Landscape ecology

Landscape ecology is the science of studying and improving relationships between ecological processes in the environment and particular ecosystems. This is done within a variety of landscape scales, development spatial patterns, and organizational levels of research and policy.[1][2][3]

As a highly interdisciplinary field in systems science, landscape ecology integrates biophysical and analytical approaches with humanistic and holistic perspectives across the natural sciences and social sciences. Landscapes are spatially heterogeneous geographic areas characterized by diverse interacting patches or ecosystems, ranging from relatively natural terrestrial and aquatic systems such as forests, grasslands, and lakes to human-dominated environments including agricultural and urban settings.[2][4][5] The most salient characteristics of landscape ecology are its emphasis on the relationship among pattern, process and scale, and its focus on broad-scale ecological and environmental issues. These necessitate the coupling between biophysical and socioeconomic sciences. Key research topics in landscape ecology include ecological flows in landscape mosaics, land use and land cover change, scaling, relating landscape pattern analysis with ecological processes, and landscape conservation and sustainability.[6]

NLCD landcover MSN area
Land cover surrounding Madison, WI. Fields are colored yellow and brown, water is colored blue, and urban surfaces are colored red.
NLCD impervious MSN area
Impervious surfaces surrounding Madison, WI
NLCD canopy MSN area
Canopy cover surrounding Madison, WI


The German term Landschaftsökologie–thus landscape ecology–was coined by German geographer Carl Troll in 1939.[7] He developed this terminology and many early concepts of landscape ecology as part of his early work, which consisted of applying aerial photograph interpretation to studies of interactions between environment and vegetation.


Heterogeneity is the measure of how parts of a landscape differ from one another. Landscape ecology looks at how this spatial structure affects organism abundance at the landscape level, as well as the behavior and functioning of the landscape as a whole. This includes studying the influence of pattern, or the internal order of a landscape, on process, or the continuous operation of functions of organisms.[8] Landscape ecology also includes geomorphology as applied to the design and architecture of landscapes.[9] Geomorphology is the study of how geological formations are responsible for the structure of a landscape.


Evolution of theory

One central landscape ecology theory originated from MacArthur & Wilson's The Theory of Island Biogeography. This work considered the biodiversity on islands as the result of competing forces of colonization from a mainland stock and stochastic extinction. The concepts of island biogeography were generalized from physical islands to abstract patches of habitat by Levins' metapopulation model (which can be applied e.g. to forest islands in the agricultural landscape[10]). This generalization spurred the growth of landscape ecology by providing conservation biologists a new tool to assess how habitat fragmentation affects population viability. Recent growth of landscape ecology owes much to the development of geographic information systems (GIS)[11] and the availability of large-extent habitat data (e.g. remotely sensed datasets).

Development as a discipline

Landscape ecology developed in Europe from historical planning on human-dominated landscapes. Concepts from general ecology theory were integrated in North America. While general ecology theory and its sub-disciplines focused on the study of more homogenous, discrete community units organized in a hierarchical structure (typically as ecosystems, populations, species, and communities), landscape ecology built upon heterogeneity in space and time. It frequently included human-caused landscape changes in theory and application of concepts.[12]

By 1980, landscape ecology was a discrete, established discipline. It was marked by the organization of the International Association for Landscape Ecology (IALE) in 1982. Landmark book publications defined the scope and goals of the discipline, including Naveh and Lieberman[13] and Forman and Godron.[14][15] Forman[5] wrote that although study of "the ecology of spatial configuration at the human scale" was barely a decade old, there was strong potential for theory development and application of the conceptual framework. Today, theory and application of landscape ecology continues to develop through a need for innovative applications in a changing landscape and environment. Landscape ecology relies on advanced technologies such as remote sensing, GIS, and models. There has been associated development of powerful quantitative methods to examine the interactions of patterns and processes.[4] An example would be determining the amount of carbon present in the soil based on landform over a landscape, derived from GIS maps, vegetation types, and rainfall data for a region. Remote sensing work has been used to extend landscape ecology to the field of predictive vegetation mapping, for instance by Janet Franklin.

Definitions/conceptions of landscape ecology

Nowadays, at least six different conceptions of landscape ecology can be identified: one group tending toward the more disciplinary concept of ecology (subdiscipline of biology; in conceptions 2, 3, and 4) and another group—characterized by the interdisciplinary study of relations between human societies and their environment—inclined toward the integrated view of geography (in conceptions 1, 5, and 6):[16]

  1. Interdisciplinary analysis of subjectively defined landscape units (e.g. Neef School[17]): Landscapes are defined in terms of uniformity in land use. Landscape ecology explores the landscape's natural potential in terms of functional utility for human societies. To analyse this potential, it is necessary to draw on several natural sciences.
  2. Topological ecology at the landscape scale (e.g. Forman & Godron[18]): 'Landscape' is defined as a heterogeneous land area composed of a cluster of interacting ecosystems (woods, meadows, marshes, villages, etc.) that is repeated in similar form throughout. It is explicitly stated that landscapes are areas at a kilometres wide human scale of perception, modification, etc. Landscape ecology describes and explains the landscapes' characteristic patterns of ecosystems and investigates the flux of energy, mineral nutrients, and species among their component ecosystems, providing important knowledge for addressing land-use issues.
  3. Organism-centred, multi-scale topological ecology (e.g. John A. Wiens[19]): Explicitly rejecting views expounded by Troll, Zonneveld, Naveh, Forman & Godron, etc., landscape and landscape ecology are defined independently of human perceptions, interests, and modifications of nature. 'Landscape' is defined – regardless of scale – as the 'template' on which spatial patterns influence ecological processes. Not humans, but rather the respective species being studied is the point of reference for what constitutes a landscape.
  4. Topological ecology at the landscape level of biological organisation (e.g. Urban et al.[20]): On the basis of ecological hierarchy theory, it is presupposed that nature is working at multiple scales and has different levels of organisation which are part of a rate-structured, nested hierarchy. Specifically, it is claimed that, above the ecosystem level, a landscape level exists which is generated and identifiable by high interaction intensity between ecosystems, a specific interaction frequency and, typically, a corresponding spatial scale. Landscape ecology is defined as ecology that focuses on the influence exerted by spatial and temporal patterns on the organisation of, and interaction among, functionally integrated multispecies ecosystems.
  5. Analysis of social-ecological systems using the natural and social sciences and humanities (e.g. Leser;[21] Naveh;[22] Zonneveld[23]): Landscape ecology is defined as an interdisciplinary super-science that explores the relationship between human societies and their specific environment, making use of not only various natural sciences, but also social sciences and humanities. This conception is grounded in the assumption that social systems are linked to their specific ambient ecological system in such a way that both systems together form a co-evolutionary, self-organising unity called 'landscape'. Societies' cultural, social and economic dimensions are regarded as an integral part of the global ecological hierarchy, and landscapes are claimed to be the manifest systems of the 'Total Human Ecosystem' (Naveh) which encompasses both the physical ('geospheric') and mental ('noospheric') spheres.
  6. Ecology guided by cultural meanings of lifeworldly landscapes (frequently pursued in practice[24] but not defined, but see, e.g., Hard;[25] Trepl[26]): Landscape ecology is defined as ecology that is guided by an external aim, namely, to maintain and develop lifeworldly landscapes. It provides the ecological knowledge necessary to achieve these goals. It investigates how to sustain and develop those populations and ecosystems which (i) are the material 'vehicles' of lifeworldly, aesthetic and symbolic landscapes and, at the same time, (ii) meet societies' functional requirements, including provisioning, regulating, and supporting ecosystem services. Thus landscape ecology is concerned mainly with the populations and ecosystems which have resulted from traditional, regionally specific forms of land use.

Relationship to ecological theory

Some research programmes of landscape ecology theory, namely those standing in the European tradition, may be slightly outside of the "classical and preferred domain of scientific disciplines" because of the large, heterogeneous areas of study. However, general ecology theory is central to landscape ecology theory in many aspects. Landscape ecology consists of four main principles: the development and dynamics of spatial heterogeneity, interactions and exchanges across heterogeneous landscapes, influences of spatial heterogeneity on biotic and abiotic processes, and the management of spatial heterogeneity. The main difference from traditional ecological studies, which frequently assume that systems are spatially homogenous, is the consideration of spatial patterns.[27]

Important terms

Landscape ecology not only created new terms, but also incorporated existing ecological terms in new ways. Many of the terms used in landscape ecology are as interconnected and interrelated as the discipline itself.


Certainly, 'landscape' is a central concept in landscape ecology. It is, however, defined in quite different ways. For example:[28] Carl Troll conceives of landscape not as a mental construct but as an objectively given 'organic entity', a harmonic individuum of space.[29] Ernst Neef[30] defines landscapes as sections within the uninterrupted earth-wide interconnection of geofactors which are defined as such on the basis of their uniformity in terms of a specific land use, and are thus defined in an anthropocentric and relativistic way. According to Richard Forman and Michael Godron,[31] a landscape is a heterogeneous land area composed of a cluster of interacting ecosystems that is repeated in similar form throughout, whereby they list woods, meadows, marshes and villages as examples of a landscape's ecosystems, and state that a landscape is an area at least a few kilometres wide. John A. Wiens[32] opposes the traditional view expounded by Carl Troll, Isaak S. Zonneveld, Zev Naveh, Richard T. T. Forman/Michel Godron and others that landscapes are arenas in which humans interact with their environments on a kilometre-wide scale; instead, he defines 'landscape'—regardless of scale—as "the template on which spatial patterns influence ecological processes".[33] Some define 'landscape' as an area containing two or more ecosystems in close proximity.[12]

Scale and heterogeneity (incorporating composition, structure, and function)

A main concept in landscape ecology is scale. Scale represents the real world as translated onto a map, relating distance on a map image and the corresponding distance on earth.[34] Scale is also the spatial or temporal measure of an object or a process,[27] or amount of spatial resolution.[5] Components of scale include composition, structure, and function, which are all important ecological concepts. Applied to landscape ecology, composition refers to the number of patch types (see below) represented on a landscape and their relative abundance. For example, the amount of forest or wetland, the length of forest edge, or the density of roads can be aspects of landscape composition. Structure is determined by the composition, the configuration, and the proportion of different patches across the landscape, while function refers to how each element in the landscape interacts based on its life cycle events.[27] Pattern is the term for the contents and internal order of a heterogeneous area of land.[14]

A landscape with structure and pattern implies that it has spatial heterogeneity, or the uneven distribution of objects across the landscape.[5] Heterogeneity is a key element of landscape ecology that separates this discipline from other branches of ecology. Landscape heterogeneity is able to quantify with agent-based methods as well.[35]

Patch and mosaic

Patch, a term fundamental to landscape ecology, is defined as a relatively homogeneous area that differs from its surroundings.[5] Patches are the basic unit of the landscape that change and fluctuate, a process called patch dynamics. Patches have a definite shape and spatial configuration, and can be described compositionally by internal variables such as number of trees, number of tree species, height of trees, or other similar measurements.[5]

Matrix is the "background ecological system" of a landscape with a high degree of connectivity. Connectivity is the measure of how connected or spatially continuous a corridor, network, or matrix is.[5] For example, a forested landscape (matrix) with fewer gaps in forest cover (open patches) will have higher connectivity. Corridors have important functions as strips of a particular type of landscape differing from adjacent land on both sides.[5] A network is an interconnected system of corridors while mosaic describes the pattern of patches, corridors, and matrix that form a landscape in its entirety.[5]

Boundary and edge

Landscape patches have a boundary between them which can be defined or fuzzy.[12] The zone composed of the edges of adjacent ecosystems is the boundary.[5] Edge means the portion of an ecosystem near its perimeter, where influences of the adjacent patches can cause an environmental difference between the interior of the patch and its edge. This edge effect includes a distinctive species composition or abundance.[5] For example, when a landscape is a mosaic of perceptibly different types, such as a forest adjacent to a grassland, the edge is the location where the two types adjoin. In a continuous landscape, such as a forest giving way to open woodland, the exact edge location is fuzzy and is sometimes determined by a local gradient exceeding a threshold, such as the point where the tree cover falls below thirty-five percent.[27]

Ecotones, ecoclines, and ecotopes

A type of boundary is the ecotone, or the transitional zone between two communities.[9] Ecotones can arise naturally, such as a lakeshore, or can be human-created, such as a cleared agricultural field from a forest.[9] The ecotonal community retains characteristics of each bordering community and often contains species not found in the adjacent communities. Classic examples of ecotones include fencerows, forest to marshlands transitions, forest to grassland transitions, or land-water interfaces such as riparian zones in forests. Characteristics of ecotones include vegetational sharpness, physiognomic change, occurrence of a spatial community mosaic, many exotic species, ecotonal species, spatial mass effect, and species richness higher or lower than either side of the ecotone.[36]

An ecocline is another type of landscape boundary, but it is a gradual and continuous change in environmental conditions of an ecosystem or community. Ecoclines help explain the distribution and diversity of organisms within a landscape because certain organisms survive better under certain conditions, which change along the ecocline. They contain heterogeneous communities which are considered more environmentally stable than those of ecotones.[37]

An ecotope is a spatial term representing the smallest ecologically distinct unit in mapping and classification of landscapes.[5] Relatively homogeneous, they are spatially explicit landscape units used to stratify landscapes into ecologically distinct features. They are useful for the measurement and mapping of landscape structure, function, and change over time, and to examine the effects of disturbance and fragmentation.

Disturbance and fragmentation

Disturbance is an event that significantly alters the pattern of variation in the structure or function of a system. Fragmentation is the breaking up of a habitat, ecosystem, or land-use type into smaller parcels.[5] Disturbance is generally considered a natural process. Fragmentation causes land transformation, an important process in landscapes as development occurs.

An important consequence of repeated, random clearing (whether by natural disturbance or human activity) is that contiguous cover can break down into isolated patches. This happens when the area cleared exceed a critical level, which means that landscapes exhibit two phases: connected and disconnected.[38]


Landscape ecology theory stresses the role of human impacts on landscape structures and functions. It also proposes ways for restoring degraded landscapes.[13] Landscape ecology explicitly includes humans as entities that cause functional changes on the landscape.[12] Landscape ecology theory includes the landscape stability principle, which emphasizes the importance of landscape structural heterogeneity in developing resistance to disturbances, recovery from disturbances, and promoting total system stability.[14] This principle is a major contribution to general ecological theories which highlight the importance of relationships among the various components of the landscape. Integrity of landscape components helps maintain resistance to external threats, including development and land transformation by human activity.[4] Analysis of land use change has included a strongly geographical approach which has led to the acceptance of the idea of multifunctional properties of landscapes.[15] There are still calls for a more unified theory of landscape ecology due to differences in professional opinion among ecologists and its interdisciplinary approach (Bastian 2001).

An important related theory is hierarchy theory, which refers to how systems of discrete functional elements operate when linked at two or more scales. For example, a forested landscape might be hierarchically composed of drainage basins, which in turn are composed of local ecosystems, which are in turn composed of individual trees and gaps.[5] Recent theoretical developments in landscape ecology have emphasized the relationship between pattern and process, as well as the effect that changes in spatial scale has on the potential to extrapolate information across scales.[27] Several studies suggest that the landscape has critical thresholds at which ecological processes will show dramatic changes, such as the complete transformation of a landscape by an invasive species due to small changes in temperature characteristics which favor the invasive's habitat requirements.[27]


Research directions

Developments in landscape ecology illustrate the important relationships between spatial patterns and ecological processes. These developments incorporate quantitative methods that link spatial patterns and ecological processes at broad spatial and temporal scales. This linkage of time, space, and environmental change can assist managers in applying plans to solve environmental problems.[4] The increased attention in recent years on spatial dynamics has highlighted the need for new quantitative methods that can analyze patterns, determine the importance of spatially explicit processes, and develop reliable models.[27] Multivariate analysis techniques are frequently used to examine landscape level vegetation patterns. Studies use statistical techniques, such as cluster analysis, canonical correspondence analysis (CCA), or detrended correspondence analysis (DCA), for classifying vegetation. Gradient analysis is another way to determine the vegetation structure across a landscape or to help delineate critical wetland habitat for conservation or mitigation purposes (Choesin and Boerner 2002).[39]

Climate change is another major component in structuring current research in landscape ecology.[40] Ecotones, as a basic unit in landscape studies, may have significance for management under climate change scenarios, since change effects are likely to be seen at ecotones first because of the unstable nature of a fringe habitat.[36] Research in northern regions has examined landscape ecological processes, such as the accumulation of snow, melting, freeze-thaw action, percolation, soil moisture variation, and temperature regimes through long-term measurements in Norway.[41] The study analyzes gradients across space and time between ecosystems of the central high mountains to determine relationships between distribution patterns of animals in their environment. Looking at where animals live, and how vegetation shifts over time, may provide insight into changes in snow and ice over long periods of time across the landscape as a whole.

Other landscape-scale studies maintain that human impact is likely the main determinant of landscape pattern over much of the globe.[42] Landscapes may become substitutes for biodiversity measures because plant and animal composition differs between samples taken from sites within different landscape categories. Taxa, or different species, can “leak” from one habitat into another, which has implications for landscape ecology. As human land use practices expand and continue to increase the proportion of edges in landscapes, the effects of this leakage across edges on assemblage integrity may become more significant in conservation. This is because taxa may be conserved across landscape levels, if not at local levels.[43]

Land change modeling

Land change modeling is an application of landscape ecology designed to predict future changes in land use. Land change models are used in urban planning, geography, GIS, and other disciplines to gain a clear understanding of the course of a landscape.[44] In recent years, much of the Earth's land cover has changed rapidly, whether from deforestation or the expansion of urban areas.[45]

Relationship to other disciplines

Landscape ecology has been incorporated into a variety of ecological subdisciplines. For example, a recent development has been the more explicit consideration of spatial concepts and principles applied to the study of lakes, streams, and wetlands in the field of landscape limnology. Seascape ecology is a marine and coastal application of landscape ecology.[46] In addition, landscape ecology has important links to application-oriented disciplines such as agriculture and forestry. In agriculture, landscape ecology has introduced new options for the management of environmental threats brought about by the intensification of agricultural practices. Agriculture has always been a strong human impact on ecosystems.[15] In forestry, from structuring stands for fuelwood and timber to ordering stands across landscapes to enhance aesthetics, consumer needs have affected conservation and use of forested landscapes. Landscape forestry provides methods, concepts, and analytic procedures for landscape forestry.[47] Landscape ecology has been cited as a contributor to the development of fisheries biology as a distinct biological science discipline,[48] and is frequently incorporated in study design for wetland delineation in hydrology.[37] It has helped shape integrated landscape management.[49] Lastly, landscape ecology has been very influential for progressing sustainability science and sustainable development planning. In example, a recent study assessed sustainable urbanization across Europe using evaluation indices, country-landscapes, and landscape ecology tools and methods.[50]

Landscape ecology has also been combined with population genetics to form the field of landscape genetics, which addresses how landscape features influence the population structure and gene flow of plant and animal populations across space and time[51] and on how the quality of intervening landscape, known as "matrix," influences spatial variation.[52] After the term was coined in 2003, the field of landscape genetics had expanded to over 655 studies by 2010,[53] and continues to grow today. As genetic data has become more readily accessible, it is increasingly being used by ecologists to answer novel evolutionary and ecological questions,[54] many with regard to how landscapes effect evolutionary processes, especially in human-modified landscapes, which are experiencing biodiversity loss.[55]

See also


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Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, isolation and habitat area. Phytogeography is the branch of biogeography that studies the distribution of plants. Zoogeography is the branch that studies distribution of animals.

Knowledge of spatial variation in the numbers and types of organisms is as vital to us today as it was to our early human ancestors, as we adapt to heterogeneous but geographically predictable environments. Biogeography is an integrative field of inquiry that unites concepts and information from ecology, evolutionary biology, geology, and physical geography.Modern biogeographic research combines information and ideas from many fields, from the physiological and ecological constraints on organismal dispersal to geological and climatological phenomena operating at global spatial scales and evolutionary time frames.

The short-term interactions within a habitat and species of organisms describe the ecological application of biogeography. Historical biogeography describes the long-term, evolutionary periods of time for broader classifications of organisms. Early scientists, beginning with Carl Linnaeus, contributed to the development of biogeography as a science. Beginning in the mid-18th century, Europeans explored the world and discovered the biodiversity of life.

The scientific theory of biogeography grows out of the work of Alexander von Humboldt (1769–1859), Hewett Cottrell Watson (1804–1881), Alphonse de Candolle (1806–1893), Alfred Russel Wallace (1823–1913), Philip Lutley Sclater (1829–1913) and other biologists and explorers.

Cross-boundary subsidy

Cross-boundary subsidies are caused by organisms or materials that cross or traverse habitat patch boundaries, subsidizing the resident populations. The transferred organisms and materials may provide additional predators, prey, or nutrients to resident species, which can affect community and food web structure. Cross-boundary subsidies of materials and organisms occur in landscapes composed of different habitat patch types, and so depend on characteristics of those patches and on the boundaries in between them. Human alteration of the landscape, primarily through fragmentation, has the potential to alter important cross-boundary subsidies to increasingly isolated habitat patches. Understanding how processes that occur outside of habitat patches can affect populations within them may be important to habitat management.

Disturbance (ecology)

In ecology, a disturbance is a temporary change in environmental conditions that causes a pronounced change in an ecosystem. Disturbances often act quickly and with great effect, to alter the physical structure or arrangement of biotic and abiotic elements. Disturbance can also occur over a long period of time and can impact the biodiversity within an ecosystem. Major ecological disturbances may include fires, flooding, storms, insect outbreaks and trampling. Earthquakes, various types of volcanic eruptions, tsunami, firestorms, impact events, climate change, and the devastating effects of human impact on the environment (anthropogenic disturbances) such as clearcutting, forest clearing and the introduction of invasive species can be considered major disturbances. Not only invasive species can have a profound effect on an ecosystem, but also naturally occurring species can cause disturbance by their behavior. Disturbance forces can have profound immediate effects on ecosystems and can, accordingly, greatly alter the natural community. Because of these and the impacts on populations, disturbance determines the future shifts in dominance, various species successively becoming dominant as their life history characteristics, and associated life-forms, are exhibited over time.

Ecological niche

In ecology, a niche (CanE, UK: or US: ) is the match of a species to a specific environmental condition. It describes how an organism or population responds to the distribution of resources and competitors (for example, by growing when resources are abundant, and when predators, parasites and pathogens are scarce) and how it in turn alters those same factors (for example, limiting access to resources by other organisms, acting as a food source for predators and a consumer of prey). "The type and number of variables comprising the dimensions of an environmental niche vary from one species to another [and] the relative importance of particular environmental variables for a species may vary according to the geographic and biotic contexts".A Grinnellian niche is determined by the habitat in which a species lives and its accompanying behavioral adaptations. An Eltonian niche emphasizes that a species not only grows in and responds to an environment, it may also change the environment and its behavior as it grows. The Hutchinsonian niche uses mathematics and statistics to try to explain how species coexist within a given community.

The concept of ecological niche is central to ecological biogeography, which focuses on spatial patterns of ecological communities. "Species distributions and their dynamics over time result from properties of the species, environmental variation..., and interactions between the two—in particular the abilities of some species, especially our own, to modify their environments and alter the range dynamics of many other species." Alteration of an ecological niche by its inhabitants is the topic of niche construction.The majority of species exist in a standard ecological niche, sharing behaviors, adaptations, and functional traits similar to the other closely related species within the same broad taxonomic class, but there are exceptions. A premier example of a non-standard niche filling species is the flightless, ground-dwelling kiwi bird of New Zealand, which feeds on worms and other ground creatures, and lives its life in a mammal-like niche. Island biogeography can help explain island species and associated unfilled niches.

Ecosystem engineer

An ecosystem engineer is any organism that creates, significantly modifies, maintains or destroys a habitat. These organisms can have a large impact on the species richness and landscape-level heterogeneity of an area. As a result, ecosystem engineers are important for maintaining the health and stability of the environment they are living in. Since all organisms impact the environment they live in in one way or another, it has been proposed that the term "ecosystem engineers" be used only for keystone species whose behavior very strongly affects other organisms.


An ecotone is a transition area between two biomes. It is where two communities meet and integrate. It may be narrow or wide, and it may be local (the zone between a field and forest) or regional (the transition between forest and grassland ecosystems). An ecotone may appear on the ground as a gradual blending of the two communities across a broad area, or it may manifest itself as a sharp boundary line.

The word ecotone was coined from a combination of eco(logy) plus -tone, from the Greek tonos or tension – in other words, a place where ecologies are in tension.


In evolutionary ecology, an ecotype, sometimes called ecospecies, describes a genetically distinct geographic variety, population or race within a species, which is genotypically adapted to specific environmental conditions.

Typically, though ecotypes exhibit phenotypic differences (such as in morphology or physiology) stemming from environmental heterogeneity, they are capable of interbreeding with other geographically adjacent ecotypes without loss of fertility or vigor.

Edge effects

In ecology, edge effects are changes in population or community structures that occur at the boundary of two or more habitats. Areas with small habitat fragments exhibit especially pronounced edge effects that may extend throughout the range. As the edge effects increase, the boundary habitat allows for greater biodiversity.

Freshwater ecology of Maharashtra

The state of Maharashtra in India has several major river systems including those of the Narmada, Tapti, Godavari and Krishna rivers. The ecology of these rivers and associated wetlands is covered in this article.


In ecology, a habitat is the type of natural environment in which a particular species of organism lives. It is characterized by both physical and biological features. A species' habitat is those places where it can find food, shelter, protection and mates for reproduction.

The physical factors are for example soil, moisture, range of temperature, and light intensity as well as biotic factors such as the availability of food and the presence or absence of predators. Every organism has certain habitat needs for the conditions in which it will thrive, but some are tolerant of wide variations while others are very specific in their requirements. A habitat is not necessarily a geographical area, it can be the interior of a stem, a rotten log, a rock or a clump of moss, and for a parasitic organism it is the body of its host, part of the host's body such as the digestive tract, or a single cell within the host's body.

Habitat types include polar, temperate, subtropical and tropical. The terrestrial vegetation type may be forest, steppe, grassland, semi-arid or desert. Fresh water habitats include marshes, streams, rivers, lakes, and ponds, and marine habitats include salt marshes, the coast, the intertidal zone, estuaries, reefs, bays, the open sea, the sea bed, deep water and submarine vents.

Habitats change over time. This may be due to a violent event such as the eruption of a volcano, an earthquake, a tsunami, a wildfire or a change in oceanic currents; or the change may be more gradual over millennia with alterations in the climate, as ice sheets and glaciers advance and retreat, and as different weather patterns bring changes of precipitation and solar radiation. Other changes come as a direct result of human activities; deforestation, the plowing of ancient grasslands, the diversion and damming of rivers, the draining of marshland and the dredging of the seabed. The introduction of alien species can have a devastating effect on native wildlife, through increased predation, through competition for resources or through the introduction of pests and diseases to which the native species have no immunity.

Insular biogeography

Insular biogeography or island biogeography is a field within biogeography that examines the factors that affect the species richness and diversification of isolated natural communities. The theory was originally developed to explain the pattern of the species–area relationship occurring in oceanic islands. Under either name it is now used in reference to any ecosystem (present or past) that is isolated due to being surrounded by unlike ecosystems, and has been extended to mountain peaks, seamounts, oases, fragmented forests, and even natural habitats isolated by human land development. The field was started in the 1960s by the ecologists Robert H. MacArthur and E. O. Wilson, who coined the term island biogeography in their inaugural contribution to Princeton's Monograph in Population Biology series, which attempted to predict the number of species that would exist on a newly created island.


A landscape is the visible features of an area of land, its landforms, and how they integrate with natural or man-made features. A landscape includes the physical elements of geophysically defined landforms such as (ice-capped) mountains, hills, water bodies such as rivers, lakes, ponds and the sea, living elements of land cover including indigenous vegetation, human elements including different forms of land use, buildings, and structures, and transitory elements such as lighting and weather conditions. Combining both their physical origins and the cultural overlay of human presence, often created over millennia, landscapes reflect a living synthesis of people and place that is vital to local and national identity.

The character of a landscape helps define the self-image of the people who inhabit it and a sense of place that differentiates one region from other regions. It is the dynamic backdrop to people's lives. Landscape can be as varied as farmland, a landscape park or wilderness. The Earth has a vast range of landscapes, including the icy landscapes of polar regions, mountainous landscapes, vast arid desert landscapes, islands, and coastal landscapes, densely forested or wooded landscapes including past boreal forests and tropical rainforests, and agricultural landscapes of temperate and tropical regions. The activity of modifying the visible features of an area of land is referred to as landscaping.

Landscape epidemiology

Landscape epidemiology draws some of its roots from the field of landscape ecology. Just as the discipline of landscape ecology is concerned with analyzing both pattern and process in ecosystems across time and space, landscape epidemiology can be used to analyze both risk patterns and environmental risk factors. This field emerges from the theory that most vectors, hosts and pathogens are commonly tied to the landscape as environmental determinants control their distribution and abundance. In 1966, Evgeniy Pavlovsky introduced the concept of natural nidality or focality, defined by the idea that microscale disease foci are determined by the entire ecosystem. With the recent availability of new computing technologies such as geographic information systems, remote sensing, statistical methods including spatial statistics and theories of landscape ecology, the concept of landscape epidemiology has been applied analytically to a variety of disease systems, including malaria, hantavirus, Lyme disease and Chagas' disease.

Landscape limnology

Landscape limnology is the spatially explicit study of lakes, streams, and wetlands as they interact with freshwater, terrestrial, and human landscapes to determine the effects of pattern on ecosystem processes across temporal and spatial scales. Limnology is the study of inland water bodies inclusive of rivers, lakes, and wetlands; landscape limnology seeks to integrate all of these ecosystem types.

The terrestrial component represents spatial hierarchies of landscape features that influence which materials, whether solutes or organisms, are transported to aquatic systems; aquatic connections represent how these materials are transported; and human activities reflect features that influence how these materials are transported as well as their quantity and temporal dynamics.


A metapopulation consists of a group of spatially separated populations of the same species which interact at some level. The term metapopulation was coined by Richard Levins in 1969 to describe a model of population dynamics of insect pests in agricultural fields, but the idea has been most broadly applied to species in naturally or artificially fragmented habitats. In Levins' own words, it consists of "a population of populations".A metapopulation is generally considered to consist of several distinct populations together with areas of suitable habitat which are currently unoccupied. In classical metapopulation theory, each population cycles in relative independence of the other populations and eventually goes extinct as a consequence of demographic stochasticity (fluctuations in population size due to random demographic events); the smaller the population, the more chances of inbreeding depression and prone to extinction.

Although individual populations have finite life-spans, the metapopulation as a whole is often stable because immigrants from one population (which may, for example, be experiencing a population boom) are likely to re-colonize habitat which has been left open by the extinction of another population. They may also emigrate to a small population and rescue that population from extinction (called the rescue effect). Such a rescue effect may occur because declining populations leave niche opportunities open to the "rescuers".

The development of metapopulation theory, in conjunction with the development of source-sink dynamics, emphasised the importance of connectivity between seemingly isolated populations. Although no single population may be able to guarantee the long-term survival of a given species, the combined effect of many populations may be able to do this.

Metapopulation theory was first developed for terrestrial ecosystems, and subsequently applied to the marine realm. In fisheries science, the term "sub-population" is equivalent to the metapopulation science term "local population". Most marine examples are provided by relatively sedentary species occupying discrete patches of habitat, with both local recruitment and recruitment from other local populations in the larger metapopulation. Kritzer & Sale have argued against strict application of the metapopulation definitional criteria that extinction risks to local populations must be non-negligible.Finnish biologist Ilkka Hanski of the University of Helsinki was an important contributor to metapopulation theory.

Patch dynamics

Patch dynamics is an ecological perspective that the structure, function, and dynamics of ecological systems can be understood through studying their interactive patches. Patch dynamics, as a term, may also refer to the spatiotemporal changes within and among patches that make up a landscape. Patch dynamics is ubiquitous in terrestrial and aquatic systems across organizational levels and spatial scales. From a patch dynamics perspective, populations, communities, ecosystems, and landscapes may all be studied effectively as mosaics of patches that differ in size, shape, composition, history, and boundary characteristics.

The idea of patch dynamics dates back to the 1940s when plant ecologists studied the structure and dynamics of vegetation in terms of the interactive patches that it comprises. A mathematical theory of patch dynamics was developed by Simon Levin and Robert Paine in the 1970s, originally to describe the pattern and dynamics of an intertidal community as a patch mosaic created and maintained by tidal disturbances. Patch dynamics became a dominant theme in ecology between the late 1970s and the 1990s.

Patch dynamics is a conceptual approach to ecosystem and habitat analysis that emphasizes dynamics of heterogeneity within a system (i.e. that each area of an ecosystem is made up of a mosaic of small 'sub-ecosystems').Diverse patches of habitat created by natural disturbance regimes are seen as critical to the maintenance of this diversity (ecology). A habitat patch is any discrete area with a definite shape, spatial and configuration used by a species for breeding or obtaining other resources. Mosaics are the patterns within landscapes that are composed of smaller elements, such as individual forest stands, shrubland patches, highways, farms, or towns.

Shade tree

A shade tree is a large tree whose primary role is to provide shade in the surrounding environment due to its spreading canopy and crown, where it may give shelter from sunlight in the heat of the summer for people who seek recreational needs in urban parks and house yards, and thus, also protecting them from the sun's harmful UV rays and sunburns. Therefore, some shade trees may be grown specifically for the comfort of the population due to their convenient shelter.

Furthermore, shade trees are also effective in reducing the energy used in cooling homes.

Spatial ecology

Spatial ecology studies the ultimate distributional or spatial unit occupied by a species. In a particular habitat shared by several species, each of the species is usually confined to its own microhabitat or spatial niche because two species in the same general territory cannot usually occupy the same ecological niche for any significant length of time.


A windbreak (shelterbelt) is a planting usually made up of one or more rows of trees or shrubs planted in such a manner as to provide shelter from the wind and to protect soil from erosion. They are commonly planted in hedgerows around the edges of fields on farms. If designed properly, windbreaks around a home can reduce the cost of heating and cooling and save energy. Windbreaks are also planted to help keep snow from drifting onto roadways or yards. Farmers sometimes use windbreaks to keep snow drifts on farm land that will provide water when the snow melts in the spring. Other benefits include contributing to a microclimate around crops (with slightly less drying and chilling at night), providing habitat for wildlife, and, in some regions, providing wood if the trees are harvested.

Windbreaks and intercropping can be combined in a farming practice referred to as alleycropping. Fields are planted in rows of different crops surrounded by rows of trees. These trees provide fruit, wood, or protect the crops from the wind. Alley cropping has been particularly successful in India, Africa, and Brazil, where coffee growers have combined farming and forestry.A further use for a shelterbelt is to screen a farm from a main road or motorway. This improves the farm landscape by reducing the visual incursion of the motorway, mitigating noise from the traffic and providing a safe barrier between farm animals and the road.

The term "windbreak" is also used to describe an article of clothing worn to prevent wind chill; this term is favored by Europeans whereas Americans tend to use the term "windbreaker".

Fences called "windbreaks" are also used. Normally made from cotton, nylon, canvas, and recycled sails, windbreaks tend to have three or more panels held in place with poles that slide into pockets sewn into the panel. The poles are then hammered into the ground and a windbreak is formed. Windbreaks or "wind fences" are used to reduce wind speeds over erodible areas such as open fields, industrial stockpiles, and dusty industrial operations. As erosion is proportional to wind speed cubed, a reduction of wind speed of 1/2 (for example) will reduce erosion by over 80%.

Food webs
Example webs
Ecology: Modelling ecosystems: Other components
Related fields

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