Patterns within patterns: fractals and landscape ecology It should be noted that due to copyright laws, no actual graph or image from any book or journal appears on this webpage. All figures below are simplified representations of the actual images. What are fractals? A central goal of landscape ecology is to detect patterns through space or time and to extrapolate those patterns across multiple scales (Urban et al. 1987, Turner 1989, Turner et al., 2001). Fractal geometry is a method used to aid in achieving these goals. This approach is valuable to the field of landscape ecology not only because few tools are currently available that explicitly address and measure scale, but it additionally allows investigators to generalize a pattern across a range of scales. This detection of patterns can then enable investigators to entertain questions concerning the processes creating the observed patterns and to look for these processes at the appropriate scales. Analytically, the fractal dimension (D) is derived from the slope of a double log plot and describes shape complexity. It is a statistical descriptor like the mean and mode and therefore does not necessarily explain the process in question. Rather, it provides a way of describing a geometric shape. Any linkages made between D and an ecological process results from experiments studying the process of interest, not from D alone. For an object to be considered fractal, two requirements must be met. The first requirement is that a fractal must be “a shape made of parts similar to the whole in some way” (Mandelbrot 1982). As the grain of measurement increases or decreases for the self-similar object, the shape or pattern tends to repeat itself. Mandelbrot developed a broad definition for fractal geometry because of its usefulness as an analytical tool in fields such as physics, biology, mathematics and ecology. Furthermore, the definition purposely does not differentiate between exactly self-similar objects or statistically selfsimilar objects. Exact self-similar structures, such as the Sierpinski carpet and triangle (figure 1), only exist in the mathematical sense. In nature, there is an inherent randomness that prevents perfect mathematical fractals from forming. What we can observe in landscapes are fractal shapes and dimensions that are statistically self-similar. In statistical self-similarity, a portion of an object or phenomenon will look similar to the whole if enlarged and/or reduced to the proper scales (Leduc et al. 1994). This definition implies that a fractal relationship found in nature will not hold across all scales but rather will only pertain to a certain range of scales. Figure 1. Recreated Sierpinski Carpet and Sierpinski Triangle. Both are examples of mathematical fractals. The second requirement for a fractal object is that it must have a non-integer dimension. In simple Euclidean space, a point has a dimension of zero, a line a dimension of one (length), a polygon a dimension of two (length and width), and a volume has three dimensions (length, width and height). Many objects in nature, however, do not fit so neatly into one of these categories. A landscape surface with ridges is more than two dimensional, but since those ridges are not found everywhere on the surface, it is not entirely three-dimensional either (figure 2) (Li 2000). Its true dimension is some fraction between two and three, which is where the term fractal originated (Mandelbrot 1982). Figure 2. The surface above fills more than just the two-dimensional plane, but the mountains do not fill the entire three-dimensional space. This surface can therefore be described by some fractional dimension between two and three. Fractals and Landscape Ecology: A Land Use Case Study Using fractal geometry, patterns can be described across multiple spatial scales. Any generalizations made across scales, however, only hold true where D does not change. As the slope changes D will also change, indicating that a new set of processes is becoming important in determining what is observed in the landscape. Krummel et al. (1987) used this idea of changing slopes as an indicator of different processes operating in the landscape. The perimeter/area method (link to a page of links) was used to derive the fractal dimensions of forest patches in Mississippi and Louisiana. Once D values were plotted against the forest patch size, it became apparent that two distinct processes were affecting the size of the deciduous patches (figure 3). Figure 3. The fractal dimension (D), as a function of forest area. The break in D at approximately 22 m2, illustrates how different processes are affecting the size of patches found. Image adapted from Krummel et al., 1987. Below a patch size of approximately 22 m2, the values of D are quite low. Low D values signify that geometric shapes are relatively simple and were most likely a product of anthropogenic processes constraining the patch shape and size. After 22 m2, D increases rapidly before becoming relatively constant again reflecting larger patches with more complex shape. These patches are a result of natural processes and topography creating landscape patterns as opposed to the smaller more simple patches shaped by human influences (Krummel et al. 1987, De Cola 1989, O’Neill et al. 1988). The use of fractals to describe a landscape is not limited to only spatial scales. Fractal geometry can also involve the temporal study of changes to a landscape. For example, fractal dimensions may provide insight into the human impact of urban sprawl (link to Heather) on natural ecosystems. Using the perimeter/area method (link to page of links), D can be found for different land use categories by using a time series of two remotely-sensed images (link to Ken) at a given location. This method of change detection (link to Jon) can then be used to examine changes in the shape complexity of land use types over many temporal scales (link to Jennie) (De cola 1989). Fractals and Landscape Ecology: Organismal Case Studies A fox may view an entire forest as its landscape while an insect, whose entire life cycle begins and ends on a singe tree within that forest, perceives the landscape at a much finer scale. As a landscape ecologist, it is important to understand at what scale the organism of interest perceives the landscape and responds to landscape features. Finding and defining the proper scales for organisms has implications for many fields in ecology including restoration ecology (link to Polly), conservation biology (link to Jenn) and even the modeling of avian communities (link to Alison). To better understand how organisms view their surroundings, Kimberly With (1994) measured and plotted the path length of three different sized grasshopper species using different ruler lengths. From these plotted points, the fractal dimension for each species was determined. The three species of grasshopper had different values of D, indicating that they took different paths of varying complexity. Higher D values signify that a grasshopper species took a more convoluted path, while lower values of D reflect a more linear path through the grassland habitat. The different D values for the three grasshopper species imply that each species views the landscape differently. Species with high values of D may consider a portion of their habitat inhospitable which could be why they took a more convoluted path. Alternatively, because the different species were of different body sizes, a blade of grass that a small species may have considered too large to travel over might be ignored by a larger species. Not only does this mean that a species with a lower fractal dimension can travel through a landscape more quickly and directly, but it also suggests that larger species are integrating landscape information at a broader scale (With 1994). That the species with the lowest D did not travel below –3 cm (this is a natural log number) indicates that it may be assimilating landscape information at a scale above –3 cm and does not consider habitat below this number. It should be noted, however, that because the fractal relationship in Kimberly With’s study is only over a small spatial scale, it may not truly describe a scaling relationship. It has been suggested that a scaling relationship must be constant over two orders of magnitude to be considered statistically self similar (Milne 1991). In a different application of fractals to organisms, Milne (1997) mapped habitat availability at different scales to better understand how two different species that varied in body size may view the same landscape. With an equation containing a fractal exponent relating body size to home range area, Milne determined the home range area of a pocket gopher and a black tailed jack rabbit. Using a remotely sensed image (link to Ken), a moving window (link to page of links) was placed over the digital map, and each pixel was classified as either grassland habitat or forest habitat. Moving window sizes were based on each species’ characteristic home range length, as found from the body size/home range area equation. The visual result of such an experiment provides a striking difference between how both rabbits and gophers might view the same habitat (figure 5). Figure 5. Habitat as viewed by the gopher (A) and rabbit (B). Areas in white represent available habitat while dark areas represent forest/inhospitable habitat. Image adapted from Milne, 1997. The boundary between grassland and forest is much more distinct for the gopher (figure 5A) as compared to the rabbit (figure 5B). Because the window used to map the gopher habitat was similar to the resolution of the image, the grassland habitat boundary is much more defined. The sharp boundary indicates that this particular gopher species may be more constrained by the forest edge habitat than the rabbit species. Since the rabbit is a larger animal, its home range incorporates a much larger area and it might be able to utilize some of the edge habitat that the gopher may find inhospitable. Therefore, the boundary for the rabbit species is softer, resulting in a more fuzzy definition of rabbit edge habitat. Similar analyses may be of use in other studies of herbivory by different species (link to Fleur) to better ascertain how different herbivores view the same area and consequently how much habitat each species could have available to it. Fractals and Landscape Ecology: Multifractals Although multifractals, an extension of the concept of fractal geometry, are becoming more common, they are only briefly mentioned here. Multifractals assume that no single D describes patterns observed in natural landscapes. Rather, there is a spectrum of scaling exponents creating the observed landscape pattern (Scheuring et al. 1994, Feder 1988, Milne 1991). Different agents of disturbance, for example, can have different scales at which they affect the landscape. Ecologists could therefore look at fractal moments, analogous to the mean, variance, skewness and higher moments based on a probability distribution, to see how the various pattern forming factors change with length scale (Milne 1991). Fractals do not explain all the patterns that are observed in nature, but they do identify the presence of patterns at multiple scales. Part of the appeal of fractals is that a single statistic can be used to describe potentially complex patterns in natural environments. Furthermore, because shapes with fractal dimensions are prevalent in many disciplines, D also provides a means for scientists in various disciplines to communicate with each other. The addition of fractal geometry should enable landscape ecologists to better entertain questions relating scale to various patterns, processes and predictions. The use of fractal geometry, however, should only be viewed as another tool to be used by landscape ecologists to aid in answering questions relating to scale, not the answers in and of themselves. Bibliography Bradbury, R. H., R.E. Reichelt and D.G. Green 1984. Fractals in ecology: methods and interpretation. Marine Ecology Progress Series 14: 295-296. De Cola, L. 1989. 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