University of Copenhagen

Landscape Ecology

Christensen, Andreas Aagaard; Brandt, Jesper; Svenningsen, Stig Roar

Published in: The International Encyclopedia of Geography

DOI: 10.1002/9781118786352.wbieg1168

Publication date: 2017

Document version Version created as part of publication process; publisher's layout; not normally made publicly available

Citation for published version (APA): Christensen, A. A., Brandt, J., & Svenningsen, S. R. (2017). Landscape Ecology. In D. Richardson, N. Castree, M. F. Goodchild, A. Kobayashi, W. Liu, & R. A. Marston (Eds.), The International Encyclopedia of Geography: People, the Earth, Environment, and Technology (pp. 1-10). Wiley. Major Reference Works https://doi.org/10.1002/9781118786352.wbieg1168

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for Troll and his contemporaries to integrate Landscape ecology analytical perspectives from a range of otherwise discrete sciences in their analysis of landscape Andreas Aagaard Christensen processes. In this way, the development of Copenhagen University, Denmark landscape ecology, characterized by its specific focus on the ecological significance of spatial Jesper Brandt form and pattern, was closely associated both Roskilde University Denmark with the novel perspective provided by aerial Stig Roar Svenningsen photography and also with older geographical The Royal Library, Denmark perspectives of analysis inherited from classical cultural and physical geography. Troll had been trained in the traditions of The foundations of landscape ecology geography defined by Alexander von Humboldt and Carl Ritter, the two great initiators of The term landscape ecology (in the original the field as a modern science in the middle of German Landschaftsökologie) was first used in the nineteenth century. From them and their 1939 by the German geographer Carl Troll immediate successors he had picked up a certain as the name for a particular way of looking breadth of scope and an interest in the intercon- upon the landscape (Anschauungsweise)based nected nature of geographic phenomena, which on a holistic perspective integrating geograph- he emphasized in his writings. He had studied k ical and ecological insights into the nature of the work of Humboldt and Ritter extensively k terrestrial environments (Troll 1939). Troll’s and had been impressed especially by their ability previous research into the coupled relationship to correlate natural and cultural phenomena. between vegetation, environment, and land use From Humboldt’s scholarship he had understood had brought him to conduct extensive fieldwork the need to abstain from disentangling research registrations of vegetation patterns in landscapes objects from their environment, but rather to in Northern Europe. These experiences, in turn, delve into the multifaceted empirical relation- had inspired him to take up aerial photography ships linking otherwise seemingly disparate as an instrument to identify and describe spatial elements with larger patterns of distribution and units of vegetation cover, the heterogeneous causality. For Troll and his contemporaries, it pattern of which he was then able to relate was never enough to understand the nature of analytically to both social and environmental a species or habitat type in its own right – the processes of change. On these foundations, Troll aim was to explain how species and habitats proposed a science of landscape processes that interact with each other and other facets of their would be based on the new technology of aerial environment, with reference to actual, empirical photograph interpretation, to classify the earth’s histories of change. surface into discrete land units. On the basis On this basis Troll was increasingly criti- of such inventory classifications, it was possible cal of the growing specialization of academic

The International Encyclopedia of Geography. Edited by Douglas Richardson, Noel Castree, Michael F. Goodchild, Audrey Kobayashi, Weidong Liu, and Richard A. Marston. © 2017 John Wiley & Sons, Ltd. Published 2017 by John Wiley & Sons, Ltd. DOI: 10.1002/9781118786352.wbieg1168

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geography in his day. Since the late nineteenth socioecological analysis developed by Ritter century the universalist approaches advocated by through his work on the relationship between Ritter and Humboldt had been superseded in land use, culture, and civilization (Troll 1960). mainstream scholarship by an outspoken (and These concepts and instruments came to in Troll’s view unwarranted) reductionist stance inspire the development of landscape ecology, towards the organization of scientific knowledge. as a counterbalance to the fragmentation or Researchers within geography had sought to disintegration of geographic–ecological thought establish biological, physical, and demographic in the twentieth century that motivated land- modes of analysis as distinct fields of research, scape ecologists to establish their field as an and within these fields it had become the norm explicitly integrative science. In the 1950s and to seek for the smallest possible and most basic 1960s, landscape ecology consisted of a synthesis units of analysis in order to be able to study of geography with soil science and ecology. each environmental subprocess individually, as From the late 1960s especially, island biogeog- if in a laboratory. Troll saw this development raphy increased in importance (MacArthur and as an understandable but tragic detour from Wilson 1967). Parallel to this, a corresponding, the unified perspectives of earlier geography, spatially-oriented vegetation science developed, and one that made it increasingly difficult to and among conservation-oriented zoologists a correctly observe and describe the cohesive, strong school of dispersal ecology and metapop- interdependent nature of geographic phenom- ulation theory developed (Gilpin and Hanski ena. Therefore, Troll sought to conserve and 1991). The methods and perspectives of these k reformulate a unified perspective on terrestrial fields were combined by applying them to k surface processes. Hence, landscape ecology from the same spatially defined land units at various the outset was conceived as an approach to envi- scales – from single landscape elements or land ronmental research that was to be unrestrained units to larger complexes of functionally and by disciplinary and methodological dogma. historically associated sets of elements, up to When, in 1959, Troll was asked to address the whole landscapes defined by heterogeneous geographers of the British Association for the patterns of landscape elements. Advancement of Science on the centenary of the Figure 1 illustrates the type of method death of Humboldt and Ritter, he made his views for spatial analysis that is typically used. on contemporary geography remarkably clear. In It allows researchers to organize the many his speech he located the inspiration for a range mono-disciplinary insights flowing into land- of seminal landscape ecological concepts and scape ecology into a common empirical instruments within the work of his two academic understanding of landscape dynamics, by refer- idols. These included: forms of spatial pattern encing all relevant explanatory understandings analysis, which Troll became inspired to apply to a common set of land units and the landscapes and further develop upon reading Humboldt’s they constitute. In this way it becomes possible pioneering work on ecological zones and associ- to examine the spatial distribution and overlap ated patterns of plant distribution; the method of of explanatory understandings spatially based regionalization developed by Ritter, which Troll on map analysis. As the field developed, this employed to classify heterogeneous areas (i.e., basic method of integrating different perspec- landscapes) into analytical units based on similar tives by way of a common spatial framework form, genesis or character; and types of early remained unchanged, even though the range

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D C

0120.5 Kilometers Landscape patterns N Forest oundary b rural areased surfaceilt up areas w u Grasslands B Plo Open Land unit Figure 1 Landscape patterns analysis typically starts with a mosaic of vertical aerial imagery such as the sample shown here (left). Through imagery analysis supported by fieldwork observations, the remote sensed data are transformed into a classified map of internally homogeneous land cover units (center). Considered as a whole, the land cover units form a set of heterogeneous landscape patterns (right), each indicating a particular processual relationship between life forms on the earth surface (including humans) and their geoecological basis. The k sample shown here illustrates how rural land use is adapted to the geoecological conditions in an agricultural k landscape near Skive in Northern Jutland, Denmark. In the north there is an intensively used and closely settled agricultural landscape dominated by arable fields situated on well-drained loamy soils (A). A narrow erosion valley covered by grassland habitats intersects the arable land from the north, draining into a broad glacial valley of low-lying waterlogged soils covered by paddocks and remnants of moorland at the center of the area (B). South of the central valley there is a relatively complex pattern of agricultural land use, which is adapted spatially and functionally to the heterogeneous soils of the area, characterized by a variety of gravel, sand, and clay deposits (C & D). Analytical approaches of this kind, which link spatial pattern with landscape functionality and landscape history, are used within landscape ecology to integrate perspectives from ecology, soil science, cultural geography, sociology, and the humanities into a common, spatially explicit framework of analysis. On this basis, otherwise potentially incongruent theories explaining landscape dynamics can be tested and compared. Source: Imagery recorded by the Royal Airforce in 1980, held by the Royal library of Denmark.

of disciplinary components included within it management practices through applied research. broadened considerably over time. One of the major driving forces for this develop- When the various adverse effects of modern ment was the realization that ongoing processes land use development and associated destructive of industrialization in the agricultural sector ecological transformations came under increas- were leading to widespread habitat fragmenta- ing public scrutiny in the 1960s and onwards, tion and loss of ecosystem services in European many landscape ecologists saw a potential in and postcolonial landscapes. These changes their field to provide integrated solutions for were pioneered in Eastern Europe, where the development of more sustainable landscape they were associated with centrally-planned

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collectivization of agricultural properties into landscape context. Such research demands a specialized, large-scale agroindustrial units. In highly developed model for integrative research, Western Europe and North America, agricul- because human landscape management is a pro- tural industrialization developed later, in most cess that mediates both between the ecological cases driven by a combination of government and social sides of human existence and between subsidies and competition among producers, so natural and anthropogenic facets of ecosystem landscape ecology as an applied scientific practice functioning. As the German geographer Ernst developed more slowly and later here than in Neef, one of the founders of socioecological Eastern Europe (Ruzicka and Miklos 1990). research within the field, expressed it: “The transfer from societal changes to the natural Cultural landscapes and holism systems is based on the spontaneous effect of the laws of nature, whereas the transfer from the societal area to the regulation of natural processes The growing influence of applied perspectives is achieved by cultural forces, human perception meant that landscape ecologists came into sus- and decision making” (Neef 1984, 6). As such, tained conflict with traditional ideals of scientific cultural landscapes are a human construct and a practice, leading some researchers to define biophysical system at the same instance, and any their field as an explicitly action-oriented, attempt to analyze, theorize or interfere with solution-driven practice integrating scientific landscape dynamics must be able to account discovery with direct intervention in landscape for both of these dimensions. Within landscape management practices. This has remained a k ecology the challenge of accounting for these k widespread perspective within landscape ecol- two sides of landscapes in an integrated way ogy, which now includes a comprehensive array has motivated the formulation of a landscape of planning and policy-related fields of research. concept where the landscape is seen as a holon: In addition to these developments, landscape an assemblage of interrelated phenomena that ecological research has also extended in the direction of historical and social disciplines. together forms a complex whole, which is This has taken place gradually as researchers “more than the sum of its elements” because “all within the field have cultivated an increasingly parts are internally related to each other by the integrated relationship with scholarship from the general state of the whole” (Naveh 2000, 11). human and social sciences, which was considered This perspective has become a characteris- critical component to include within landscape tic feature of landscape ecological research. It ecology to fully appreciate the complex trajec- explicitly challenges both the basic division tories of change in human-dominated cultural between anthropogenic and natural phenomena, landscapes. as well as the mono-disciplinary reductionism Cultural landscapes have remained a dom- prevalent in mainstream science, by claiming inant research object throughout the history that the socioecological processes constituting of landscape ecology, which has progressed landscape holons and their constituent parts from descriptive accounts of human–landscape cannot be understood without accounting for interaction to more intricate attempts at under- the multifaceted relationships that unite indi- standing, conceptualizing, and quantifying the vidual elements into a spatially and functionally nature of socioecological relationships in a integrated whole.

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In terms of empirical work, such relationships It was the advent of airplanes in the early have been investigated by focusing on the pro- twentieth century that kicked off what was to cesses or “functions” linking ecosystems with become an explosion of vertical aerial photog- each other and with human agency. In this raphy. At first, images were recorded in limited way, empirical evidence of the organization of numbers and with rudimentary instruments, landscape holons is collected by characterizing but technological development and funding for and mapping the extent, magnitude and coupled recording campaigns soon became fueled by interplay of ecosystems processes (ecosystem public investment as the potentials for military functionality), social processes (societal func- use of the imagery became evident. During tionality), and intentional practice (transcending World War I recording efforts grew rapidly. functionality) within landscapes (Brandt and Mobile laboratories for photographic processing Vejre 2004). By integrating intentional human became available in 1915, making it possible practices related to human culture, cognition, to analyze images swiftly at locations along the belief, planning, and decision-making with front lines. This meant that aerial photography respect to landscapes into the field on equal became a critical technology to the war effort, terms with other more tangible types of func- and the increasing entrenchment of the western tionality, landscape ecologists were able to front raised a demand for intensive small-scale outline a broad-ranging holistic science devoted scrutiny of long stretches of frontline, to a point to examining the full interplay between socioe- where at the height of the war the French side cological processes and biophysical patterns in alone was processing more than 10 000 images k landscapes (Nassauer 1997). each night. In the period after World War I, k images and recording technology became available for other uses, forming a growing resource Aerial vision: a new perspective in the for landscape research, and a broad spectrum of study of landscapes researchers, planning, and policy professionals picked up the new technology. From the outset, landscape ecology was char- The idea of an all-encompassing birds-eye acterized by a visual approach to analyzing view of the earth’s surface goes back to antiquity landscapes from a birds-eye perspective, making and cartographic representations of the earth it possible to detect minute nuances in land as seen from above formed part of established cover patterns, while at the same time retain- practices within the scientific community long ing a distanced landscape-scale overview. This before airplanes made it possible to actually perspective was made possible mainly due to see the earth’s surface from above. In spite of the new technology of vertical mono- and this, however, it had an unanticipated effect on stereographic aerial photography, which became landscape research when actual remote sensed gradually more widespread in the period after imagery became available. Earlier map-makers World War I. The technology of aerial pho- based their delineations of areas, lines, and point tography had been pioneered with recordings features on fieldwork observations collected on from balloons over Paris in the 1850s and in the earth’s surface, which were then transposed Boston some years later, but due to navigational onto a spherical or flat medium in order for challenges the method proved unsuited for patterns of landscape elements to be made systematic recordings of large areas. available for visual inspection “as if” seen from

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above. In this sense, all earlier maps reflect a analysis. This spurred a rapid development of spa- form of inquiry where research design, sam- tial analysis tools within the field, supported by pling, and data collection precedes the rendition the new technology of user-centered computing, of raw data spatially. In such mapping processes, which became available from the 1970s onwards. the data becomes spatial in coverage only by way As such, landscape ecology became one of of analysis, not beforehand. several arenas where geographical information When remote sensed imagery eventually systems (GIS) and associated geostatistical tools became available, it represented an altogether were pioneered and developed. In conjunction different type of resource for map-makers than with these efforts, an array of computational previous types of spatial data. Mosaics of aerial approaches and indexing-methods for measuring imagery offered researchers a continuous field and comparing the configuration, complexity, of spatial data to begin with, making the earth’s and diversity of land cover patches was devel- surface an observable research object in its own oped, including the important Fragstats software right, rather than a canvas on which to drape package that empowered individual researchers existing observations. And when series of images to conduct quantitative spatial analysis (McGari- of the same areas at different points in time gal and Marks 1995). This initiated a shift in were recorded, it also became possible to classify landscape ecology away from analogue pattern processes of landscape change by way of direct recognition towards the use of computational overlay analysis. This was the key condition for tools able to quantitatively assess landscape pat- the development of early landscape ecological terns and associated processes across vast expanses analysis methods, and for the particular concept k of land. k of landscape processes that still characterizes the field. In the period since World War II, the range, Current methods coverage, and resolution of remote sensed imagery available for scientific analysis has grown rapidly. The Cold War, which succeeded World Current landscape ecological methods involve War II and culminated with the formal dissolu- a combination of fieldwork and analysis of tion of the USSR in December 1991, meant that imagery to identify the relationship between American and Russian intelligence gathering patterns and processes. Pattern in this sense refers efforts were in constant preparation for armed to heterogeneity in the horizontal dimension conflict. New, more precise recording devices of the landscape, i.e. differences between land and new platforms for carrying them, such as the units, while ecological processes typically transpire U2 high altitude jets and the Corona and Landsat within a single land unit and refer to the vertical satellite programs, are examples of the tremen- relationships between organisms and their abiotic dous development of remote sensing technology environment. The various approaches within that took place. When imagery from these the field relate to either of these two parallel programs was gradually declassified and older analytical perspectives: (1) the chorologic, dealing platforms began to be replaced by new in a suc- with the horizontal patterns and processes of cession of technological improvements, a broad land units on the earth surface, and (2) the range of new materials were added to the reper- topologic, dealing with the vertical processes of toire of data available for landscape ecological energetic, informational and material exchange

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between organisms within their habitats (Neef ways. Key parameters include the connectivity, 1963; Zonneveld 1989). diversity, and composition of landscapes, which In a chorologic perspective the primary dimen- have been shown to be systematically associated sions of the landscape are its geometrical and both with types of human land-use practices temporal characteristics. Here landscape dynam- affecting landscapes, with biodiversity, and with ics are approached analytically by measuring ecosystem functioning within patches. alterations in the shape, extent, and distribution or pattern of land units in the landscape, and by analyzing exchanges occurring between indi- Landscape ecology today and its vidual units. Trajectories of landscape change challenges through time can be appraised by determining the rate of change, its frequency, and magnitude Landscape ecology has grown to become a (Antrop 2000). widely recognized approach to environmental In a topologic perspective the character of a research, distinguishable from other scientific land unit is investigated by examining its internal traditions by the type of spatially explicit, inter- functioning as defined by the societies of species disciplinary, and empirically-focused analytical within it and its geoecological potentials in perspectives detailed above. By 1980 researchers the form of substrates, climate, and hydrology, from across the world had become involved including the flows of information, matter, and in the development of the field, with a world energy taking place through vertical vectors conference convened in the Netherlands in k within it. 1981. The following year the International k Today, the patch-corridor-matrix model intro- Association for Landscape Ecology (IALE) was duced by Richard T.T. Forman and Michael established as a focal point for dialog and mutual Godron has become the most widespread exchange of ideas by organizing regular con- conceptual model with which to approach ferences and through publication of newsletters pattern-process relationships (Forman and and journals. These organizational activities take Godron 1986). It consists of a spatial language place at international, regional, and national designed to describe landscape patterns and levels through local chapters of the associa- related processes. The basic idea of the model tion. They have proven to be instrumental for is that landscapes are made up of a mosaic of the further development of landscape ecology, patches (areas differing from their surround- because the local working environment of many ings), connected by corridors (strips of land landscape ecologists is delimited by traditional that infiltrate the landscape and support flows mono-disciplinary or sectoral organizational of information, matter, and energy) in a matrix structures and associated domains of expertise (defined as the dominant, most extensive, and within the academic community. Therefore, coherent landscape element type) (Forman most landscape ecological research relies on the 1995). In this perspective a landscape is defined application of a combination of skills and insights by the pattern formed by patch, corridor, and from researchers and practitioners who retain matrix elements repeated throughout its extent. their commitment to one or more disciplines By measuring the size, shape, and distribution while contributing to landscape ecology through of these three types of elements, landscapes cross-, inter- and transdisciplinary cooperation. can be compared quantitatively in a number of On this basis, a substantial literature on how to

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facilitate interdisciplinary exchange and organize the others, namely that between approaches action research has developed within landscape advocating theoretical pluralism on the one ecology, which has come to form a significant hand, and approaches characterized by exclusive scientific contribution in its own right (Tress, theories of truth and validity on the other. This Tress, and Fry 2005). appears to be a division line that cuts across other But, at the same time, landscape ecology is also variations within the field, and it has become haunted by its fragmented character, and one of particularly clear with respect to research identi- the most persistent threats to its continued suc- fied either with a strict positivist stance or with cess is the challenge of translating concepts and a constructivist conception of truth. methods that have been harvested eclectically This may illustrate that while landscape from other disciplines into a common analytical ecologists have succeeded in reconciling their framework. These efforts are made increasingly multifaceted perspectives on the landscape when difficult by the fact that the field itself has become dealing with tangible empirical problems, policy subdivided to some degree, because researchers advice, and action research, it has proven more from different parts of the world are basing their difficult to resolve differences pertaining to analytical efforts on differing theories of science the variety of underlying, a priori theoretical and validity, and thus also on different criteria for foundations within the field. Landscape ecology scientific achievement within the field. Today, was united not by a common theoretical debate mainstream American and European approaches but rather by a set of methods and research differ in research priorities and theoretical per- interests, and a fascination with a common k spectives for example, which is reflected in books empirical domain. At times this has been a great k designed to provide an overview of the field from advantage to landscape ecology, which has been either of the two perspectives. The introductory able to include and cultivate a host of different text by Bastian and Steinhardt is a European perspectives, but it also makes it a challenge to example, while the reader compiled and com- uphold a unified perspective able to make good mented by Wiens et al. is an American parallel on Carl Troll’s original ambition to understand (Bastian and Steinhardt 2002; Wiens et al. 2007). landscapes holistically. In a global perspective, the main differences This would entail the further parallel develop- between perspectives within landscape ecol- ment and amalgamation of three existing subject ogy, including those between American and areas within landscape ecology. European scholars, have tended to reflect underlying disparities concerning: (i) the way in 1 Research focusing on the basic, persis- which human interference or engagement with tent structural and processual character of landscape processes is handled analytically and landscape types and processes of landscape theoretically; (ii) the degree to which researchers change, which provides insights into the lean towards epistemologies derived from the long-term natural history of landscapes, human sciences and/or the natural sciences; and enabling society to better understand and (iii) the degree of practical engagement with adapt to general conditions for land-use landscape management and policymaking. management. When landscape ecological research is reviewed 2 Studies of historic and actual, anthropogenic along these lines it is clear that a particular dispar- and natural landscape structures and their ity within the field tends to inform and support development through time, which illustrate

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past and present relationships and barriers Gilpin, M., and I. Hanski, eds. 1991. Metapopula- within landscapes, supporting improved tion Dynamics: Empirical and Theoretical Investigations. understandings of how to conserve and/or London: Academic Press. improve valuable landscape resources. MacArthur, R. H., and E. O. Wilson. 1967. The The- 3 Studies of the cultural and ideological ory of Island Biogeography. Princeton, NJ: Princeton dimensions of human engagement with University Press. landscapes and associated types of man- McGarigal, K., and B.J. Marks. 1995. agement practices, making it possible to Fragstats – Spatial Pattern Analysis Program for understand the sociocultural and politi- Quantifying Landscape Structure. Corvallis: Oregon cal background for sustainable landscape State University. Nassauer, Joan Iverson, ed. 1997. Placing Nature: Cul- management. ture and Landscape Ecology. Washington, DC Island Press. The main challenge for landscape ecology, Naveh, Zev. 2000. “What Is Holistic Landscape Ecol- therefore, is not to focus or broaden its scope, ogy? A Conceptual Introduction.” Landscape and but rather to improve the way in which insights Urban Planning, 50(1–3): 7–26. DOI: 10.1016/ gained in one area of the field are communi- S0169-2046(00)00077-3. cated, compared, and combined with insights Neef, E. 1963. “Topologische Und Chorologische from other areas. Arbeitsweisen in Der Landschaftsforschung [Topo- logic and Chorologic Approaches in Landscape Research].” Petermanns Geographische Mitteilungen, k 107(4): 249–259. k Wbieg0803 SEE ALSO: Cultural geography; Landforms Neef, E. 1984. “Applied Landscape Research.” Paper Wbieg1144 and physiography; Landscape Dedicated to the Participants of The First Interna- Wbieg0794 tional Seminar of The International Association for References Landscape Ecology (IALE), Roskilde, Denmark. Ruzicka, M., and L. Miklos. 1990. “Basic Premises and Methods in Landscape Ecological Planning and Antrop, Marc. 2000. “Background Concepts for Inte- Optimization.” In Changing Landscapes. An Ecolog- grated Landscape Analysis.” Agriculture, Ecosystems ical Perspective., edited by Isaak S. Zonneveld and & Environment, 77 (1–2): 17–28. DOI: 10.1016/ Richard T.T. Forman. New York: Springer Verlag. S0167-8809(99)00089-4. Tress, Gunther, Bärbel Tress, and Gary Fry. 2005. Bastian, Olaf, and Uta Steinhardt, eds. 2002. Develop- “Clarifying Integrative Research Concepts in ment and Perspectives of Landscape Ecology. Dordrecht: Kluwer Academic. Landscape Ecology.” Landscape Ecology, 20(4): Brandt, J., and H. Vejre. 2004. “Multifunctional 479–493. DOI: 10.1007/s10980-004-3290-4. Landscapes – Motives, Concepts and Perspectives.” Troll, Carl. 1939. “Luftbildplan Und ökologische In Multifunctional Landscapes – Theory Values and Bodenforschung : Ihr Zweckmäßiger Einsatz Für History, edited by J. Brandt and H. Vejre, 13–31. Die Wissenschaftliche Erforschung Und Praktis- Ashurst, UK: WIT Press. che Erschließung Wenig Bekannter Länder [Aerial Forman, Richard T.T. 1995. Land Mosaics: The Ecology Imagery and Ecological Earth Science].” Zeitschrift of Landscapes and Regions.Cambridge:Cambridge der Gesellschaft für Erdkunde zu Berlin, 7: 241–298. University Press. Troll, Carl. 1960. “The Work of Alexander von Hum- Forman, Richard T.T., and Michel Godron. 1986. boldt and Carl Ritter. A Centenary Address.” The Landscape Ecology. New York: John Wiley & Sons. Advancement of Science, 16(64): 441–452.

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