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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] Landscape ecology can be described as the science of "landscape diversity" as the synergetic result of biodiversity and geodiversity.^[4]

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]^[5]^[6]

The most salient characteristics of landscape ecology are its emphasis on the relationship among pattern, process and scales, 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.^[7] Landscape ecology also studies the role of human impacts on landscape diversity in the development and spreading of new human pathogens that could trigger epidemics.^[8]^[9]

Terminology

The German term Landschaftsökologie – thus landscape ecology – was coined by German geographer Carl Troll in 1939.^[10] 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.

Explanation

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.^[11] Landscape ecology also includes geomorphology as applied to the design and architecture of landscapes.^[12] Geomorphology is the study of how geological formations are responsible for the structure of a landscape.

History

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^[13]). 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)^[14] 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.^[when?] 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.^[15]

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^[16] and Forman and Godron.^[17]^[18] Forman^[6] 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.^[5] 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):^[19]

Interdisciplinary analysis of subjectively defined landscape units (e.g. Neef School^[20]^[21]): 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.
Topological ecology at the landscape scale^[22]^[23] '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.
Organism-centered, multi-scale topological ecology (e.g. John A. Wiens^[24]^[25]): 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.
Topological ecology at the landscape level of biological organisation (e.g. Urban et al.^[26]): 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.
Analysis of social-ecological systems using the natural and social sciences and humanities (e.g. Leser;^[27] Naveh;^[28]^[29] Zonneveld^[30]): 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.
Ecology guided by cultural meanings of lifeworldly landscapes (frequently pursued in practice^[31] but not defined, but see, e.g., Hard;^[32] Trepl^[19]): 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.^[33]

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.

Landscape

Certainly, 'landscape' is a central concept in landscape ecology. It is, however, defined in quite different ways. For example:^[19] Carl Troll conceives of landscape not as a mental construct but as an objectively given 'organic entity', a harmonic individuum of space.^[34] Ernst Neef^[20]^[21] 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 Michel Godron,^[22] 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^[24]^[25] 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".^[25]^[35] Some define 'landscape' as an area containing two or more ecosystems in close proximity.^[15]

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

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General	Abiotic component Abiotic stress Behaviour Biogeochemical cycle Biomass Biotic component Biotic stress Carrying capacity Competition Ecosystem Ecosystem ecology Ecosystem model Green world hypothesis Keystone species List of feeding behaviours Metabolic theory of ecology Productivity Resource Restoration
Producers	Autotrophs Chemosynthesis Chemotrophs Foundation species Kinetotrophs Mixotrophs Myco-heterotrophy Mycotroph Organotrophs Photoheterotrophs Photosynthesis Photosynthetic efficiency Phototrophs Primary nutritional groups Primary production
Consumers	Apex predator Bacterivore Carnivores Chemoorganotroph Foraging Generalist and specialist species Intraguild predation Herbivores Heterotroph Heterotrophic nutrition Insectivore Mesopredators Mesopredator release hypothesis Omnivores Optimal foraging theory Planktivore Predation Prey switching
Decomposers	Chemoorganoheterotrophy Decomposition Detritivores Detritus
Microorganisms	Archaea Bacteriophage Lithoautotroph Lithotrophy Marine Microbial cooperation Microbial ecology Microbial food web Microbial intelligence Microbial loop Microbial mat Microbial metabolism Phage ecology
Food webs	Biomagnification Ecological efficiency Ecological pyramid Energy flow Food chain Trophic level
Example webs	Lakes Rivers Soil Tritrophic interactions in plant defense Marine food webs cold seeps hydrothermal vents intertidal kelp forests North Pacific Gyre San Francisco Estuary tide pool
Processes	Ascendency Bioaccumulation Cascade effect Climax community Competitive exclusion principle Consumer–resource interactions Copiotrophs Dominance Ecological network Ecological succession Energy quality Energy systems language f-ratio Feed conversion ratio Feeding frenzy Mesotrophic soil Nutrient cycle Oligotroph Paradox of the plankton Trophic cascade Trophic mutualism Trophic state index
Defense, counter	Animal coloration Anti-predator adaptations Camouflage Deimatic behaviour Herbivore adaptations to plant defense Mimicry Plant defense against herbivory Predator avoidance in schooling fish


Population ecology	Abundance Allee effect Consumer-resource model Depensation Ecological yield Effective population size Intraspecific competition Logistic function Malthusian growth model Maximum sustainable yield Overpopulation Overexploitation Population cycle Population dynamics Population modeling Population size Predator–prey (Lotka–Volterra) equations Recruitment Small population size Stability Resilience Resistance Random generalized Lotka–Volterra model
Species	Biodiversity Density-dependent inhibition Ecological effects of biodiversity Ecological extinction Endemic species Flagship species Gradient analysis Indicator species Introduced species Invasive species / Native species Latitudinal gradients in species diversity Minimum viable population Neutral theory Occupancy–abundance relationship Population viability analysis Priority effect Rapoport's rule Relative abundance distribution Relative species abundance Species diversity Species homogeneity Species richness Species distribution Species–area curve Umbrella species
Species interaction	Antibiosis Biological interaction Commensalism Community ecology Ecological facilitation Interspecific competition Mutualism Parasitism Storage effect Symbiosis
Spatial ecology	Biogeography Cross-boundary subsidy Ecocline Ecotone Ecotype Disturbance Edge effects Foster's rule Habitat fragmentation Ideal free distribution Intermediate disturbance hypothesis Insular biogeography Land change modeling Landscape ecology Landscape epidemiology Landscape limnology Metapopulation Patch dynamics r/K selection theory Resource selection function Source–sink dynamics
Niche	Ecological trap Ecosystem engineer Environmental niche modelling Guild Habitat Marine Semiaquatic Terrestrial Limiting similarity Niche apportionment models Niche construction Niche differentiation Ontogenetic niche shift
Other networks	Assembly rules Bateman's principle Bioluminescence Ecological collapse Ecological debt Ecological deficit Ecological energetics Ecological indicator Ecological threshold Ecosystem diversity Emergence Extinction debt Kleiber's law Liebig's law of the minimum Marginal value theorem Thorson's rule Xerosere
Other	Allometry Alternative stable state Balance of nature Biological data visualization Ecological economics Ecological footprint Ecological forecasting Ecological humanities Ecological stoichiometry Ecopath Ecosystem based fisheries Endolith Evolutionary ecology Functional ecology Industrial ecology Macroecology Microecosystem Natural environment Regime shift Sexecology Systems ecology Urban ecology Theoretical ecology
Outline of ecology


General	Degradation Development/Conversion Planning Conflict Land management Landscaping Integrated landscape management Land grabbing Land consumption Land loss Habitat destruction Illegal construction Land reclamation Land rehabilitation Landscape ecology Rangeland management Environmental planning Leopold matrix Watertable control Developed environments Built-up area Locally unwanted land use
Property	Property Subdivision (land) Real estate development Land development bank Land (economics) Customary land
Related fields	Soil Soil science Soil compaction Soil contamination Alkali soil Pollution Deforestation Urban planning Agriculture Drainage system (agriculture) Land change modeling
Categories: Land use


Methodology	Field ecology Quantitative ecology Theoretical ecology
Spatial scale	Microecology Macroecology
Organisation or scope	Autecology Synecology Population ecology Community ecology Ecosystem ecology
Taxonomy or taxon	Microbial ecology Plant ecology Insect ecology Human ecology
Biome or biogeographical	Polar ecology Arctic ecology Desert ecology Forest ecology Soil ecology Tropical ecology Urban ecology
Ecological phenomena	Behavioral ecology Chemical ecology Disease ecology Ecophysiology Ecotoxicology Evolutionary ecology Fire ecology Functional ecology Genetic ecology Landscape ecology Molecular ecology Paleoecology Social ecology Spatial ecology Thermal ecology
Interdisciplinarity	Agroecology Ecological anthropology Ecological economics Ecological engineering Political ecology Systems ecology
Other	History of ecology Glossary of ecology Applied ecology Restoration ecology Ecosophy Natural history Biogeography

Authority control databases
National	Germany United States France BnF data Japan Czech Republic Israel
Other	Encyclopedia of Modern Ukraine Yale LUX


Atmospheric science / Meteorology Biogeography / Phytogeography Climatology / Paleoclimatology / Palaeogeography Coastal geography / Oceanography Soil science / Pedology / Edaphology Geobiology Geology Geomorphology Geostatistics Glaciology Hydrology / Limnology Landscape ecology Quaternary science
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