View source for Landscape ecology

{{Short description|Relationships between ecological processes in the environment and particular ecosystems}}
[[Image:NLCD landcover MSN area.png|thumb|Land cover surrounding Madison, Wisconsin. Fields are colored yellow and brown and urban surfaces are colored red.]]
[[Image:NLCD impervious MSN area.png|thumb|[[Impervious surface|Impervious surfaces]] surrounding Madison, Wisconsin]]
[[Image:NLCD canopy MSN area.png|thumb|[[Canopy (biology)|Canopy]] cover surrounding Madison, Wisconsin]]

'''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.<ref>{{cite journal | vauthors = Wu J | title = Landscape ecology, cross-disciplinarity, and sustainability science. | date = January 2006 | journal = Landscape Ecology | volume = 21 | issue =1 | pages = 1–4 | doi = 10.1007/s10980-006-7195-2 | s2cid = 27192835 | url = http://elib.bsu.by/handle/123456789/173085 }}</ref><ref name="wu-2007">{{cite book | veditors = Wu J, Hobbs R | date = 2007 | title = Key Topics in Landscape Ecology. | publisher = Cambridge University Press | location = Cambridge }}</ref><ref>{{cite book | vauthors = Wu J | date = 2008 | chapter = Landscape ecology. | veditors = Jorgensen SE | title = Encyclopedia of Ecology | publisher = Elsevier | location = Oxford }}</ref> Landscape ecology can be described as the science of "landscape diversity" as the synergetic result of [[biodiversity]] and [[geodiversity]].<ref>{{cite book |last1=Leser |first1=Hartmut |last2=Nagel |first2=Peter |chapter=Landscape diversity — a holistic approach | name-list-style = vanc | doi = 10.1007/978-3-662-06071-1_9 |title=Biodiversity |year=2001 |publisher=Springer  |pages=129–143 |isbn=978-3-642-08370-9 }}</ref>

As a highly interdisciplinary field in [[systems science]], landscape ecology integrates [[biophysical]] and [[Analytic induction|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.<ref name="wu-2007" /><ref name="Turner-2001">{{cite book | vauthors = Turner MG, Gardner RH, O'Neill RV | date = 2001 | title = Landscape Ecology in Theory and Practice. | publisher = Springer-Verlag | location = New York, NY, USA }}</ref><ref name="Forman_1995">{{cite book | vauthors = Forman RT | date = 1995 | title = Land Mosaics: The Ecology of Landscapes and Regions. | publisher = Cambridge University Press | location = Cambridge, UK }}</ref>

The most salient characteristics of landscape ecology are its emphasis on the relationship among pattern, process and [[Scale (geography)|scales]], and its focus on broad-scale ecological and environmental issues. These necessitate the coupling between biophysical and [[Socioeconomics|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]].<ref>Wu & Hobbs 2002</ref> Landscape ecology also studies the role of human impacts on landscape diversity in the development and spreading of new human pathogens that could trigger [[epidemic]]s.<ref>{{Cite journal|last1=Bloomfield|first1=Laura S. P.|last2=McIntosh|first2=Tyler L.|last3=Lambin|first3=Eric F. | name-list-style = vanc |date=2020-04-01|title=Habitat fragmentation, livelihood behaviors, and contact between people and nonhuman primates in Africa|journal=Landscape Ecology|language=en|volume=35|issue=4|pages=985–1000|doi=10.1007/s10980-020-00995-w|s2cid=214731443|issn=1572-9761|doi-access=free|hdl=2078.1/243632|hdl-access=free}}</ref><ref>{{cite journal | vauthors = Bausch DG, Schwarz L | title = Outbreak of ebola virus disease in Guinea: where ecology meets economy | journal = PLOS Neglected Tropical Diseases | volume = 8 | issue = 7 | article-number = e3056 | date = 2014-07-31 | pmid = 25079231 | pmc = 4117598 | doi = 10.1371/journal.pntd.0003056 | doi-access = free }}</ref>

== Terminology ==

The German term {{Lang|de|Landschaftsökologie}} – thus ''landscape ecology'' – was coined by German [[geographer]] [[Carl Troll]] in 1939.<ref>{{cite journal | vauthors = Troll C | date = 1939 | title = Luftbildplan und ökologische Bodenforschung | trans-title = Aerial photography and ecological studies of the earth | language = de | journal =  Zeitschrift der Gesellschaft für Erdkunde | location = Berlin | pages = 241–298 }}</ref>  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.<ref>{{cite journal | vauthors = Turner MG | title = Landscape ecology: the effect of pattern on process. | journal = Annual Review of Ecology and Systematics | volume = 20 | date = 1989 | pages = 171–197 | doi = 10.1146/annurev.es.20.110189.001131 }}</ref>  Landscape ecology also includes geomorphology as applied to the design and architecture of landscapes.<ref name="Allaby-1998">{{cite book | vauthors = Allaby M | date = 1998 | title = Oxford Dictionary of Ecology.  | publisher = Oxford University Press | location = New York, NY }}</ref> [[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 [[Robert MacArthur|MacArthur]] & [[Edward Osborne Wilson|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 (ecology)|habitat]] by Levins' [[metapopulation]] model (which can be applied e.g. to forest islands in the agricultural landscape<ref>{{cite book | veditors = Banaszak J | date = 2000 | title = Ecology of Forest Islands | publisher = Bydgoszcz University Press | location = Bydgoszcz, Poland | page = 313 }}</ref>).  This generalization spurred the growth of landscape ecology by providing [[conservation biology|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 system]]s (GIS)<ref name="sstein-2009">{{cite journal | vauthors = Steiniger S, Hay GJ | title = Free and open source geographic information tools for landscape ecology. | journal = Ecological Informatics | date = September 2009 | volume = 4 | issue = 4 | pages = 183–95 | doi = 10.1016/j.ecoinf.2009.07.004 | url = https://sourceforge.net/projects/jump-pilot/files/w_other_freegis_documents/articles/sstein_freegitools_ecoinf2009.pdf }}</ref> and the availability of large-extent habitat data (e.g. [[remote sensing|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|date=May 2014}}  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 [[ecosystem]]s, [[population]]s, [[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.<ref name="Sanderson-2000">{{cite book | veditors = Sanderson J, Harris LD | date = 2000 | title = Landscape Ecology: A Top-Down Approach. | publisher = Lewis Publishers | location = Boca Raton, Florida, USA }}</ref>

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<ref name="Naveh-1984">{{cite book | vauthors = Naveh Z, Lieberman A | date = 1984 | title = Landscape ecology: theory and application. | publisher = Springer-Verlag | location = New York, NY, USA }}</ref> and Forman and Godron.<ref name="Forman-1986">{{cite book | vauthors = Forman RT, Godron M | date = 1986 | title = Landscape Ecology | publisher = John Wiley and Sons, Inc. | location = New York, NY, USA }}</ref><ref name="Ryszkowski-2002">{{cite book | veditors = Ryszkowski L | date = 2002 | title = Landscape Ecology in Agroecosystems Management. | publisher = CRC Press, Boca Raton | location = Florida, USA }}</ref>  Forman<ref name="Forman_1995"/> 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 [[computer simulation|models]].  There has been associated development of powerful quantitative methods to examine the interactions of patterns and processes.<ref name="Turner-2001" />  An example would be determining the amount of [[carbon cycle|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 geography|integrated]] view of [[geography]] (in conceptions 1, 5, and 6):<ref name = "Kirchhoff_2012">{{cite journal | vauthors = Kirchhoff T, Trepl L, Vicenzotti V | title = What is landscape ecology? An analysis and evaluation of six different conceptions. | journal = Landscape Research | date = February 2013 | volume = 38 | issue = 1 | pages = 33–51 | doi = 10.1080/01426397.2011.640751 | s2cid = 145421450 }} All the following quotations and descriptions come from this source.</ref>
# ''Interdisciplinary analysis of subjectively defined landscape units'' (e.g. [[Ernst Neef|Neef]] School<ref name = "Neef_1967">{{cite book | vauthors = Neef E | date = 1967 | title = Die theoretischen Grundlagen der Landschaftslehre | trans-title = The theoretical basics of landscape science | language = de | location = Gotha | publisher = Haack }}</ref><ref name = "Haase_1990">{{cite journal | vauthors = Haase G | date = 1990 | title =  Approaches to, and methods of landscape diagnosis as a basis of landscape planning and landscape management | journal = Ekológia | volume = 9 | issue = 1 | pages = 31–44 }}</ref>):  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''<ref name = "Forman_1981">{{cite journal | vauthors = Forman RT, Godron M | title = Patches and structural components for a landscape ecology. | journal = BioScience | date = November 1981 | volume = 31 | issue = 10 | pages = 733–40 | doi = 10.2307/1308780 | jstor = 1308780 }}</ref><ref>{{cite book | vauthors = Forman RT, Godron M | date = 1986 | title = Landscape ecology | publisher = Wiley | location = NY }}</ref> '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]]<ref name = "Wiens_1989">{{cite journal | vauthors = Wiens JA, Milne BT | title = Scaling of 'landscapes' in landscape ecology, or, landscape ecology from a beetle's perspective. | journal = Landscape Ecology | date = December 1989 | volume = 3 | issue = 2 | pages = 87–96 | doi = 10.1007/BF00131172 | s2cid = 15683804 }}</ref><ref name = "Wiens_1999">{{cite book | vauthors = Wiens JA | date = 1999 | chapter = The science and practice of landscape ecology. | veditors = Klopatek JM, Gardner RH | title = Landscape ecological analyses: Issues and applications. | publisher = Springer | location = NY | pages = 371–383 }}</ref>):  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.<ref>{{cite journal | vauthors = Urban DL, O'Neill RV, Shugart Jr HH | title = A hierarchical perspective can help scientists understand spatial patterns. | journal = BioScience | date = February 1987 | volume = 37 | issue = 2 | pages = 119–27 | doi = 10.2307/1310366 | jstor = 1310366 | url = http://www.edc.uri.edu/nrs/classes/nrs534/NRS_534_readings/Urbanetal1987.pdf | archive-date = 2021-05-04 | access-date = 2020-05-20 | archive-url = https://web.archive.org/web/20210504130937/http://www.edc.uri.edu/nrs/classes/nrs534/NRS_534_readings/Urbanetal1987.pdf }}</ref>):  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;<ref>{{cite book | vauthors = Leser H | date = 1991 | title = Landschaftsökologie. Ansatz, Modelle, Methodik, Anwendung. | publisher = Ulmer | location = Stuttgart }}</ref> Naveh;<ref>{{cite book | vauthors = Naveh Z, Lieberman AS | date = 1984 | title = Landscape ecology. Theory and application. | publisher = Springer | location = NY }}</ref><ref>{{cite journal | vauthors = Naveh N | date = 2000 | title = What is holistic landscape ecology? A conceptual introduction. | journal = Landscape and Urban Planning | volume = 50 | issue = 1–3 | pages = 7–26 | doi = 10.1016/S0169-2046(00)00077-3 }}</ref> Zonneveld<ref>{{cite book | vauthors = Zonneveld IS | date = 1995 | title = Land ecology: an introduction to landscape ecology as a base for land evaluation, land management and conservation. | publisher = SPB | location = Amsterdam }}</ref>):  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<ref>However, not always under the designation 'landscape ecology', but as part of landscape stewardship, landscape architecture and, first and foremost, environmental or urban and landscape planning.</ref> but not defined, but see, e.g., Hard;<ref>{{cite book | vauthors = Hard G | date = 1973 | title = Die Geographie. Eine wissenschaftstheoretische Einführung. | publisher = deGruyter | location = Berlin | pages =  92–95 }}</ref> Trepl<ref name = "Kirchhoff_2012" />):  Landscape ecology is defined as ecology that is guided by an external aim, namely, to maintain and develop [[lifeworld]]ly [[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 pattern]]s.<ref name="Turner-1991">{{cite book | veditors = Turner MG, Gardner RH | date = 1991 | title = Quantitative Methods in Landscape Ecology. | publisher = Springer-Verlag | location = New York, NY, USA }}</ref>

== 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 ===
{{Main|Landscape}}

Certainly, 'landscape' is a central concept in landscape ecology. It is, however, defined in quite different ways. For example:<ref name = "Kirchhoff_2012" /> [[Carl Troll]] conceives of landscape not as a mental construct but as an objectively given 'organic entity', a ''harmonic individuum of space''.<ref>{{cite book | vauthors = Troll C | date = 2007 | chapter = The geographic landscape and its investigation. | veditors = Wiens JA, Moss MR, Turner MG, Mladenoff DJ | title = Foundation papers in landscape ecology. | location = New York | publisher = Columbia University Press | pages = 71–101 }} First published as: {{cite book | vauthors = Troll C | title = Studium Generale | chapter = Die geographische Landschaft und ihre Erforschung | date = 1950 | volume = 3 | issue = 4–5 | pages = 163–181 | doi = 10.1007/978-3-662-38240-0_20 | isbn = 978-3-662-37475-7 }}</ref> 
[[Ernst Neef]]<ref name = "Neef_1967" /><ref name = "Haase_1990" /> 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 [[anthropocentrism|anthropocentric]] and relativistic way.
According to [[Richard Forman]] and [[Michel Godron]],<ref name = "Forman_1981" /> 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]]<ref name = "Wiens_1989" /><ref name = "Wiens_1999" /> 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".<ref name = "Wiens_1999" /><ref>{{cite book | vauthors = Wiens JA | date = 2005 | chapter = Toward a unified landscape ecology | veditors = Wiens JA, Moss MR | title = Issues and perspectives in landscape ecology. | publisher = Cambridge University Press | location = Cambridge | pages = 365–373 }}</ref> Some define 'landscape' as an area containing two or more [[ecosystem]]s in close proximity.<ref name="Sanderson-2000"/>

=== Scale and heterogeneity (incorporating composition, structure, and function) ===
{{Main|Scale (geography)|Spatial heterogeneity}}

A main concept in landscape ecology is ''[[scale (geography)|scale]]''. Scale represents the real world as translated onto a map, relating distance on a map image and the corresponding distance on earth.<ref>{{cite book | vauthors = Malczewski J | date =  1999 | title = GIS and Multicriteria Decision Analysis. | publisher = John Wiley and Sons, Inc. | location = New York, NY, USA }}</ref> Scale is also the spatial or temporal measure of an object or a process,<ref name="Turner-1991" /> or amount of spatial resolution.<ref name="Forman_1995" /> 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 [[road]]s 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.<ref name="Turner-1991" /> ''Pattern'' is the term for the contents and internal order of a heterogeneous area of land.<ref name="Forman-1986" />

A landscape with structure and pattern implies that it has spatial ''heterogeneity'', or the uneven distribution of objects across the landscape.<ref name="Forman_1995" /> 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.<ref>{{cite journal| vauthors = Wirth E, Szabó G, Czinkóczky A |date=2016-06-07|journal=ISPRS Annals of the Photogrammetry, Remote Sensing and Spatial Information Sciences|language=en|volume=III-8|pages=145–151|doi=10.5194/isprs-annals-iii-8-145-2016|issn=2194-9042|title=Measure of Landscape Heterogeneity by Agent-Based Methodology|bibcode=2016ISPAnIII8..145W|doi-access=free}}</ref>

===Patch and mosaic===<!-- This section is linked from [[Patch]] -->

{{See also|Patch dynamics}}
''Patch'', a term fundamental to landscape ecology, is defined as a relatively homogeneous area that differs from its surroundings.<ref name="Forman_1995"/>  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.<ref name="Forman_1995"/>

''Matrix'' is the "background ecological system" of a landscape with a high degree of [[Landscape connectivity|connectivity]].  ''Connectivity'' is the measure of how connected or spatially continuous a corridor, network, or matrix is.<ref name="Forman_1995"/>  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.<ref name="Forman_1995"/>  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.<ref name="Forman_1995"/>

=== Boundary and edge ===
{{Main|Edge effect}}

Landscape patches have a boundary between them which can be defined or fuzzy.<ref name="Sanderson-2000" /> The zone composed of the edges of adjacent ecosystems is the ''boundary''.<ref name="Forman_1995" /> ''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.<ref name="Forman_1995" /> 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.<ref name="Turner-1991" />

=== Ecotones, ecoclines, and ecotopes ===
{{Main|Ecotone|Ecocline|Ecotype}}

A type of boundary is the ''ecotone'', or the transitional zone between two communities.<ref name="Allaby-1998" /> Ecotones can arise naturally, such as a [[Shore|lakeshore]], or can be human-created, such as a cleared [[agricultural field]] from a forest.<ref name="Allaby-1998" /> The ecotonal community retains characteristics of each bordering community and often contains species not found in the adjacent communities. Classic examples of ecotones include [[fencerow]]s, forest to marshlands transitions, forest to grassland transitions, or land-water interfaces such as [[riparian zone]]s 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 [[Biodiversity|species richness]] higher or lower than either side of the ecotone.<ref name="Walker_2003">{{cite journal | vauthors = Walker S, Wilson JB, Steel JB, Rapson GL, Smith B, King WM, Cottam YH | title = Properties of ecotones: evidence from five ecotones objectively determined from a coastal vegetation gradient. | journal = Journal of Vegetation Science | date = August 2003 | volume = 14 | issue = 4 | pages = 579–90 | doi = 10.1111/j.1654-1103.2003.tb02185.x | doi-access = free }}</ref>

An ''[[wikt:ecocline|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.<ref name="Attrill-2002">{{cite journal | vauthors = Attrill MJ, Rundle SD | title = Ecotone or ecocline: ecological boundaries in estuaries. | journal = Estuarine, Coastal and Shelf Science | date = December 2002 | volume = 55 | issue = 6 | pages = 929–36 | doi = 10.1006/ecss.2002.1036 | bibcode = 2002ECSS...55..929A }}</ref> An ''ecotope'' is a spatial term representing the smallest ecologically distinct unit in mapping and classification of landscapes.<ref name="Forman_1995" /> 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 ===
{{Main|Disturbance (ecology)|Habitat 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.<ref name="Forman_1995" /> 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 exceeds a critical level, which means that landscapes exhibit two phases: connected and disconnected.<ref name="Green-2006">{{cite book | vauthors = Green DG, Klomp NI, Rimmington GR, Sadedin S | date = 2006 | title = Complexity in Landscape Ecology | publisher = Springer | location = Amsterdam | url = http://www.csse.monash.edu.au/~dgreen/books/springer/ | access-date=2008-03-22 |archive-url = https://web.archive.org/web/20080619025437/http://www.csse.monash.edu.au/~dgreen/books/springer/ | archive-date=2008-06-19 }}</ref>

== Theory ==
Landscape ecology theory stresses the role of human impacts on landscape structures and functions. It also proposes ways for restoring degraded landscapes.<ref name="Naveh-1984"/> Landscape ecology explicitly includes humans as entities that cause functional changes on the landscape.<ref name="Sanderson-2000"/> Landscape ecology theory includes the landscape stability principle, which emphasizes the importance of landscape structural heterogeneity in developing resistance to disturbances, [[regeneration (ecology)|recovery from disturbances]], and promoting total system stability.<ref name="Forman-1986"/> 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.<ref name="Turner-2001" /> 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.<ref name="Ryszkowski-2002" /> 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 basin]]s, which in turn are composed of local ecosystems, which are in turn composed of individual trees and gaps.<ref name="Forman_1995"/>  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.<ref name="Turner-1991"/>  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 (ecology)|habitat]] requirements.<ref name="Turner-1991"/>

== Application ==

===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 [[Natural environment|environmental]] problems.<ref name="Turner-2001"/>  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.<ref name="Turner-1991"/> [[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).<ref>{{cite journal | vauthors = Lyon J, Sagers CL | title = Structure of herbaceous plant assemblages in a forested riparian landscape. | journal = Plant Ecology | date = September 1998 | volume = 138 | issue = 1 | pages = 1–6 | doi = 10.1023/A:1009705912710 | s2cid = 28628830 }}</ref>

[[Climate change]] is another major component in structuring current research in landscape ecology.<ref name="raul">{{cite journal | vauthors = Ochoa-Hueso R, Delgado-Baquerizo M, King PT, Benham M, Arca V, Power SA |title=Ecosystem type and resource quality are more important than global change drivers in regulating early stages of litter decomposition |journal=Soil Biology and Biochemistry |date=February 2019 |volume=129 |pages=144–152 |doi=10.1016/j.soilbio.2018.11.009 |s2cid=92606851 |hdl=10261/336676 |hdl-access=free }}</ref>  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.<ref name="Walker_2003"/>  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.<ref>{{cite journal | vauthors = Löffler J, Finch OD | title = Spatio-temporal gradients between high mountain ecosystems of central Norway. | journal = Arctic, Antarctic, and Alpine Research | date = November 2005 | volume = 37 | issue = 4 | pages = 499–513 | doi = 10.1657/1523-0430(2005)037[0499:sgbhme]2.0.co;2 | s2cid = 131326887 | doi-access = free }}</ref>  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.<ref>{{Cite journal |last1=Ellis |first1=Erle C. |last2=Gauthier |first2=Nicolas |last3=Klein Goldewijk |first3=Kees |last4=Bliege Bird |first4=Rebecca |last5=Boivin |first5=Nicole |last6=Díaz |first6=Sandra |last7=Fuller |first7=Dorian Q. |last8=Gill |first8=Jacquelyn L. |last9=Kaplan |first9=Jed O. |last10=Kingston |first10=Naomi |last11=Locke |first11=Harvey |last12=McMichael |first12=Crystal N. H. |last13=Ranco |first13=Darren |last14=Rick |first14=Torben C. |last15=Shaw |first15=M. Rebecca |date=2021-04-27 |title=People have shaped most of terrestrial nature for at least 12,000 years |journal=Proceedings of the National Academy of Sciences |language=en |volume=118 |issue=17 |article-number=e2023483118 |doi=10.1073/pnas.2023483118 |issn=0027-8424 |pmc=8092386 |pmid=33875599 |doi-access=free }}</ref><ref>{{cite journal | vauthors = Wilson JB, King WM | title = Human-mediated vegetation switches as processes in landscape ecology. | journal = Landscape Ecology | date = August 1995 | volume = 10 | issue = 4 | pages = 191–6 | doi = 10.1007/BF00129253 | s2cid = 772430 }}</ref>  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.<ref>{{cite journal | vauthors = Dangerfield JM, Pik AJ, Britton D, Holmes A, Gillings M, Oliver IA, Briscoe D, Beattie AJ | title = Patterns of invertebrate biodiversity across a natural edge. | journal = Austral Ecology | date = June 2003 | volume = 28 | issue = 3 | pages = 227–36 | doi = 10.1046/j.1442-9993.2003.01240.x | doi-access = free }}</ref>

=== 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, [[Geographic information science|GIS]], and other disciplines to gain a clear understanding of the course of a landscape.<ref>{{Cite book|url=https://www.nap.edu/read/18385/chapter/3|title=Advancing Land Change Modeling: Opportunities and Research Requirements|last=National Research Council|publisher=National Academies Press|year=2014|isbn=978-0-309-28833-0|pages=Chapter 1|language=en|doi=10.17226/18385}}</ref> In recent years, much of the Earth's land cover has changed rapidly, whether from [[deforestation]] or the [[Urbanization|expansion of urban areas]].<ref>{{Cite web|url=http://glcf.umd.edu/services/landcoverchange/|title=GLCF: Global Land Cover Change|last=University of Maryland|website=glcf.umd.edu|access-date=2018-12-27|archive-url=https://web.archive.org/web/20190609023650/http://www.glcf.umd.edu/services/landcoverchange/|archive-date=2019-06-09}}</ref>

===Relationship to other disciplines===
Landscape ecology has been incorporated into a variety of ecological subdisciplines. For example, it is closely linked to [[land change science]], the interdisciplinary of land use and land cover change and their effects on surrounding ecology. Another 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.<ref>{{cite book | veditors = Pittman SJ | date = 2017 | title = Seascape Ecology | publisher = Wiley & Sons }}</ref> 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.<ref name="Ryszkowski-2002"/>

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.<ref>{{cite book | vauthors = Boyce SG | date = 1995 | title = Landscape Forestry | publisher = John Wiley and Sons, Inc. | location = New York, NY }}</ref> Landscape ecology has been cited as a contributor to the development of [[fishery|fisheries]] biology as a distinct biological science discipline,<ref>{{cite journal | vauthors = Magnuson JJ | title = Fish and fisheries ecology. | journal = Ecological Applications | date = February 1991 | volume = 1 | issue = 1 | pages = 13–26 | doi = 10.2307/1941844 | jstor = 1941844 | pmid = 27755677 }}</ref> and is frequently incorporated in study design for wetland delineation in [[hydrology]].<ref name="Attrill-2002" /> It has helped shape [[integrated landscape management]].<ref name="Sayer">{{cite journal| vauthors = Sayer J |title=Reconciling conservation and development: are landscapes the answer?|journal=Biotropica|date=2009|volume=41|issue=6|pages=649–652|doi=10.1111/j.1744-7429.2009.00575.x|s2cid=85171847 }}</ref>  Lastly, landscape ecology has been very influential for progressing sustainability science and sustainable development planning.  For example, a recent study assessed sustainable urbanization across Europe using evaluation indices, country-landscapes, and landscape ecology tools and methods.<ref>{{cite journal | vauthors = Shaker RR | title = The well-being of nations: an empirical assessment of sustainable urbanization for Europe. | journal = International Journal of Sustainable Development & World Ecology | date = September 2015 | volume = 22 | issue = 5 | pages = 375–87 | doi = 10.1080/13504509.2015.1055524 | s2cid = 154904536 }}</ref>

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<ref>{{Cite journal|last1=Manel|first1=Stéphanie|last2=Schwartz|first2=Michael K.|last3=Luikart|first3=Gordon|last4=Taberlet|first4=Pierre | name-list-style = vanc |date=April 2003|title=Landscape genetics: combining landscape ecology and population genetics|journal=Trends in Ecology & Evolution|volume=18|issue=4|pages=189–197|doi=10.1016/S0169-5347(03)00008-9|s2cid=2984426 }}</ref> and on how the quality of intervening landscape, known as "matrix", influences spatial variation.<ref>{{cite journal | vauthors = Storfer A, Murphy MA, Evans JS, Goldberg CS, Robinson S, Spear SF, Dezzani R, Delmelle E, Vierling L, Waits LP | display-authors = 6 | title = Putting the "landscape" in landscape genetics | journal = Heredity | volume = 98 | issue = 3 | pages = 128–42 | date = March 2007 | pmid = 17080024 | doi = 10.1038/sj.hdy.6800917 | doi-access = free }}</ref> After the term was coined in 2003, the field of landscape genetics had expanded to over 655 studies by 2010,<ref>{{cite journal | vauthors = Storfer A, Murphy MA, Spear SF, Holderegger R, Waits LP | title = Landscape genetics: where are we now? | journal = Molecular Ecology | volume = 19 | issue = 17 | pages = 3496–514 | date = September 2010 | pmid = 20723061 | doi = 10.1111/j.1365-294X.2010.04691.x | s2cid = 16435893 | doi-access = free }}</ref> 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,<ref>{{Cite book|url=https://books.google.com/books?id=UVPKCQAAQBAJ&q=landscape+genetics+concepts+methods&pg=PR11|title=Landscape Genetics: Concepts, Methods, Applications|last1=Balkenhol|first1=Niko|last2=Cushman|first2=Samuel|last3=Storfer|first3=Andrew|last4=Waits|first4=Lisette | name-list-style = vanc |date=2015-11-09|publisher=John Wiley & Sons|isbn=978-1-118-52529-6|language=en}}</ref> many with regard to how landscapes effect evolutionary processes, especially in human-modified landscapes, which are experiencing [[biodiversity loss]].<ref>{{cite journal | vauthors = Manel S, Holderegger R | title = Ten years of landscape genetics | journal = Trends in Ecology & Evolution | volume = 28 | issue = 10 | pages = 614–21 | date = October 2013 | pmid = 23769416 | doi = 10.1016/j.tree.2013.05.012 }}</ref>

== See also ==
{{Portal|Environment|Ecology|Earth sciences|Biology}}
{{div col|colwidth=22em}}
* [[Agroecology]] 
* [[Biogeography]]
* [[Conservation communities]]
* [[Concepts and Techniques in Modern Geography]]
* [[Ecology]]
* [[Ecotope]]
* [[European Landscape Convention]]
* [[Historical ecology]]
* [[Integrated landscape management]]
* [[Land change modeling]]
* [[Landscape epidemiology]]
* [[Landscape limnology]]
* [[Landscape planning]]
* [[Landscape connectivity]]
* [[Patch dynamics]]
* [[Total human ecosystem]]
* [[Sustainable landscaping]]
* [[Landscape architecture]]
* [[Land development]]
* [[Tobler's first law of geography]]
* [[Tobler's second law of geography]]
* [[Artificialization]]{{div col end}}

== References ==
{{Reflist|2}}

== External links ==
* [http://www.complexification.net/gallery/machines/substrate/index.php Computer sumulation "Substrate" launch applet creates fractal iterations that resemble urban streetscape. Algorithm written 2004 by Jared Tarbell]
* [https://www.landscape-ecology.org/ International Association for Landscape Ecology] 
* [http://www.napolisoundscape.com Napolisoundscape Urban Space Research]

{{Physical geography topics}}
{{modelling ecosystems|expanded=other}}
{{land use}}
{{Branches of ecology}}
{{Authority control}}

{{DEFAULTSORT:Landscape Ecology}}
[[Category:Landscape ecology| ]]
[[Category:Systems ecology]]
[[Category:Biogeography]]
[[Category:Ecological restoration]]
[[Category:Environmental soil science]]
[[Category:Environmental design]]
[[Category:Habitat]]
[[Category:Landscape]]
[[Category:Applications of geographic information systems]]