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Introduction to Spatial Ecology and Its Relevance for Conservation: Applications with R


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How space directly and indirectly affects biodiversity and ecosystem functioning is the focus of several subdisciplines in the life sciences. All of these subdisciplines share concepts and analytical methods that stem from the field of spatial ecology. Spatial ecology focuses on the study and modeling of the role(s) of space on ecological processes that in turn affects ecological patterns. Our goal is to introduce why space is important for ecology and conservation, and how various components of space can inform ecological processes and be modeled. We introduce these issues by tracking the history and development of spatial ecology and conservation, and how different types of modeling frameworks have been advanced over the years to capture spatial problems. Spatial ecology largely arose from key empirical and theoretical developments in the 1950s and 1970s that emphasized how spatial heterogeneity could promote population persistence and how dispersal could have major impacts on populations and communities. With more recent growth in spatial models, the availability of spatial data, computing capacity, and ongoing large-scale environmental change, spatial ecology has matured as a discipline. The maturation has led to spatial ecology becoming integral to the entire fields of ecology and conservation.
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Chapter 1
Introduction to Spatial Ecology and Its
Relevance for Conservation
1.1 What Is Spatial Ecology?
Space: The nal frontierKareiva (1994)
All aspects of ecology play out in space. From Darwins entangled bank to
Hutchinsons ecological theater (Hutchinson 1965; Darwin 1859), space is inherent
to all processes and research in ecology. The importance of space has captured the
imagination of biologists interested in a wide variety of topics, such as migration,
species coexistence, deforestation, and the spread of invasive species. Therefore,
how space directly and indirectly affects biodiversity and ecosystem functioning is
implicitly and/or explicitly the focus of several subdisciplines in the life sciences
(Fig. 1.1).
All of these subdisciplines share concepts and analytical methods that stem from the
eld of spatial ecology: a eld coined by Tilman and Karieva in 1997. Since then, the
term spatial ecologyhas been used in a wide range of ways depending on each
ecological subdiscipline and eld. Biogeography focuses on species geographic distri-
butions (Lomolino 2017). Landscape ecology relates spatial heterogeneity to ecological
processes and species distribution (Turner and Gardner 2015). Movement ecology
focuses on organismal dispersal and migration (Nathan et al. 2008). Macroecology
investigates the relation of processes and species at large spatial scales (Gaston and
Blackburn 2000). Metaecology considers dispersal and spatial interactions at different
spatial scales to model ecological processes that affect species distribution and dynam-
ics (i.e., metapopulations, metacommunities, metaecosystems; Massol et al. 2011).
Spatial and landscape genetics relate how landscape features affect gene glow and
local adaptation (Manel et al. 2003; Guillot et al. 2009). Finally, conservation biology
develops and applies spatial solutions to a variety of problems, including mitigating the
effects of roads, protected area networks, and spatial prioritization in conservation
planning (Primack 2014)(Fig.1.1).
Throughout this book, we use the term spatial ecology in a broad sense referring
to the study and modeling of the role(s) of space on ecological processes (e.g.,
©Springer Nature Switzerland AG 2018
R. Fletcher, M.-J. Fortin, Spatial Ecology and Conservation Modeling,
population dynamics, species interactions, dispersal) that in turn affects ecological
patterns, such as species distributions. This denition shares similarities with some
early denitions of landscape ecology (Pickett and Cadenasso 1995; Turner 1989).
Yet over the years, landscape ecology evolved to include socio-economic aspects of
landscapes as well (Wu 2017).
Research in spatial ecology aims to understand the processes that affect species
distributions and dynamics, and how these processes play out across space. Endog-
enous processes are related to the dynamics of each ecological entity (e.g., movement,
dispersal, and migration) and the interactions among entities within and across
species (population demographics, genetic variation, behavior, competition, facilita-
tion, trophic interactions, etc.). Exogenous processes are related to the response of
organisms to environmental factors that are themselves spatially structured (climate,
local habitat features, microhabitat heterogeneity, patch disturbance-succession,
environmental ltering, historical contingencies, etc.). Overall, it is the combined
action and feedback effects of these endogenous and exogenous processes that result
in the spatial patterns observed at different levels of organization though space (e.g.,
metapopulations, metacommunities, and metaecosystems) (Fig. 1.2).
Spatial ecology is increasingly applied to conservation and management to help
deliver more effective ways to conserve biodiversity. The rapid rate at which
landscapes are altered is creating spatially heterogeneous environmental conditions
that affect species ability to disperse and ultimately persist. Yet, even in homoge-
neous environments, endogenous processes alone can shape species spatial distri-
butions (Okubo 1974). This is why many of the core ecological theories and
analytical models used in spatial ecology are process-based ones. Therefore, one
of the most important cornerstones of spatial ecology as a discipline is the way in
Animal Movement
Plant Dispersal
pattern /
Fig. 1.1 Spatial
subdisciplines derived from
ecological disciplines using
a spatial ecology framework
to tackle current
conservation issues
2 1 Introduction to Spatial Ecology and Its Relevance for Conservation
which the challenges of understanding the processes underlying the spatial distribu-
tion of ecological entities are tackled. Spatial ecology offers concepts and tools to
understand, predict, and map how biodiversity responds to environmental change.
1.2 The Importance of Space in Ecology
Species dynamics occur over space and time. Space affects species in multiple ways
from how they use resources and occupy space within their home range and
throughout their geographical range, how they move, disperse, and migrate through
heterogeneous landscapes, as well as how they interact with other species
(Table 1.1).
To determine the relative importance of space on ecological patterns and pro-
cesses, both mathematical and statistical models are frequently used (Dale and Fortin
2014; Cantrell et al. 2009; Fortin et al. 2012; Ovaskainen et al. 2016). These two
modeling approaches encompass stark differences from data needs, model assump-
tions, and epistemologies (Fig. 1.2). Both process-based (e.g., mathematical, sto-
chastic simulations and computational models) and phenomenological approaches
(e.g., statistical regression models) have a long history of contributing to our under-
standing of the spatial distribution of ecological entities from ne to broad scales
(Levin 1976; MacArthur and Wilson 1967). Such spatial models aim to improve our
understanding of the underlying processes acting on species distributions (e.g., to
estimate the relative importance of environmental drivers versus dispersal to species
distributions) and to perform ecological forecasting (e.g., to predict species distribu-
tions based on such processes; Pagel and Schurr 2012; Dietze 2017).
The foundation for spatial ecology can be traced largely to the seminal paper of
Watt (1947) on the relationship between spatial pattern and ecological processes.
Watt (1947) emphasized that plants occurred in bounded communitiespatches
that form a dynamic mosaic across the landscape, what has become known as the
SA-sp: Spatial autocorrelation of the species
SA-env: Spatial structure of the environment
SD: Spatial dependence of species response to
spatially structured environment
Environment Species
SA-env SA-sp
Fig. 1.2 How space affects
both the spatial structure of
the environmental
conditions and species
distribution. Species
distribution is also affected
by the spatial structure of the
environmental data (adapted
from Wagner and Fortin
1.2 The Importance of Space in Ecology 3
shifting-mosaic steady stateconcept (Bormann and Likens 1979). Then, in
the 1950s and 1960s, there were three key areas of research that emphasized the
importance of space for ecological processes and its relevance for conservation. First,
some inuential experimental studies highlighted the importance of space for ecol-
ogy. In a seminal experiment, Huffaker (1958) showed how predatorprey dynamics
could be stable when including the potential for spatial refugia of prey, while stability
was not possible in small, homogenous habitats. This result was important because
prior to that time, spatial concepts had not been formally considered in theory and
concepts regarding species coexistence. This experiment emphasized the role of
movement in altering species interactions and community structure, a theme that
has persisted and grown over time.
A second area of conceptual development came from theoretical ecology
(Hastings and Gross 2012), where ecologists investigated how diffusion of organ-
isms through space can alter population and community dynamics (Skellam 1951;
Okubo and Levin 2001; Hilborn 1979). Skellam (1951) pioneered these ideas by
applying reactiondiffusion models originally derived for molecular processes to the
problem of dispersal and population dynamics. In this model, Skellam (1951)
assumed diffusion (or random movement) of organisms. While it is clear that
organisms do not move in a simple random manner, the utility of this approach is
that this simple formulation can go a long way in explaining observed patterns in
ecology (Kareiva 1982,1983), and it can be extended to capture non-random issues
(e.g., advection; Reeve et al. 2008). In addition, Skellams work set the stage for
modeling invasive spread, a topic of great importance to conservation biology.
The third area is the application of biogeographic concepts to our understanding
of speciesarea relationships by Preston (1948,1962) and later MacArthur and
Table 1.1 Examples of how space can be incorporated into spatial analyses and their effects on
ecological processes (adapted from Fortin et al. (2012))
Spatial aspects Effects on ecological processes and data
xycoordinates Location of data according to positions of other locations (Euclidean or
relative distance)
The magnitude, spatial scale, and directionality of data values as a function of
distances between data point locations
Locations of abiotic predictors affect the responses of biotic/ecological
Spatial legacy Inuence of past spatial pattern on current ecological processes and species
current spatial pattern
Inuence of nearby locations (local neighbors) on ecological processes and
species spatial pattern
How the intervening landscape features affect daily animal movement and
species dispersal ability
Multiple spatial
Additive spatial scales inuence current spatial pattern
4 1 Introduction to Spatial Ecology and Its Relevance for Conservation
Wilson (1963,1967). This area was particularly crucial in developing the application
of spatial ecology to practical issues of conservation (Higgs 1981). Indeed, many
ecological theories and conservation concepts, including practical solutions, stem
from island biogeography theory, where the size of islands/patches and their spatial
conguration (spacing/isolation) are critical for species persistence through variation
in colonization and extinction events (MacArthur and Wilson 1967; Laurance 2008).
The current era of spatial ecology has grown from island biogeography, where
dispersal of individuals is key and can act as a rescue effect or spatial insurance
(Loreau et al. 2003a) that protects a population from local extinction. Here, species
are often considered to act as metapopulations (Hanski 1999; Levins 1969). The
concept of spatial insurance has been extended to dispersal of several species to
maintain species assemblages and communities as metacommunities (Leibold et al.
2017,2004) and to maintain ecosystem functions as metaecosystems (Loreau et al.
2003b; Guichard 2017).
1.3 The Importance of Space in Conservation
Conservation biologists have increasingly embraced the importance of space in the
conservation of biodiversity and ecosystem services (Schagner et al. 2013; Moilanen
et al. 2009). Space is relevant for conservation in four major ways: (1) it is essential
for spatial mapping of biodiversity and ecosystem services; (2) it provides guidance
for mitigating effects of environmental change; (3) it facilitates effective prioritiza-
tion of areas for conservation; and (4) it provides key components of tools and
models used in conservation.
Several biogeography and macroecology theories provide spatial foundations for
understanding and mapping biodiversity across the planet. The emphasis on spatial
components rst emerged in the eld of biogeography, where there was interest in
identifying and understanding species distributions and geographic gradients in
biodiversity throughout the world. For instance, early on scientists emphasized the
latitudinal gradient of diversity, where diversity was greater in the tropics than in the
temperate zone (Currie and Paquin 1987). Understanding this and other biogeo-
graphic (and macroecological) patterns have been, and continue to be, of interest in
conservation as it helps identify hotspots of biodiversity and endemism of conser-
vation relevance (Myers et al. 2000; Dawson et al. 2017; Orme et al. 2005).
Many approaches to mitigating the effects of environmental change embrace spatial
concepts. For example, the use of corridors in conservation explicitly emphasizes how
the spatial conguration of the environment can promote biodiversity (Crooks and
Sanjayan 2006). Translocations and re-introduction programs require understanding
how potential release locations may inhibit or foster the success of such programs
(Seddon et al. 2014). Adaptation strategies to mitigate the effects of climate change
often emphasize spatial ecological concepts (Heller and Zavaleta 2009).
1.3 The Importance of Space in Conservation 5
Conservation prioritization and planning, one of the major foci for conservation
biology, also emphasizes the importance of spatial ecology. Early rules for conser-
vation planning embraced the need to limit isolation of protected areas and maximize
their area (Diamond 1975). Later work has embraced explicit mapping of conserva-
tion prioritization strategies and how issues such as complementarity of biodiversity
among protected areas is essential for efcient conservation planning (Margules and
Pressey 2000). More recently, conservation planning for climate change emphasizes
how key areas are currently connected and how connectivity may change as climate
and land use continue to change (Pressey et al. 2007; Schmitz et al. 2015; Carroll
et al. 2017). Throughout, spatial concepts are essential for guiding effective strate-
gies for both biodiversity and ecosystem service conservation (Chan et al. 2006;
Moilanen and Wintle 2007).
Ecological concepts and analytical tools developed in the elds of landscape
ecology, geography, and spatial statistics are now commonly used in conservation so
that informed decisions about planning strategies and management can be made
(e.g., Moilanen et al. 2009). Indeed, most conservation planning and management
requires knowledge and the explicit spatial modeling of space and its major conse-
quences on species spatial variation and responses to global change. The inclusion of
space is therefore crucial when modeling species ecology and responses to a
changing world such as (1) species dispersal, (2) species interactions, (3) disturbance
dynamics, and (4) environmental change. Furthermore, as the eld of conservation
aims to provide better management recommendations to mitigate threats to biodi-
versity, implicit and explicit aspects of space need to be incorporated into applied
solutions such as restoration, species reintroductions, and maintaining connectivity
among habitat patches. In all these conservation applications the spatial scale of
implementation is key (Wiens 1989; Levin 1992,2000; McGarigal et al. 2016; Doak
et al. 1992; Fletcher et al. 2013; Gering et al. 2003).
1.4 The Growth of Frameworks for Spatial Modeling
Before modeling species dispersal, response to environmental conditions, and species
interactions, quantication of their spatial distribution is needed. This is why in
ecology and conservation the rst steps toward a better understanding and manage-
ment of biodiversity often consist of (1) mapping species distributions, and (2) quan-
tifying spatial patterns of both species distributions and environmental conditions
(Ferrier 2002; Gaston and Blackburn 2000; Guisan and Thuiller 2005). Once such
quantitative information is obtained, the next modeling steps frequently aim at relating
and modeling the responses of species to environmental conditions across space and/or
the species (intraspecic and interspecic) spatial interactions (Synes et al. 2017).
Modeling the processes that affect species distribution can be done using different
degrees of complexity in the analytical tools used. The level of complexity depends on
the processes modeled and ecological theories considered. Then, knowledge gaps about
6 1 Introduction to Spatial Ecology and Its Relevance for Conservation
species distribution can be gained by combining data on species behavior from empir-
ical studies and theoretical models of dispersal and related ows across space. Early
dispersal models set the stage for the development of ecological theories that embrace
space (Fig. 1.3), such as island biogeography (MacArthur and Wilson 1967), patch
dynamics (Pickett and White 1984), hierarchical theory (Wu and Loucks 1995;Allen
and Starr 1982), species coexistence (Chesson 2000), metapopulation (Hanski 1999),
metacommunity (Leibold and Chase 2017), and metaecosystem theory (Guichard
2017). Although these disciplines can be seen as separate elds, spatial ecology brings
them together through theory, models, and data analysis (Massol et al. 2011).
The emergence of modeling frameworks for spatial ecology was also fostered by
several technological advances ranging from the availability of aerial photographs,
remote sensing captors, and computing power. This allowed for conceptual and
modeling developments in spatial ecology to advance more realistic ways to repre-
sent and incorporate space into statistical and modeling approaches (Fig. 1.3).
Indeed, the ability to explicitly include the effects of space in ecological models
was also pivotal in the explosion of novel ecological questions and analytical ways
to address them over the last few decades.
Implicitly: Relative positions (i, j)
in a regular grid
Explicitly: x-y coordinates; Euclidean distance
between patches (black lines); effective
functional distance based on network topology
(grey lines)
Incorporating Space
i, j
Abiotic factors
generating spatial
in environmental
and human impacts,
such as habitat loss
and spread of invasives
Biotic processes
creating spatial
in distributions,
Ecological responses Environmental covariates
Spatial structure of
one species
Spatial structure of
several species
Fig. 1.3 How spatial processes affect species (response variables) and covariates (predictors), and
how space can be incorporated into models
1.4 The Growth of Frameworks for Spatial Modeling 7
The quantum leap in spatial ecology modeling frameworks involved considering
and incorporating space into modeling: implicitly (kernels, moving windows, rela-
tive topological position, etc.), explicitly (xy, diffusion, spread, individual/agent-
based models, etc.), and realistically (explicit network structure, spatial weights,
multiple spatial scales, etc.) (Fig. 1.3). It started by considering space as discrete
units. Such discretization of space opened a multitude of novel ways to model
ecological systems either in a spatially implicit fashion, where species occupancy
and abundance are modeled considering the effects of relative neighbors based on
grid topology (e.g., cellular automata models), or in a spatially explicit way, where
the actual Euclidean distances among cells (quadrats, pixels, sampling locations) are
used to model the spread of disturbance, disease, or species using dispersal kernels.
Then space was represented by the exact xycoordinates of each individual in a
given area such that the spatially explicit movement of individuals could be modeled
using individual/agent-based modeling approaches (Grimm et al. 2005; Matthews
et al. 2007). For example, this approach enabled modeling the dynamics and
succession of tree species at the tree-level using SORTIE (Pacala et al. 1996).
Using xycoordinates of individuals or sampling locations also allowed the spatially
explicit modeling of movement and connectivity while accounting for species
dispersal ability through spatially heterogeneous landscapes (Urban and Keitt
2001). Lastly, the spatially explicit representation of space permits us to model
processes acting over several spatial scales using meta-models (Urban 2005; Talluto
et al. 2016). The ability to model species and their responses to global change
explicitly in space opens the door to investigate the effects of the spatial legacy
(Wallin et al. 1994; James et al. 2007; Peterson 2002) of heterogeneity on ecological
processes and species persistence.
1.5 The Path Ahead
Spatial ecology and conservation has rapidly advanced over the past 20 years. With
an increasing emphasis on the use of spatial data and modeling to address both
fundamental and applied problems, the topic has matured. Spatial ecology embraces
spatial modeling and analysis, which is often applied to conservation issues.
In the remainder of this book, our path will be to provide an introduction to
several issues in spatial ecology and conservation, with an emphasis on spatial
modeling of applied ecological problems. We emphasize learning-by-doing, where
we illustrate these topics with real data and the application of spatial modeling to
these topics. We rst cover topics regarding the quantication of spatial pattern in
ecological data and we then focus more specically on topics regarding how species
respond to spatial pattern and its relevance for conservation (Table 1.2). We hope
that this coverage will deliver a strong foundation for students and professionals
alike to begin tackling ongoing issues of ecological and conservation importance.
8 1 Introduction to Spatial Ecology and Its Relevance for Conservation
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Table 1.2 Examples of spatial analytical methods (and book chapter(s) where they are presented)
used in spatial ecology and the quantitative components that they are estimating
Spatial analytical methods Spatial components addressed
Multiscale analysis (Chap. 2) Determine key spatial scales affecting the response variables
Categorical pattern analysis
(Chap. 3)
Quantify land-use and land-cover patterns
Spatial point processes
(Chap. 4)
Identifying the spatial pattern of points (events) and understand
the potential processes generating those patterns
Spatial-, geo-statistics
(Chap. 5)
Magnitude, range, and directionality of spatial variance
Spatial regressions (Chap. 6) Accounting for spatial structure of the response (spatial nuisance)
and independent variables (spatial contingency) in estimating
Species distribution models
(Chap. 7)
Interpolation, projections, and forecasting
Animal movement models
(Chaps. 8and 9)
Accounting for spatial heterogeneity and quantifying trajectories
Spatial network analysis
(Chap. 9)
Topological network, Euclidean, and functional distances
Spatial population dynamics
(Chap. 10)
Population dynamics accounting for spatial heterogeneity
Beta diversity (Chap. 11) Spatial species turnover
Spatial community analysis
(Chap. 11)
Spatial components of species interactions and environmental
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References 13
... To some extent metapopulation theory superseded the refuge design criteria in the 1990s (Hanski and Simberloff 1997), and it remains an active area of research in conservation biology and ecology (Hanski and Gaggiotti 2004;Begon and Townsend 2021). The equilibrium theory and metapopulation theory have together spawned research on metacommunities (e.g., Leibold and Miller 2004;Logue et al. 2011;Massol et al. 2011) and metaecosystems (e.g., Loreau et al. 2003;Gravel et al. 2011;Massol et al. 2011;Leibold and Chase 2017), key components in the rise of spatial ecology Fletcher and Fortin 2018). ...
... One of the most important aspects of animal behavior, particularly as it relates to wildlife ecology and conservation, is the way animals move and use space on a landscape. e eld of spatial ecology has grown rapidly over the last several decades as ecologists increasingly recognize the importance of scale, as well as the relationship between ecological processes and landscape composition and heterogeneity (Fletcher and Fortin 2018). As global environments become increasingly fragmented and dominated by anthropogenic drivers, the interaction between ecological systems and spatial heterogeneity becomes all the more important to understand and incorporate into ecological studies and biodiversity conservation e orts alike. ...
The Wood Turtle has experienced significant population declines across its range in the United States and Canada, where it is a species emblematic of cool, remote, clean rivers from Nova Scotia to Minnesota and south to Virginia. This richly illustrated book is the first solely dedicated to the natural history, ecology, and conservation of the Wood Turtle. More than 20 scientists and managers from across the species' range have collaborated in this volume to explore the Wood Turtle's evolution, landscape ecology, distribution, habitat, biology, and behavior, and to evaluate its conservation needs and outlook.
... Dans ce but, ce travail mobilisera différentes approches de l'écologie spatiale (c. à d., aspect de l'écologie où la dimension spatiale joue un rôle clé ; Fletcher & Fortin, 2018) pour améliorer Introduction les connaissances sur l'utilisation de l'espace d'espèces animales associées à des zoonoses préoccupantes pour la santé et l'économie publique. ...
Prévenir les risques d’épidémies est devenu un enjeu sanitaire et économique mondial, comme en témoigne l’émergence récente du SARS-COV-2. Cette Thèse vise à améliorer les connaissances sur l’utilisation de l’espace des chauves-souris frugivores (Pteropodidae) dans des environnements modifiés par l’homme. Ce travail mobilise des données de télémétrie satellitaire chez (i) la roussette de Lyle (Pteropus lylei), espèce réservoir du virus Nipah en Asie, et (ii) la chauve-souris à tête de marteau (Hypsignathus monstrosus), impliquée dans la circulation du virus Ebola en Afrique. La population étudiée de roussette de Lyle était déjà connue pour se nourrir préférentiellement dans les zones résidentielles d’un environnement fragmenté au Cambodge. La chauve-souris à tête de marteau, dont l’utilisation des habitats était méconnue, a été étudiée dans une région forestière en République du Congo – épicentre d’épidémies humaine d’Ebola en 2001–2005. De plus, des données de captures directes de chauves-souris ont été collectées dans cette dernière région. Il ressort de ces travaux que la chauve-souris à tête de marteau se nourrit préférentiellement dans les terres agricoles qui entourent les petits villages forestiers. Les individus de roussette de Lyle visitent davantage d’aires d’alimentation dans l’habitat préférentiel durant la nuit, tandis que les chauves-souris à tête de marteau y passent plus de temps sans multiplier le nombre d’aires visitées. Ces deux espèces bénéficient ainsi des ressources anthropiques à l’échelle de la population selon deux stratégies de déplacements individuels, qui sont possiblement ajustées selon le degré de fragmentation de l’environnement. Chez la chauve-souris à tête de marteau, les aires d’alimentation dans la forêt sont délaissées par les individus qui restent longtemps dans le site d’accouplement durant la nuit, ce qui suggère un rôle des terres agricoles dans l’établissement et le maintien des sites d’accouplement. Au cours de nuits successives, les deux espèces revisitent davantage une aire d’alimentation lorsqu’elles y ont passé beaucoup de temps lors de leur dernière visite. Par ailleurs, une communauté de sept espèces de chauves-souris frugivores a été identifiée dans la région étudiée en Afrique. La probabilité d’occurrence de quatre espèces était plus importante dans les villages, tandis que les autres espèces n’étaient pas influencées par l’habitat. L’ensemble de ces travaux fournit de nouvelles informations sur l’utilisation de l’espace des chauves-souris frugivores dans le cadre de leurs activités nocturnes d’alimentation et de reproduction. Ces données pourraient être intégrées dans des modélisations épidémiologiques visant à mieux comprendre les interactions entre les chauves-souris frugivores, les humains ou les animaux domestiques, ainsi que les voies de transmission de pathogènes.
... The combination of ecological exogenous and endogenous processes determines spatial characteristics of a species or population, such as, movement and dispersal, resource acquisition, and intra and interspecies interactions (Fletcher and Fortin 2018). Aided by advanced technology and data analysis methodologies, research on the spatial ecology of jaguars has already yielded valuable information on movement and spatial patterns, which are key to arresting habitat loss and, therefore the fragmentation and subsequent decline in biodiversity. ...
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ContextMovement ecology contributes valuable information about animal interactions with the environment, and their responses to landscape-level anthropogenic impacts. Big cats are vulnerable to such changes, but the current deficit of information about home range movements, limits the scope of conservation initiatives. Objectives Describe the home range size, interactions, and differences between jaguar populations across its distribution in MexicoMethods We used 41,008 GPS-generated data points obtained from 28 tagged jaguars ( Panthera onca ) in five different states of Mexico over an 18-year period to describe home range size, differences between male and female territories, interactions in overlapping territories, and territory differences among populations. ResultsOur data shows that jaguar home range is smaller than tiger’s but larger than leopard’s. Male mean home range size (285.28 km ² , n=13) tends to be larger than that of females (152.2 km ² , n = 15), the difference was not statistically significant. While the home range for at least one male was 633.44 km ² , contrasted with the much smaller 48.89 km ² for some female jaguars. Data of overlapping ranges showed 34.71% of female territory overlaps male territory, 32.46% of female territory is shared with other females, 18.97% of male territory is shared with other males, and only 16.89% male territory overlaps with female territory. Conclusions The absence of significant differences in home range sizes among the habitats suggest jaguar territory is not highly dependent on the type of habitat it occupies. Our findings of the spatial parameters of jaguar movements can be applied to identifying ecological corridors and the design of protected areas for this species.
... One of the fundamental interests in ecology is understanding which factors and processes drive species' occurrence in space. This knowledge stands as the basis of several research lines and supports the conservation and management of species and habitats worldwide [1]. Many studies made use of both the ecological niche theory and habitat suitability modelling to predict and map species distributions in relation to biotic (e.g., vegetation type or land use) and abiotic (e.g., climate) factors [2]. ...
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One of the fundamental interests in ecology is understanding which factors drive species’ distribution. We aimed to understand the drivers of bat distribution and co-occurrence patterns in a remote, insular system. The two bat species known to occur in the Azores archipelago were used as a model. Echolocation calls were recorded at 414 point-locations haphazardly distributed across the archipelago. Calls were analysed and assigned to each species. Binominal generalised linear models were adjusted using different descriptors at two scales: archipelago and island. The presence of the co-occurring species was included at both scales. The results show that island isolation, habitat and climate play an essential role on the archipelago and island scales, respectively. However, the positive interaction between bat species was the most critical driver of species’ distribution at the island scale. This high co-occurrence pattern at the island scale may result from both species’ maximising foraging profit in a region where prey abundance may be highly variable. However, further research is necessary to clarify the mechanisms behind this positive interaction. Both species are threatened and lack specific management and protection measures. Maintaining this positive interaction between the two species may prove to be fundamental for their conservation.
... In this context, attaining knowledge of the red deer's spatial ecology is crucial if effective measures with which to manage populations and mitigate human-wildlife conflicts are to be designed (Apollonio et al., 2017). The study of spatial ecology makes it possible to understand key ecological processes (e.g., activity, population dynamics, inter-species interactions, dispersal…) that operate on a spatial scale and that, in turn, affect key ecological patterns, such as species distribution and population abundance (Fletcher and Fortin, 2018). These patterns are far from being general, as land use, periods of the year, hunting management and their combined effects are assumed to determine spatial behaviour (e.g., Gallego-Zamorano et al., 2020), as has previously been observed in red deer in central and northern Europe (Kamler et al., 2007;Náhlik et al., 2009;Allen et al., 2014). ...
The knowledge regarding the spatial ecology of red deer (Cervus elaphus) in different environments is crucial if effective management actions are to be designed. However, this knowledge continues to be scarce in the complex contexts of mixed land use and management circumstances. This study describes the spatial ecology of red deer monitored using GPS collars in Mediterranean ecosystems of South-Central Spain, considering the effect of individual and seasonal (food shortage period, rut, hunting season and food abundance period) factors on different land use and management scenarios, namely protected areas, mixed farms and fenced hunting estates. Our results showed less activity (ACT), a shorter daily range (DR) and a smaller home range (HR) during the food shortage period: ACT: 0.38 ± (SD) 0.12; DR: 3010.9 ± 727.3 m; and weekly HR: 122.2 ± 59.6 ha. With regard to land use, individuals were less ACT and had a smaller DR on fenced hunting estates (ACT: 0.24 ± 0.12; DR: 1946.3 ± 706.7 m) than in protected areas (ACT: 0.59 ± 0.12; DR: 4071.4 ± 1068.2 m) or on mixed farms (ACT: 0.57 ± 0.29; DR: 5431.1 ± 1939.5 m) in all the periods studied. Red deer selected land cover with forage and shelter when foraging and resting, respectively. When drive hunt events occurred (mixed farms and fenced hunting estates), the deer were more prone to select safer habitats (scrublands) and avoid open areas (crops or grasslands) than were their counterparts in protected areas. The patterns observed can be explained by sexual and seasonal differences as regards requirements, the response to disturbances and, interestingly, population management. Our results provide useful information with which to design scientifically-based species adaptive management in response to relevant and timely situations in Europe, such as the potential transmission of shared infections, vehicle collisions, and damage to crops and ecosystems.
... Nowadays ecology is widely considered not only a natural scientific discipline, a subdivision of biology, but also a point of view and perspective into the problems derived from the interrelationships between human society and its natural environment. Spatial ecology grew in importance recently as a subdiscipline that focuses specifically on the study and modeling of the roles of space on ecological processes that in turn affects ecological patterns (Fletcher & Fortin, 2018). The development and maturation of the subdiscipline of spatial ecology in the last few decades was conditioned by the growing availability of spatial data, constantly increased computing capacities, as well as the advances in spatial modeling techniques. ...
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The main concepts and methods of ecological geomorphometry as a research field aimed at studying relationships between terrain morphometric characteristics and ecological factors and processes are reviewed in the paper. The progress in this research field has been conditioned by the propagation of high-resolution digital elevation models in free access and of methods of their digital analysis, namely: the calculation of derivative quantitative characteristics (attributes and indices) of terrain and the statistical models of analyzing the relationships between the latter and the ecological properties and factors (those relevant for a certain ecological subject). A peculiar feature of ecological approach to regarding terrain (its morphology) is subjectcentrism (that is, regarding it from a point of view of a certain subject). The subject of ecological relationships can be living entities (populations, species, communities) as well as a human, social entities, economy and its branches. Three main concepts of ecological geomorphometry are put forward: terrain attributes (relatively simple quantitative characteristics of terrain form that characterize its geometry and some elementary physical processes); topographic indices (quantitative surrogates for some complex physical or biophysical processes of ecological significance); morphotops (spatial units that are distinguished by terrain morphology, using criteria of ecological homogeneity relevant from a viewpoint of a certain ecological subject). Morphotops can be distinguished with different level of detail (and, as a result, with different characteristic dimensions), relative to the study aim, to the geographic features of the area, and to the available data and the methods of their analysis. While morphotops are distinguished with strictly defined quantitative morphometric parameters (terrain attributes, topographic indices), this enables using formalized methods with their advantages of reproducibility and possibility of automatizing. In our studies aimed at morphotop mapping for a small area in the hilly terrain of Davydiv range near Lviv and for a larger area in the central part of Ukrainian Carpathians, morphotops delineation was based on topographic indices that characterize insolation level (solar radiation incidence on terrain elements of different aspect and slope values), lateral redistribution of water on slopes and redistribution of solid matter by washout on slopes. Morphotops were distinguished with cluster analysis method, which allows to distinguish natural groupings of data in the attribute space. Presetting different number of clusters to be distinguished, morphotopes can be distinguished with different levels of detail, larger number of clusters corresponding to more homogenous morphotops with smaller characteristic sizes. Key words: ecological geomorphology, ecological geomorphometry, morphotops, terrain attributes, topographic indices.
... Spatial ecology aims to uncover the causes and consequences of organism distribution and its changes over space and time [204,205]. Special attention has been devoted to biological invasions due to their significant adverse ecological and economic effects. However, there is no single tool or unified approach to understand and predict invasion impacts [206,207]. ...
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Biological invasions represent some of the most severe threats to local communities and ecosystems. Among invasive species, the vector-borne pathogen Xylella fastidiosa is responsible for a wide variety of plant diseases and has profound environmental, social and economic impacts. Once restricted to the Americas, it has recently invaded Europe, where multiple dramatic outbreaks have highlighted critical challenges for its management. Here, we review the most recent advances on the identification, distribution and management of X. fastidiosa and its insect vectors in Europe through genetic and spatial ecology methodologies. We underline the most important theoretical and technological gaps that remain to be bridged. Challenges and future research directions are discussed in the light of improving our understanding of this invasive species, its vectors and host-pathogen interactions. We highlight the need of including different, complimentary outlooks in integrated frameworks to substantially improve our knowledge on invasive processes and optimize resources allocation. We provide an overview of genetic, spatial ecology and integrated approaches that will aid successful and sustainable management of one of the most dangerous threats to European agriculture and ecosystems.
Nowadays the process of global urbanization is unstoppable, leading to a serious threat to local biodiversity. Urbanization may result in biodiversity decline or even species extinction, while sometimes help maintain species abundance in some developed countries. Different land-cover and land-use types affect species diversity in different aspects and directions, so it’s important to understand the pattern of species distribution across different characteristics of urban landscape, which helps city-designers and decision-makers to mitigate detrimental influences of urbanization on local biodiversity by rational urban planning and effective conservation protection. This study uses bird, which are highly sensitive to environmental changes, as the ecological indicators.This paper studies the differences of species richness, abundance and community composition from five urban land-cover and seven land-use types, and analyses patterns of bird distribution in different land use purposes on the same land cover landscape. This study used bird species richness, Shannon and Simpson diversity index across Greater Manchester to evaluate bird diversity. This study also used Generalized Linear Model to model the relationship between bird species richness and land-cover or land-use density, and used Redundancy Analysis (RDA) to interpret the response of bird communities to land-cover and land-use density. Green spaces (especially for public parks land use) and water bodies have relatively higher bird species richness. Built areas have the lowest species richness, especially the institutional land use (including religious grounds, school grounds, and institutional grounds). Considering different land-use purposes, public parks and recreation have the highest bird diversity in green spaces land-cover, followed by amenity land and domestic gardens. In built-up areas, species diversity in institutional land use is higher than previously developed land use. Clear understanding the relationships between land-cover and land-use types and bird species diversity and communities composition will help better policy making for potential future land-cover and land-use planning.KeywordsSpatial ecologyAvianGreater ManchesterSpecies richnessLand coverLand use
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Knowledge about spatial distribution of owl species is important for inferring species coexistence mechanisms. In the present study, we explore spatial patterns of distribution and habitat selection of four owl species-Eurasian pygmy owl (Glaucidium passerinum), boreal owl (Aegolius funereus), tawny owl (Strix aluco) and Ural owl (Strix uralensis)-ranging in body mass from 50 g to 1300 g, with sympatric occurrence in temperate continuous montane forests in the Veľká Fatra Mts., Western Carpathians, central Slovakia. Locations of hooting owl males were surveyed between 2009-2015 in an area of 317 km 2. Spatial point pattern analysis was used for analysis of owl distribution. Random patterns of owls' spatial arrangement dominate at both intra and interspecific levels within the studied area. Only intraspecific distribution of pygmy owls and interspecific distribution of Ural owls toward tawny owls exhibited positive associations. This discrepancy with other studies can be explained in terms of pygmy owls' preference for highquality nest sites and/or spatial clustering in their prey distribution, and due to aggressive behaviour of dominant Ural owls toward subdominant tawny owls, respectively. Moreover, we found considerable overlap in habitat preferences between owl species, considering stand age, stand height, tree species richness, distance to open area, elevation , slope, percentage of coniferous tree species and position on hillslope, although pygmy owls were not registered in pure broadleaved stands, Ural owls were not registered in pure coniferous stands, and boreal and Ural owls were more common on slope summits and shoulders than tawny and pygmy owls. The observed patterns of spatial arrangement might suggest developed coexistence mechanisms in these owl species; differences between studies may indicate complex interactions between intra and interspecific associations and habitat quality and quantity, food availability and owl species involved in those interactions on a landscape scale.
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Human-mediated transport beyond biogeographic barriers has led to the introduction and establishment of alien species in new regions worldwide. However, we lack a global picture of established alien species richness for multiple taxonomic groups. Here, we assess global patterns and potential drivers of established alien species richness across eight taxonomic groups (amphibians, ants, birds, freshwater fishes, mammals, vascular plants, reptiles and spiders) for 186 islands and 423 mainland regions. Hotspots of established alien species richness are predominantly island and coastal mainland regions. Regions with greater gross domestic product per capita, human population density, and area have higher established alien richness, with strongest effects emerging for islands. Ants and reptiles, birds and mammals, and vascular plants and spiders form pairs of taxonomic groups with the highest spatial congruence in established alien richness, but drivers explaining richness differ between the taxa in each pair. Across all taxonomic groups, our results highlight the need to prioritize prevention of further alien species introductions to island and coastal mainland regions globally.
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Metacommunity theory has provided many insights into the general problem of local versus regional control of species diversity and relative abundance. The metacommunity framework has been extended from competitive interactions to whole food webs that can be described as spatial networks of interaction networks. Trophic metacommunity theory greatly contributed to resolving the community complexity-stability debate by predicting its dependence on the regional spatial context. The meta-ecosystem framework has since been suggested as a useful simplification of complex ecosystems to apply this spatial context to spatial flows of both individuals and matter. Reviewing the recent literature on metacommunity and meta-ecosystem theories suggests the importance of unifying theories of interaction strength into a meta-ecosystem framework that captures how the strength of spatial, species, and ecosystem fluxes are distributed across location and trophic levels. Such integration predicts important feedback between local and regional processes that drive the assembly of species, the stability of community, and the emergence of ecosystem functions, from limited spatial fluxes of individuals and (in)organic matter. These predictions are often mediated by the maintenance of environmental or endogenous fluctuations from local to regional scales that create important challenges and opportunities for the validation of metacommunity and meta-ecosystem theories and their application to conservation.
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As most regions of the earth transition to altered climatic conditions, new methods are needed to identify refugia and other areas whose conservation would facilitate persistence of biodiversity under climate change. We compared several common approaches to conservation planning focused on climate resilience over a broad range of ecological settings across North America and evaluated how commonalities in the priority areas identified by different methods varied with regional context and spatial scale. Our results indicate that priority areas based on different environmental diversity metrics differed substantially from each other and from priorities based on spatiotemporal metrics such as climatic velocity. Refugia identified by diversity or velocity metrics were not strongly associated with the current protected area system, suggesting the need for additional conservation measures including protection of refugia. Despite the inherent uncertainties in predicting future climate, we found that variation among climatic velocities derived from different general circulation models and emissions pathways was less than the variation among the suite of environmental diversity metrics. To address uncertainty created by this variation, planners can combine priorities identified by alternative metrics at a single resolution and downweight areas of high variation between metrics. Alternately, coarse-resolution velocity metrics can be combined with fine-resolution diversity metrics in order to leverage the respective strengths of the two groups of metrics as tools for identification of potential macro- and micro-refugia that in combination maximize both transient and long-term resilience to climate change. Planners should compare and integrate approaches that span a range of model complexity and spatial scale to match the range of ecological and physical processes influencing persistence of biodiversity and identify a conservation network resilient to threats operating at multiple scales. This article is protected by copyright. All rights reserved.
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Landscape ecological modelling provides a vital means for understanding the interactions between geographical, climatic, and socio-economic drivers of land-use and the dynamics of ecological systems. This growing field is playing an increasing role in informing landscape spatial planning and management. Here, we review the key modelling approaches that are used in landscape modelling and in ecological modelling. We identify an emerging theme of increasingly detailed representation of process in both landscape and ecological modelling, with complementary suites of modelling approaches ranging from correlative, through aggregated process based approaches to models with much greater structural realismthat often represent behaviours at the level of agents or individuals. We provide examples of the considerable progress that has been made at the intersection of landscape modelling and ecological modelling, while also highlighting that the majority of this work has to date exploited a relatively small number of the possible combinations of model types from each discipline. We use this review to identify key gaps in existing landscape ecological modelling effort and highlight emerging opportunities, in particular for future work to progress in novel directions by combining classes of landscape models and ecological models that have rarely been used together.
Ecologists are aware of the importance of natural dynamics in ecosystems. Historically, the focus has been on the development in succession of equilibrium communities, which has generated an understanding of the composition and functioning of ecosystems. Recently, many have focused on the processes of disturbances and the evolutionary significance of such events. This shifted emphasis has inspired studies in diverse systems. The phrase "patch dynamics" (Thompson, 1978) describes their common focus. The Ecology of Natural Disturbance and Patch Dynamics brings together the findings and ideas of those studying varied systems, presenting a synthesis of diverse individual contributions.
Recent work linking community structure and ecosystem function has primarily focused on the effects of local species richness but has neglected the dispersal-dependent processes of community assembly that are ultimately involved in determining community structure and its relation to ecosystems. Here we combine simple consumer-resource competition models and metacommunity theory with discussion of case studies to outline how spatial processes within metacommunities can alter community assembly and modify expectations about how species diversity and composition influence ecosystem attributes at local scales. We argue that when community assembly is strongly limited by dispersal, this can constrain ecosystem functioning by reducing positive selection effects (reducing the probability of the most productive species becoming dominant) even though it may often also enhance complementarity (favoring combinations of species that enhance production even though they may not individually be most productive). Conversely, excess dispersal with strong source-sink relations among heterogeneous habitats can reduce ecosystem functioning by swamping local filters that would normally favor better-suited species. Ecosystem function is thus most likely maximized at intermediate levels of dispersal where both of these effects are minimized. In this scenario, we find that the selection effect is maximized, while complementarity is often reduced and local diversity may often be relatively low. Our synthesis emphasizes that it is the entire set of community assembly processes that affect the functioning of ecosystems, not just the part that determines local species richness. This article is protected by copyright. All rights reserved.