applications for conservation assessment and planning in the northeastern United States.Methods
We characterized the biophysical setting of the landscape at the 30 m cell resolution using a parsimonious suite of settings variables. Based on these settings variables and mapped ecosystems, we computed a suite of anthropogenic stressor metrics reflecting intactness (i.e., freedom from anthropogenic stressors) and resiliency metrics (i.e., connectivity to similar neighboring ecological settings), quantile-rescaled them by ecosystem and geographic extent, and combined them in a weighted linear model to create IEI. We used the change in IEI over time under a land use scenario to compute ecoImpact.Results
We illustrated the calculation of IEI and ecoImpact to compare the ecological integrity consequences of a 70-year projection of urban growth to an alternative scenario involving securing a network of conservation core areas (reserves) from future development.Conclusions
IEI and ecoImpact offer an effective way to assess ecological integrity across the landscape and examine the potential ecological consequences of alternative land use and land cover scenarios to inform conservation decision making.8.
Context
The assessment of land-use impacts on biodiversity is one of the central themes of landscape ecology and conservation biology. However, due to the complexity of biodiversity, it is impossible to obtain complete information about the diversity of all species even for small areas, necessitating the selection of individual species or assemblages thereof as species surrogate. In parts of the world where taxonomic expertise is lacking, species identification has hindered progress in biodiversity conservation, and the only practical, relatively-accurate option, is the use of taxonomic minimalism.Objective
We carried out a rapid biodiversity assessment based on three surrogates—land-use (driver-surrogate), terrestrial arthropods (species-surrogate) and morphospecies (taxonomic-surrogate)—to determine the impacts of land-use on biodiversity of the Western Region (Ghana), an area covering ~4 % of the West African biodiversity hotspot.Method
We used diversity profiles to visualize the distribution of a total of 8848 arthropod individuals over seven land-use types which define the complete heterogeneity of the landscape.Results
Here, we present both sample and asymptotic diversity profiles of arthropod morphospecies for each land-use type and the potential of each land-use type for conserving arthropods.Conclusions
We conclude that (1) the morphospecies approach is useful for detecting differences in species diversity of land-use types; (2) the concept of asymptotic diversity may not be necessary for land-use based biodiversity comparison; and (3) maximum diversity profiles are useful for determining the land-use conservation values in cases where pristine areas are not available.9.
Context
Protected areas are a cornerstone of the global strategy for conserving biodiversity, and yet their efficacy in comparison to unprotected areas is rarely tested. In the highly fragmented forests of temperate regions, landscape context and forest history may be more important than protection status for plant species diversity.Objectives
To determine whether there are differences in plant diversity between protected areas and private lands while controlling for landscape context, forest age, and other important factors.Methods
We used a database of 156 one-hectare forest plots distributed over 120,000 km2 in the fragmented forests of southern Ontario to test whether protected areas and private forests differed in native species richness, relative abundance of exotic species, and the probability of finding species of conservation concern.Results
Plots with more forest on the surrounding landscape had higher native species richness, lower abundance of exotic species, and greater probability of supporting at least one species of conservation concern. Young forests tended to have higher abundance of exotics, and were less likely to support species of conservation concern. Surprisingly, privately owned forests had greater native species richness and were more likely to support species of conservation concern once these other factors were accounted for. In addition, there were significant interactions between ownership type, forest history, and landscape context.Conclusions
Our results highlight the importance of privately owned forests in this region, and the need to consider forest history and landscape context when comparing the efficacy of protected areas versus private land for sustaining biodiversity.10.
Context
Changes in land use have disruptive effects on community structure, causing many species to disappear, though a few thrive and become pests.Objectives
To gain understanding on how anthropogenic activity changes spatial patterns of native species diversity while favoring pests, we conducted rapid biodiversity assessments of dacine fruit flies across eight regions in Southeast Asia.Methods
Male lure traps were maintained for 2 days along transects at 233 sites, in forest, agricultural and urban environments.Results
A total of 8393 individuals were collected, belonging to 57 described and 4 new or unidentified species. The majority (78 %) of individuals belonged to 14 pest species, dominated by Bactrocera dorsalis (Hendel). The 57 species represent 38 % of those recorded from the region, indicating effective sampling. Individual flies were collected in highest numbers in urban and agricultural sites, but species diversity was low. Forest samples yielded fewer specimens but highest species diversity, suggesting a shift in community structure after disturbance, benefiting a few pest species at the expense of the broader community, even in the same genus and ecological guild.Conclusions
Dacine fruit flies may be useful in assessing habitat quality and bait systems permit the execution of rapid biodiversity and multi-species conservation assessments. Our results apply to broader patterns concerning biodiversity loss and the emergence of pest species under increasingly intensive land use gradients, and demonstrate the remarkable loss of biodiversity over very narrow distances as forest is converted into agricultural use, hence the importance in maintaining a mosaic of native habitats.11.
Jessica M. Price Janet Silbernagel Kristina Nixon Amanda Swearingen Randy Swaty Nicholas Miller 《Landscape Ecology》2016,31(5):1093-1115
Context
Broad-scale land conservation and management often involve applying multiple strategies in a single landscape. However, the potential outcomes of such arrangements remain difficult to evaluate given the interactions of ecosystem dynamics, resource extraction, and natural disturbances. The costs and potential risks of implementing these strategies make robust evaluation critical.Objectives
We used collaborative scenario modeling to compare the potential outcomes of alternative management strategies in the Two Hearted River watershed in Michigan’s Upper Peninsula to answer key questions: Which management strategies best achieve conservation goals of maintaining landscape spatial heterogeneity and conserving mature forests and wetlands? And how does an increase in wildfire and windthrow disturbances influence these outcomes?Methods
Scenarios were modeled using the VDDT/TELSA state-and-transition modeling suite, and resulting land cover maps were analyzed using ArcGIS, FRAGSTATS, and R statistical software.Results
Results indicate that blending conservation strategies, such as single-ownership forest reserves and working forest conservation easements in targeted areas of the landscape, may better achieve these goals than applying a single strategy across the same area. However, strategies that best achieve these conservation goals may increase the sensitivity of the landscape to changes in wildfire and windthrow disturbance regimes.Conclusions
These results inform decision-making about which conservation strategy or combination of strategies to apply in specific locations on the landscape to achieve optimum conservation outcomes, how to best utilize scarce financial resources, and how to reduce the financial and ecological risks associated with the application of innovative strategies in an uncertain future.12.
Xuan Guo Nicholas C. Coops Sarah E. Gergel Christopher W. Bater Scott E. Nielsen J. John Stadt Mark Drever 《Landscape Ecology》2018,33(3):491-511
Context
The application of regional-level airborne lidar (light detection and ranging) data to characterize habitat patches and model habitat connectivity over large landscapes has not been well explored. Maintaining a connected network of habitat in the presence of anthropogenic disturbances is essential for regional-level conservation planning and the maintenance of biodiversity values.Objectives
We quantified variation in connectivity following simulated changes in land cover and contrasted outcomes when different conservation priorities were emphasized.Methods
First, we defined habitat patches using vegetation structural attributes identified via lidar. Second, habitat networks were constructed for different forest types and assessed using network connectivity metrics. And finally, land cover change scenarios were simulated using a series of habitat patch removals, representing the impact of implementing different spatial prioritization schemes.Results
Networks for different forest structure types produced very different patch distributions. Conservation scenarios based on different schemes led to contrasting changes during land cover change simulations: the scheme prioritizing only habitat area resulted in immediate near-term losses in connectivity, whereas the scheme considering both habitat area and their spatial configurations maintained the overall connectivity most effectively. Adding climate constraints did not diminish or improve overall connectivity.Conclusions
Both habitat area and habitat configuration should be considered in dynamic modeling of habitat connectivity under changing landscapes. This research provides a framework for integrating forest structure and cover attributes obtained from remote sensing data into network connectivity modeling, and may serve as a prototype for multi-criteria forest management and conservation planning.13.
Rita Bastos Manuela D’Amen Bruno Marcos Mário Santos Luís Braz Joana Vicente João P. Honrado João Gonçalves António Monteiro João Alexandre Cabral 《Landscape Ecology》2018,33(12):2221-2237
Context
Current shifts in biodiversity are driven by multiple processes of environmental and landscape change. Particularly, land use/land cover (LU/LC) dynamics are among the major drivers of biodiversity loss worldwide.Objectives
In this study we aim to explore the applicability of a new modelling framework to predict top predators’ responses to LU/LC changes.Methods
The framework integrates remote-sensing based predictors, statistical inference, stochastic-dynamic simulations and spatial projections in a common and interactive approach. From an ecological modelling perspective, the main innovation of our approach lies on the integration of (1) biomass of birds of prey as an upper trophic indicator of the community characteristics that emerge from the habitat quality across multiple scales of organization and (2) fine-scale biophysical attributes to add a new level of understanding about the role of local LU/LC drivers influencing those emergent biodiversity patterns.Results
Based on species data from published atlases this approach allowed transposing species biomass to finer resolutions, overcoming the lack of detailed information for the study area. Our demonstrative case study revealed a disruptive effect of ongoing LU/LC changes in the spatio-temporal distribution of top predators’ biomass, suggesting the possibility of an emergent disturbance pattern in habitat suitability and community stability. Comparative analysis between simulations and independent field data revealed a promising model performance.Conclusions
Our modelling approach highlights the importance of integrating local LU/LC functional dynamics to predict key trophic’ responses, considered as pertinent ecological indicators for biodiversity management under realistic' future changing regional scenarios.14.
Context
East African ecosystems are characterized by the migrations of large herbivores that are highly vulnerable to the recent development of anthropogenic land use change.Objectives
We analyzed land cover changes in the Kenyan-Tanzanian borderlands of the greater Amboseli ecosystem to evaluate landscape connectivity using African elephants as an indicator species.Methods
We used multi-temporal Landsat imagery and a post classification approach to monitor land cover changes over a 43-year period. GIS based methods were accompanied by a literature review for spatial data on land cover changes and elephant migrations.Results
Land cover changed considerably between 1975 and 2017. Wood- and bushlands declined by 16.3% while open grasslands increased throughout the study region (+?10.3%). Agricultural expansion was observed (+?12.2%) occupying important wildlife habitats and narrowing migration corridors. This development has led to the isolation of Nairobi National Park which was previously part of a large contiguous ecosystem. Eight migration corridors were identified of which only one is formally protected. Two others are almost completely blocked by agriculture and three are expected to become endangered under continuing land use changes.Conclusions
Landscape connectivity is still viable for this ecosystem (except for Nairobi National Park). However, the current situation is very fragile as anthropogenic land use changes are threatening most of the identified large mammal migration corridors. Sustainable land use planning with regard to important wildlife habitats and connecting corridors is a crucial task for further conservation work to safeguard a viable future for wildlife populations in the Kenyan-Tanzanian borderlands.15.
Helena Tukiainen Janne Alahuhta Richard Field Terhi Ala-Hulkko Raino Lampinen Jan Hjort 《Landscape Ecology》2017,32(5):1049-1063
Context
‘Conserving Nature’s stage’ has been advanced as an important conservation principle because of known links between biodiversity and abiotic environmental diversity, especially in sensitive high-latitude environments and at the landscape scale. However these links have not been examined across gradients of human impact on the landscape.Objectives
To (1) analyze the relationships between land-use intensity and both landscape-scale biodiversity and geodiversity, and (2) assess the contributions of geodiversity, climate and spatial variables to explaining vascular plant species richness in landscapes of low, moderate and high human impact.Methods
We used generalized additive models (GAMs) to analyze relationships between land-use intensity and both geodiversity (geological, geomorphological and hydrological richness) and plant species richness in 6191 1-km2 grid squares across Finland. We used linear regression-based variation partitioning (VP) to assess contributions of climate, geodiversity and spatial variable groups to accounting for spatial variation in species richness.Results
In GAMs, geodiversity correlated negatively, and plant species richness positively, with land-use intensity. Both relationships were non-linear. In VP, geodiversity best accounted for species richness in areas of moderate to high human impact. These overall contributions were mainly due to variation explained jointly with climate, which dominated the models. Independent geodiversity contributions were highest in pristine environments, but low throughout.Conclusions
Human action increases biodiversity but may reduce geodiversity, at landscape scale in high-latitude environments. Better understanding of the connections between biodiversity and abiotic environment along changing land-use gradients is essential in developing sustainable measures to conserve biodiversity under global change.16.
Context
The Natura 2000 aims to promote the persistence of biodiversity and traditional uses. European landscapes have, however, undergone large transformations in the past decades, mainly associated with the abandonment of less productive lands concentration of intensive agriculture. These changes could pose management challenges and new opportunities to the achievement of the network´s goals.Objective
Evaluate changes in land cover within Natura 2000 in the last two decades.Methods
We use different Corine Land Cover datasets to construct transition matrices of land uses for measuring changes for each Natura 2000 site. We also explore the role of different drivers in observed changes and assess the impacts of these changes in the structure of landscape.Results
Landscape has been highly dynamic within Natura 2000 in the last two decades with more than 20% undergoing land cover changes. The most systematic transitions involved both, succession processes towards naturalisation in older and more abrupt protected areas (PAs) and anthropization in less steep and more recently designated PAs. Changes across land cover categories had also significant effects on the landscape configuration towards a higher homogenisation.Conclusions
Two different strategies would be needed to enhance the role of Natura 2000, (i) tighter control to ensure anthropization, mainly intensive agriculture, does not compromise conservation goals within PAs and (ii) tackle more effectively the ecological and socio-economic effects of abandonment in less productive areas to halt loss of key habitats. On the other hand, changes in composition and structure of landscape open new conservation opportunities derived from enhanced connectivity.17.
Synergistic effects of climate and land cover: grassland birds are more vulnerable to climate change
Marta A. Jarzyna Benjamin Zuckerberg Andrew O. Finley William F. Porter 《Landscape Ecology》2016,31(10):2275-2290
Context
Climate change is not occurring over a homogeneous landscape and the quantity and quality of available land cover will likely affect the way species respond to climate change. The influence of land cover on species’ responses to climate change, however, is likely to differ depending on habitat type and composition.Objectives
Our goal was to investigate responses of forest and grassland breeding birds to over 20 years of climate change across varying gradients of forest and grassland habitat. Specifically, we investigated whether (i) increasing amounts of available land cover modify responses of forest and grassland-dependent birds to changing climate and (ii) the effect of increasing land cover amount differs for forest and grassland birds.Methods
We used Bayesian spatially-varying intercept models to evaluate species- and community-level responses of 30 forest and 10 grassland birds to climate change across varying amounts of their associated land cover types.Results
Responses of forest birds to climate change were weak and constant across a gradient of forest cover. Conversely, grassland birds responded strongly to changing climatic conditions. Specifically, increasing temperatures led to higher probabilities of localized extinctions for grassland birds, and this effect was intensified in regions with low amounts of grassland cover.Conclusions
Within the context of northeastern forests and grasslands, we conclude that forests serve as a possible buffer to the impacts of climate change on birds. Conversely, species occupying open, fragmented grassland areas might be particularly at risk of a changing climate due to the diminished buffering capacity of these ecosystems.18.
Joan Marull Carme Font Enric Tello Nofre Fullana Elena Domene Manel Pons Elena Galán 《Landscape Ecology》2016,31(2):317-336
Context
The role of agricultural landscapes in biodiversity conservation is an emerging topic in a world experiencing a worrying decrease of species richness. Farm systems may either decrease or increase biological diversity, depending on land-use intensities and management.Objectives
We present an intermediate disturbance-complexity model (IDC) of cultural landscapes aimed at assessing how different levels of anthropogenic disturbance on ecosystems affect the capacity to host biodiversity depending on the land matrix heterogeneity. It is applied to the Mallorca Island, amidst the Mediterranean biodiversity hotspot.Methods
The model uses the disturbance exerted when farmers alter the Net Primary Production through land-use change as well as when they remove a share of it (HANPP), together with Shannon–Wiener index (H′) of land-cover diversity. The model is tested with a twofold-scalar experimental design (1:50,000 and 1:5000) of a set of landscape units along three time points (1956, 1989, 2011). Species richness of breeding and wintering birds, taken as a biodiversity proxy, is used in an exploratory factor analysis.Results
The results clearly show that when intermediate levels of HANPP are performed within intermediate levels of complexity (H′) in landscape patterns, like agro-forest mosaics, great bird species richness and high socio-ecological resilience can be maintained. Yet, these complex-heterogeneous landscapes are currently vanishing due to industrial farm intensification, rural abandonment and urban sprawl.Conclusions
The results make apparent the usefulness of transferring the concept of intermediate disturbance-complexity interplay to cultural landscapes. Our spatial-explicit IDC model can be used as a tool for strategic environmental assessment of land-use planning.19.
Context
The variation in spatial distribution between ecosystem services can be high. Hence, there is a need to spatially identify important sites for conservation planning. The term ‘ecosystem service hotspot’ has often been used for this purpose, but definitions of this term are ambiguous.Objectives
We review and classify methods to spatially delineate hotspots. We test how spatial configuration of hotspots for a set of ecosystem services differs depending on the applied method. We compare the outcomes to a heuristic site prioritisation approach (Marxan).Methods
The four tested hotspot methods are top richest cells, spatial clustering, intensity, and richness. In a conservation scenario we set a target of conserving 10 % of the quantity of five regulating and cultural services for the forest area of Telemark county, Norway.Results
Spatial configuration of selected areas as retrieved by the four hotspots and Marxan differed considerably. Pairwise comparisons were at the lower end of the scale of the Kappa statistic (0.11–0.27). The outcomes also differed considerably in mean target achievement, cost-effectiveness in terms of land-area needed per unit target achievement and compactness in terms of edge-to-area ratio.Conclusions
An ecosystem service hotspot can refer to either areas containing high values of one service or areas with multiple services. Differences in spatial configuration among hotspot methods can lead to uncertainties for decision-making. This also has consequences for analysing the spatial co-occurrence of hotspots of multiple services and of services and biodiversity.20.