ACKNOWLEDGMENTS Egolfs V. Bakuzis, Professor Emeritus at the University of Minnesota, College of Natural Resources, collected most of the information upon which this review is based. We express our sincere appreciation for his investment of time and energy in collecting these articles and books, in organizing the diverse material collected, and in sacrificing his personal research time to have weekly meetings with one of us (N.L.) to discuss the relevance and importance of each referenced paper and many not included here. Besides his interdisciplinary approach to the scientific literature, his extensive knowledge of forest ecosystems and his early interest in nonlinear dynamics have helped us greatly. We express appreciation to Kevin Nimerfro for generating Diagrams 1, 3, 4, 5, and the cover using the programming package Mathematica. Craig Loehle and Boris Zeide provided review comments that significantly improved the paper.

Funded Central

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USDA Forest Minnesota.

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CONTENTS i_ Introduction ....................................................................................................... I 2 2 3 4 4 4 6 6 7 7 7 9 9 12 12 13 14 15 15 16 16 16 17 17 18 18 19 20 20 22 22 23 24 26 27 28 30 31 32 34

2° Description of Fractals ..................................................................................... Fractal Objects ................................................................................................. Fractal Processes .............................................................................................. Iteration ........................................................................................................... Initiator-Generator ........................................................................................... Classical Fractals ............................................................................................. I-Julia and Mandelbrot Sets; the Complex Plane] ................................................. Natural Fractals ............................................................................................... 3, Fractional Dimension ........................................................................................ Dimension in General ....................................................................................... The Fractal Dimension ..................................................................................... I The Hausdorff and Other Dimensions] ............................................................... Computing the Fractal Dimension .................................................................... 4. Fraetals Scaling and Scale ........................................................................................... .............................................................................................

in Nature

I Other Scaling Techniques and Their Relation to Fractals I................................. Scale-free Fractals ........................................................................................... 5. Fraetals and Self-similarity ............................................................................ The Parts and the Whole .................................................................................. Local vs. Global ............................................................................................... Fraetals and Complexity ..................................................................................

6.

Jaggedness and Boundary ............................................................................... Complex Pattern and Self-organization .... ........................................................ Noise ............................................................................................................... 7. Applications to Forest Systems ....................................................................... Leaf Shape ....................................................................................................... Tree Branching ................................................................................................ Tree Crowns .................................................................................................... Tree Models ..................................................................................................... Vegetation Patches .......................................................................................... Size and Density of Organisms Related to Habitat ............................................ Species Diversity ............................................................................................. Animal Movement ............................................................................................ Forest Disturbance .......................................................................................... Modeling Soils with Sierpinski Carpets ............................................................ Spatial Heterogeneity and Sampling ................................................................. [D and Shoreline Erosion_ ................................................................................. Data Storage and Compression ........................................................................ 8. Fraetal Applications in Managing Forest Landscapes ...................................

Bibliography

...........................................................................................................

The Fracta] Forest: Fractal Geometry and Applications in Forest Science

Nancy

D. Lorimer,

Robert

G. Haight,

and Ro|fe

A. Leafy

1. INTRODUCTION Fractal geometry is a new branch of mathematics with roots in set theory, topology, and the theory of measures. As a young science (Bunge 1968), the theory of fractals is more like a collection of examples, linked by a common point of view, than an organized theory. But the mathematical underpinnings are developing; and researchers in other fields are applying the concepts and techniques, thereby extending fractal geometry beyond its mathematical beginnings (Mandelbrot 1983). Despite its newness, fractal geometry expands and sparks investigation in physics, chemistry, and biology because it offers something very important--tools for describing and analyzing irregularity. Kinds of irregularity can be classifled as new regularities, seemingly random but with precise internal organization. With fractals we see new patterns in the world (Stewart 1988b). The fractal world view is different from the world view exemplified by calculus, by Leibniz' "principle of continuity" (Mandelbrot 1983). Out tools for a forest that is smooth, continuous, linearly ordered, and at equilibrium have been Euclidean geometry, calculus, the Fourier

transform, and the Gaussian curve. But most of what we measure in the forest---crowns, roots, soils, stands, plant-animal interactions, landscapes--is discontinuous, jagged, and fragmented; and our tools, designed for smoothness, have trouble measuring them. Fractal geometry is a tool for discontinuity, a tool for the fractal forest. We are interested in mechanism as well as measurements. When we ask: What is the mechanism?, we traditionally mean: what is the linear interaction of two or more elements that cause something to happen? The mechanisms in fractal systems, however, are probably not linear cause-and-effect; they are complex scaling interactions (West 1987). Like relativity and quantum theory, fractal geometry brings the observer into the story. Consider the dimension of a ball of string. From far away, the ball looks like a dot (dimension = 0). Closer, it is a solid (= 3). Very close it is a twisted thread (= 1). Then closer yet it is a column (= 3). Closer still it is FLuetwisted hairs (= 1). Dimension in this example is a function of the observer's location (Briggs and Peat 1989). In sections 2 and 3, we will show how dimension is a function of the observer's choice of measuring unit. Euclidean geometry is one of the conventions in mathematics and science that is chosen for its utility {Giedymin 1991). But that utility is limited, when Euclidean geometry is faced with the complexity generated by nonlinear dynamical systems (Bohr and Cvitanovic 1987}. Fractal geometry is an alternative, a complementary, geometry. But as a new geometry is proclaimed, we need not feel uneasy aboutventuring beyond Euclid. Mathematician (and geometer) Poincar6 (1952) observed, "One geometry cannot be more

Nancy D. Lorimer is a Staff Entomologist with the Forest Pest Management Staff, State and Private Forestry, Washington, DC. Robert G. Haight is a Research Forester and Rolfe A. Leary is a Principal Mensurationist with the Mathematical Models of Forest Dynamics of the North Central Region research work unit, North Central Forest Experiment Station, St. Paul, Minnesota.

In section 8. Fractals may be the most promising direction for forest research in its task of representing the forest (Zeide 1990). wildlife habitat. had difficulty in defining them.g. These boxes can be skipped in an overview reading. . As a result. Our paper begins with a general description of fractals in section 2. Where needed in our review. the evidence for the potential of fractals is overwhelming (Zeide 1990). fractals have infinite detail. Damage patterns in forests and arrangements of tree crowns are fractals (Loehle 1983). the shape (the geometry) of the fractal does not change (Gefen 1987). more detailed information is presented inside a box near the general discussion. infinite length. it can only be more convenient. who called fractals to the world's attention. predicting total wood volume. and complexity. fractional dimension. scale. and complexity. using the Hausdorff-Besicovich measure of dimension. The simplest way to def'me fractals is via the self-similarity assumption. Self-similarity means that as magnification (the scale) changes. In assembling this section. the definition of "fractal" is still not definitive. the necessary and sufficient conditions for an object or a process to have fractal properties have yet to be determined. physiology. selfsimilarity. This review is a collection of fractal thought and its reference to many natural disciplines. The usefulness of the fractal concept stems from its ability to describe apparently random structures within a precise geometry (Orbach 1987). D. The purpose of our interdisciplinary approach is to make this wideranging material accessible to researchers in forestry. Besides self-similarity. there are mathematical complications. are very difficult to understand (Cipra 1989). This is not a fractal cookbook. Although there is fair agreement in the literature that the fractal properties are fractional dimension. mensuration. Mandelbrot (1967). So. scale independence. and geology. microclimatology. For example. with which fractals are usually described.. vegetation mapping. Fractal objects carmot be adequately described by two dimensions or three dimensions or n dimensions of whole numbers because fractal dimensions are fractions. In 1983 he approached the definition with the aid of measure theory. and number of branch tips. self-similarity. Despite many attempts. and they are generated by iteration (Briggs and Peat 1989). we focus on helping readers quickly understand concepts they can apply to their own research. we used a wide definition of forestry research. physics). We have used illustrations from work in forestry whenever possible. we offer thoughts on the research potential for fractals in forest resource management. Because of the fast-breaking nature of this new field. a ferromagnet is a good example of self-similarity. are fractals. chemistry.true than another. while a leaf can fllustrate fractal dimension. Although the danger exists that exciting new fields may raise false hopes (Mayer-Kress 1986). Mandelbrot (1983) offers a way to characterize fractals by the fractal dimension. Properties (Bunge 1979) can be component properties ("a tree has fractal shape") or relational properties ("stand boundaries have fractal shape"). We followed the semantic convention that ff an object or process has fractal properties. it is called a fractal. but ideas about fractals come from many disciplines (e. we have left out the full theoretical discussion these concepts deserve. although references to specific techniques are provided. 2. Section 7 describes specific applications to forest systems. stem surface area. The continuing development of fractal definition within measure theory does not hamper the utility of the concept. followed by more detailed exposition of four fractal properties in sections 3-6: fractal dimension (with a section on how to compute the fractal dimension)." Different geometries serve different purposes. Applications for fractals in forestry include forest ecology. that are nonuniform in space. a tree can be modeled as a fractal. DESCRIPTION Fractal OF FRACTAI_ Objects Self-similar objects that cannot be described in common Euclidean fashion. Rigorous definitions of topological and Hausdorff dimensions. soil science.

Vlcek and Cheung {1986) generated a one-pLxel-thick computer image of a leaf. That is. computed fkactal dimension of coastlines from 1. fractional dimension is more thoroughly described in section 3. the number of lines changes from 1 to 4 to 16 to 64. it is jagged. called the scaling factor. Because measurement of fractal objects depends on scale. like diffusion-controlled reactions with geometrical constraints (Kopelman 1988}.raThe generation of a Koch curve. the fractal dimension describes the relationship between vibrational 3 . That is. the value of the attribute diverges toward infinity. At each step the length scale changes by a factor of b = 3. n=1 n=2 the fractal 3 1. These kinds of structures are often heterogeneous. has fractional dimension. They measured the leaf outline with different measurement lengths. Start with a straight line divided into three equal segments. Mandelbrot suggested. processes may have fractal properties. on the size of the measuring unit. so does the perimeter of a leaf depend on the size of the measuring unit--in the above case. The Koch curve is self-similar: any portion magnified by a factor of three looks like the previous iteration. or irregular.15 to 1. also called fractal dimension. A coast_line. Fractal Fractals are geometrical of fractal objects The mathematical concept of fractal dimension follows. Diffusion processes (Orbach 1987). Divide each line segment into three segments. Replace the center segment with two segments that are each the same length as the original center segment. An amorphous solid has less than three dimensions because of the openness of the structure. Measuring a fractal object (e. nor is it two dimensional like a plane. the length of the line goes to infinity (Mandelbrot 1983). Just as the length of a coastline depends on the resolution of scale. a coastline is not a simple one-dimensional line. the perimeter of a maple leaf is not smooth. for example. There are "holes" in the solid. As the scale becomes infinitely fine. changes by a factor of N = 4. the usual statistical concepts (e. the perimeter of a leaf) depends on the scale of the measuring unit.g. and replace the center segment with two segments the same length as the segment being replaced. divide each line into three segments and replace the middle segment with two segments. the length of a Koch curve and other fractal curves like the coastline varies by the unit of measurement (Richardson 1960). For the Koch curve. n=3 /k n=4 _ .g. Now repeat the process with each of the four new lines. Fractal geometry can also be applied to planes and even to amorphous solids. The values for the leaf's perimeter differed depending on which measurement length did the measuring. as well (Orbach 1987).v%. and vibration distributions (Orbach 1987) have fractal dimensionality. In the diagram below. Diagram 1. or D= ln N/ ln b. then. With a video imaging system. on the number of pixels that mark off the perimeter of the leaf. The number of basic units. A Koch curve is the simplest way to show how the fractal dimension is derived (diagram 1).. sample mean) need to be re-thought when applied (Loehle to the measurement 1983). Continue for an infmite number of times. The ranges Because of fractal dimensionality.26 dimensionality is: Processes objects or constructs. D = In 4/ln [Gefen 1987). Repeat. but they may also have dynamical properties. the measurement lengths each had different numbers of pixels. The fractal dimensionality is: N= b D .25.. To use an example from the forest.Mandelbrot's famous example of a fractal is a coastline's rich twists and turns. meaning that it has a dimension somewhere between a line's one dimension and a plane's two dimensions. As the unit of measurement decreases. In the frequency distribution of vibrations.

Demonstrating the concepts. These techniques are applied to many physiological processes (West and Goldberger 1987). Fractal processes have many component frequencies. in the second column. The question becomes: What is the algorithm that produces this pattern? By first learning how fractals are generated. replacing the center segment with two segments. the statistical properties of the whole its parts (West 1987). and has dimension 2. 1986).frequency and the spatial extent of vibration. formed by the remaining points. A fractal process is one that cannot be characterized by a single time scale. we may better understand how to untangle them. In forestry we usually wish to untangle fractals. diagram 2 shows some formal examples of initiators---the beginning shapes--in the first column. For the latter. producing a broad spectrum of frequencies. we want to find the initiator and the generator. just like a fractal structure does not have a single scale length. the next figure (third column) has three images of the previous figure. Tree branching is an example of iteration in forestry. A Cantor set is constructed by removing the middle third of a line segment. self-similai_ty. The generators are the changes made to each line segment in the initiator. the (n + 1)st figure contains three copies of the nth figure. is a key idea in the whole field of dynamic systems and fractals (Devaney 1988). then removing the middle third of those segments. a technique that unravels the complexity of frequencies. On the first row of the diagram. The growth of a modular organism like a tree is determined by rules of iteration and the reaction of each growing point to the local conditions around it (Franco 1986). This happens ff the structure is "f'flled in" with atoms. the generators are sketched above the arrows {Arlinghaus 1985). The fractal dimension measures shape with shifting scale. Fractals are complex (they contain 4 There are other types of "classical" fractals. A Peano curve plane (Kramer (diagram 3) is a line that fills up a 1970). it measures self-similarity (Arlinghaus 1985). Fractals are also self-similar sets composed of modules or pieces similar to the entire set on lesser scales (Harrison 1989). The repeated change is called the generator. They are useful for illustrating fractal characteristics before moving on to more complicated forms. When the generator is applied a second time. has a fractal dimension between the zero dimension of a point and the one dimension of a line (Mandelbrot 1983). . That is. Initiator-Generator A fractal may be generated by performing a repeated operation on a beginning structure. Heterogeneity. the application of the generator to the initiator gives three copies of the initiator at reduced scale. such as the Koch curve and Julia sets (see box on page 6) that are heavily used in mathematical studies and in the development of theory about fractals. The beginning structure is called the initiator. Continuing on. Classical Fraetals Iteration Iteration. Computers iterate easily and are thereby useful in the generation and study of fractals (Harrison 1989). Cantor dust. and absence of a characteristic length of scale are therefore three properties applicable to dynamics as well as to structures. there is the category of statistical fractals. and continuing to remove middle thirds (Stewart 1989b). A modular organism has repeated units of structure at varying levels of organization (Halle et at 1978. Harper et al. are the same as that of much detail) yet simple (they can be simply generated) (Briggs and Peat 1989). A structure can have Euclidean mass properties and yet have fractal dynamic properties. repeating a process over and over. So a fractal time series can be investigated with spectral analysis. The Koch curve above was generated by the repetition of a single step. And besides geometric fractals and temporal fractals. complicated forms do not have to be generated by complicated processes. Because of iteration. but not all the atoms are participating in the vibrational activity (Orbach 1987). not generate them.

" is foUowed as each square is successively divided into four squares. r_ errors... dimensionality to less than a plane. Geografiska Annaler...S'e_Nexfes: K= 3.. "move the line to the center of each of four squares.ach [O-uesegment _v " . In the second step. rule.wGeneration of a Peano curve._Inf_ators and genm F... between 5 ... [-'] ['-] _ Diagram 3..) ii iii n = 1 n = 2 "J . S. _ _v.. 1985. is replaced by a new form.. and a third structure emerges.. Tyler and Wheatcraft (1990a) modeled soft types with Sierpinski carpets. h . (Reprinted with permission frorru" Lach. and thereby has dimension two.. Eventually the line fiUs the whole plane. the line segments in the new structure are replaced in the same way..._ j-x_. "_ "_ _ initiator. Fractals take a central place. The fractal dimenthe area of pores greater than a given size. -----_ n = 3 n = 4 Sierpinski carpets are generated by a recursion in which successively smaller holes are cut from the plane to give a self-similar pattem at levels smaller than the initial plane (diagram 4). 7 ] _erogon $OCO_Cl U J U_ " 1_" V . 67B: 83-88.Central |hill@lot Plate b'raetal _rlll " Iteratian to¢ogo_ . 4.. 2. Diagram o_-_.. The dark holes are the pore spaces among soft grains... The holes in the face of a Sierpinski carpet reduce its sions of these carpets can be used to estimate dimensions 1 and 2.. the generator.

with a mapping or a rule repeated over and over.Julia and Mandelbrot Sets. Equations can also be iterated to generate fractals. Natural fractals normally have some element of randomness.-. Different Julia sets are generated by changing the value of z0 when iterating the above relation.M. approach a limit. and partly understood... Forest fractals. They are excellent illustrations of a common theme in the study of fractais: very simple. and compute zl. nonlinear processes generate very complex structures (diagram Programs are available Mandelbrot sets. by humans. for generating and other fractals on 1986.. 5). not strictly self-similar at every scale. Iia IR ma @_ @ _."may be the most complicated figure ever to be studied. zl = c. three possibilities: Zncan Z2=Z12+c. starting from Zo = Xo+ i yo. The rich variation and form in the Mandelbrot set and Julia sets come about through nonlinearity within rigorous organization (Douady 1986). it The approach zo. Zn+_= Zn + C. The dimensional description of multifractals requires two or more dimensional exponents. Zl=Z02+C. z2. its definition. infinity." However.m. The dimension of these figures faUs between one for a line and two for aplane. are most likely multifractals.®ua. Natural Fractals Nature adds the random element to fractals. Examples of multifractals are complex surfaces and interfaces. the Complex Plane The examples of fractai generation in section 2 were generated by iteration of lines or figures. relation Start with a Step 0 Step 1 where both z and cf(z) =complex numbers. and Julia sets and the Mandelbrot set are classic examples of the results of iterating equations. stays bounded" (Strang 1986). remains remarkably simple: "M contains all numbers c for which the sequence z0 = 0.uGeneration ofa Sierpinski """"'"'""'" carpet. Deterministic fractals are the mathematical fractals discussed above. Mathematical fractals can be generated with a computer. not exactly the same at all scales. fractals in nature. Fractals not exactly self-similar. or it can do neither. Iterate. etc. personal computers (Strang Saupe 1988}._ Step 3 Remove the center square and then remove the center square from the remaining squares. can sequence. Repeat. The Mandelbrot set is related to the Julia set in the following way. c is varied rather than z0. but. but the new set M is in the cplane and is an "index" to all Julia sets. has.. . A Julia set is the set of points in the z0 plane for which zn stays bounded as n approaches infinity (Strang 1986). Strang (1986) explains how Julia and Mandelbrot sets are generated.. Peitgen and |. f(z)=z2+c. and fluid flow in porous media (Stanley and Meakin 1988). Each value of c leads to a Julia set in the Zo plane. Julia sets. he says. according to Strang. The relation is iterated as described above. Strang observes that the Mandelbrot set .. etc.||l||w| _R[]___ _l ___I" R I_I! lira U@ MlllillllmIDU Step 2 Diagram 4. Fractals generated with a computer program that includes random elements simulate natural phenomena more closely than fractals generated by deterministic methods (Saupe 1988). are called multifractals (Stewart 1989b). are z2 + c.

. Department of Agriculture.. yet dimension is elusive and very complex (Mandelbrot 1967). Natural C_racta \_ I) I i nes 1<o<2 _" L 0 a_ 200 '_0_ important properties of fractals---fractional dimension. The difficulties of defming dimension in general carry over to definitions of fractional dimension [Cipra 1991). When the number of measurement units is graphed against the size of measurement unit.. 1 "dusts" o _o < _ _r_. as mentioned in the previous section. and complexity.. 1..._ _ i i -... Euclidean dimensions are whole numbers. and 3 (volume) (diagram 6). Typical Euclidean objects have slopes of 1 (length). They are measures of the relative degree of complexity (Briggs and Peat 1989)...0 Un i t Diagram 6. The figures are generated by the iteration of simple. Boris.--The MandeIbrot set (upper figure) and a Julia set (lower figure).) 7 .... It is helpful to compare Euclidean dimensions and fractal dimensions (also see box on page 9)... Jl _ o. i 3.... 1990. 2 (area).. D . Forest Service. Irregularity and fragmentation cannot properly be understood by defining dimension simply as a number of coordinates. The loose notion of dimension turns out to have many mathematical ramifications (Mandelbrot 1983) and deFmitions {Mendes-France 1990). 4o.400. OR: U. Portland.--Dimension is the relationship between the size of measurement unit and number of measurement units. FRACTIONAL Dimension Geometry makes space... nonlinear equations. Pacific Northwest Research Station: 260-266... Geru Tectz Rep. Dimension is a strong center in the language of all geometries.. PNW-GTR-263.. their dimensions are fractions (Zeide 1990).. we describe four m '7. Natural objects fall between these slopes. despite the ambiguity about the underlying philosophy and mathematics.. Classical DIMENSION i 2o i i i i 2 ...0 S i ze of Measurement L 10. Fractal geometry and forest measurements..S. t. the dimension is read from the slope.ooo- X Natura surf_ces I Diagram 5.. i in General explicit our intuitions about Euclidean geometry is a first approximation of the structure of physical objects.The Fractal Dimension • .. self-similarity... Fractal geometry extends classical geometry with a new language (Barnsley 1989).. However.. In the next four sections.... fractal dimensions are numbers that precisely specify deviation from the expected scaling rule {Pagels 1985)... Fractal dimensions are fractions (Zeide. scale independence...

a wiggly line is identical to a circle. a straight line. Tsonis and Tsonis (b) Y . observer's yardstick (Cherbit 1990). For the four objects in the diagram. on and on the The observer and its dimension would approach 1. o E= 2. DE =3.. Tsonis.-_ Y3 _ _ x z (1987) looked at three kinds of dimensionality: Euclidean. both have dimension 1 .. This parameter is for fractal dimension // o. PoA. a crooked line in a plane. and fractat (D) dimensions of lines and surfaces.(a) In another comparison. 1-<D-< 2 (c) /. chooses the unit of measurement. topology cannot discriminate between crooked lines and straight lines (Mandelbrot 1983). The non-integral dimension may be a relative (d) or= 1. x DT = 1. topological (D_). topological. Fractals: a new look at biological shape and patterning.) . But this does not mean that measurement is arbitrary.. dimensions in the three different geometries are identical for only one object: the straight line. (Reprinted with permission frorrc Tsonis.. (Zeide 1990)._ =2. DE = 1. By defmition.__. a crooked line in a volume. Correspondence y between unit and length is maintained. 1-< _-3 0 Y notion. and the length of the object will depend on the unit chosen. and fractal (diagram 7). . 1987. A tangled worm in a stagnant pond has a trajectory with a dimension near 2. and a crooked plane in a volume. In topology. Topology is a branch of mathematics that treats form as flexible and compressible. Length becomes a process rather than an event. 2-_1_-_3 / ""_x Diagranl 7. a process controlled by a constant parameter unique calledeach object measured. 0= 1 o r = 1._Euclidean (D_). Dimension motion. 0 _ = 3. can depend on the object's state of the frame of the observer's reference.. The dimension of aa object can vary depending on the geometrical system in which it is measured. Perspectives in Biology and Medicine. 30: 35536I. A. The same worm in a current would have a stretched trajectory.

except that the Hausdorff dimension has been defined with more mathematical rigor (Harrison 1989). begin an estimate of the length of a crooked line by stepping dividers with step size st along the line. for another length estimate. and there are many examples of interest in the forest (e. then D will be greater than 1 by some fraction (Goodchild and Mark 1987). the very complex Peano curve completely fKls the plane and has dimension 2. For unequal probabilities. The more jiggles in the line. Repeat the process with a smaller step size. can be compared to Euclidean dimensions. This dimension takes the density into account. the number of pairwise correlations between points in a set scales as the size of the set is raised to some power. The relationship between the length of the line and the unit of measurement is predictable. giving a length estimate of nls_. the length of the line will differ. The capacity dimension is defined by boxcounting. the greater the increase in length between the two estimates. Calculating the fractal dimension becomes automated when the object is photographed by a video 9 . / log(sl/s2). For many fractals. Over a range of scales. is relatively simple. length measurements for a crooked line vary with the size of the measurement unit. the slope is the key. the information dimension will be less than or equal to the capacity dimension (Rasband 1990). Sometimes scaling properties emerge only asymptotically. Computing the Fractal D_zlension Loehle (1990) warned that Funding natural fractal dimensions can be difficult and computationally intensive. but computing the fractal dimension of a crooked line. depending on the unit of measurement. When the unit of measurement is plotted against length for several scales on log-log paper. then halving the step size or sampling interval will require precisely twice as many steps. is D = log N/log r where N is the number of measurement steps and r is the scale ratio (or self-similarity ratio).The Hausciorff and Other Dimensions By Mandelbrot's definition. Perimeters and outlines are simply closed crooked lines. but if the line is crooked. the line must be measured with different sizes of measurement units. The correlation dimension is less than or equal to the information dimension. r_s2. which may be greater than or equal to the previous one. Count the number nl of steps required to traverse the line. The Hausdorff dimension and the capacity dimension are inner fractal dimensions (Naudts 1988).. s2. or the local and global dimensions. the number of boxes increases by the fractal exponent. at the microscopic level or at the upper limit. As mentioned in the previous section. As set size decreases. The capacity measure depends on metric properties of the space. Therefore. The Hausdorff dimension. Very complicated fractals are difficult to measure. As the size of the box decreases. The information dimension depends on the natural probability measure of the set. A curve of D = 1 approaches D = 2 when it becomes so complex that it effectively takes up the whole plane (PhiJiips 1985). This dimension is often called the Hausdorff. the three exponents will be equal (Rasband 1990). D. Loehle 1983). which is less than or equal to the capacity dimension. If the slope is a straight line. Then: D = log(_/nl) If the line is smooth. at the extremes. In more formal terms. for example. this kind of rigor in definition has not been necessary in fractai applications to biological problems. That relationship is the fractal dimension. the fractal dimension is the same over a wide range of sampling intervals (Goodchild and Mark 1987). the number of correlations will decrease too. The more crooked the line.g. So far. the higher the fractal dimension. a fractai set is a set with its topological dimension less than its Hausdorff dimension. because the length measure is crucial. Several kinds of fractal dimensions have been identified by mathematicians. the fractal dimension or the HausdorffBesicovitch dimension. The dimensions calculated for these asymptotic regions have been called the inner (microscopic) and outer (upper limit) dimensions. in the realm of geometric measure theory. and D will equal 1 (the Euclidean dimension of a line). but there is a technical difference (Rasband 1990). For the correlation dimension. In fractal theory. that is.

With three or more grid sizes. Morse et al.. but rough surfaces can be secUoned before the fractal dimension is computed.--Ca/cu/ating thefractal dimension (D=-1. Plot grid size against number of squares entered. Box counting is a method used to describe dispersed objects... . A photograph of the object can be placed under grids of three sizes (diagram 9). R.. Canadian Journal of Forest Researctz 16: 124-12 7. • Log X Diagram 8. 1988.. .55 . (Reprinted with permissionfrorrc May. Azlother approach is to make sections parallel to the surface and measure the length of perimeters of "island" and "lakes" (Cahn 1989). How many species are there on Earth? Science..camera and the perimeter length is counted by computer program (Goodchild and Mark 1987). In two dimensions../ / / magnificat /'.--:-_ V/. The logarithm of the number of squares touched by the figure is plotted against the logarithm of the number of squares on one side of the grid. To obtain the fractal dimension of a rough surface. Frontier {1987) describes a method of using circles to compute the fractal dimension of dispersed objects (diagram 10)._. thereby providing data for computing the fractal dimension (Schroeder 1991). __ 10 100 1.21)= 1.000.. D=1..--Usbzg dimension of a plant. °6. A rough surface will have a fractal dimension greater than 2 (Goodchfld and Mark 1987).. J... ofside of grid one squares on a grid to compute the fractal Diagram 9.) For more complicated objects like whole plants.. count the number of squares that cover the object (A).5 = e 6.21) of a silver maple leaf from the slope of the log pixel length versus the log number of lengths.) using line segments. m "O 10. _\_ '-__r _ .. A r.000. (low rr{agnification) 1 1 . {1985) explain this technique in more detail. Fractal analysis of leaf shapes. Cheung.. 241:1441-1449.98 0.0 100._t I __<-_L D='!-(r=0. The slope of the plotted data gives the fractal dimension (diagram 8).. Vlcek and Cheung (1986) calculated the fractal dimension of six leaves from eight trees representing seven species..21 7. Read the slope (13).. E. Fractal dimensions of areas and surfaces can be (5 z 10..8 z. make serial sections perpendicular to the surface and measure the length of the secUon boundary 10 .. (Reprinted with permission fTom: Vlcek.oo0 No.000- /(high ion) :3 _ _ O 100. and the number of penetrated boxes is counted. The box counting is repeated for different scales. obtained using pixels or line segments. 1986.. a grid is placed on the map of dispersed objects. using a grid system may be more efficient than moving different-sized measurement units around the outline. Surfaces may be complicated.. The slope of the line is the fractal dimension (May 1988). Square pixels of different sizes will measure the area of an irregular patch.0 2 3 4 1.

and no criteria have been established for choosing scale (Loehle 1990). Algorithms are provided for computing the fractal dimension of the resulting 3-dimensional surface. fractal dimension 11 . Developments in numerical ecology. wavelet analysis separates position and scale as independent variables and allows identification of self-similar properties of fractal objects (Argoul et aL 1989). a circle centered on this point is drawn to represent the area in which the animal searches for prey during a short time interval. The slope is read from the loglog plot. The density of points inside the spheres is n/V a rd3. For each point where an animal is observed. (Reprinted with permission frorm Frontier. If d is the fractal dimension. The Fourier transform has been used for situations of irregularity and fragmentation. S.. Wavelet analysis is a mathematical technique used for interpreting seismic and accoustic data.Diagram l O. P. the height of each grid square equals the number of circles that overlap the square. Applications of fractal theory to ecology. Next. Some measures are summarized for computing in table 1. Legendre. r. In contrast to the Fourier transform. 1987. L. In: Legendre.. New York.) Loehle (1990) uses a box-counting technique to describe anknal home range. Wavelet analysis Fourier analysis is a new technique replacing for complex time series data.--Measuring the fractal dimension of the patchy distribution of biomass through space with spheres of varying radii. but it is scale dependent. NY: SpringerVertag: 335-378. the number of points intercepted by the spheres is n a r. a grid is laid over the map. eds.

some phenomena do depend on events at many scales (Wilson 1979). scaling is a primary focus. foresters and meteorologists look at vast areas of vegetation. 12 Once spatial and temporal scales have been identified. Now turbulence is modeled as a multifractal.and [Nakamura 1989). Over centuries. many scales were a barrier for physicists who wished to understand and describe turbulence. FR/kCTALS AND SCP. For mathematicians and physicists. An example from the forest illustrates how scale separates research efforts. Temporal and spatial vague when ecologists patterns of vegetational how large an area and needed for succession? areas may or may not small-scale disturbances scales are sometimes and foresters discuss change. Over years. there are seasonal processes. . Algae Home range Crystals Population size the fractai dimension Citation Mandelbrot 1967 Vlcek and Cheung 1986 Morse et aL 1985 Goodchild and Mark 1987 Frontier 1987 Loehle 199. Statistical models were insufficient because they did not give topological information about tile location of the intermittent structures in space (Argoul et aI. The information is not interchangeable (Jarvis and McNaughton 1986). a process is repeated from scale to scale. Ecologists. Sealing in Natuxe Before the fractal approach. Over days. the reality of the scale is hidden by the process itself because without a reference the scale is unknown. Although some events differing greatly in size do not influence each other. 1991).and small-scale be affected by large. Hoekstra eta/. ecologists work with individual plants. are likely to choose scales appropriate to person-centered perception (Wiens 1989. Where are the standard functions that define the appropriate units for such spacetime comparisons in ecology? Continuous linear scales may not be appropriate for discontinuous processes _Viens 1989). Observation scale is the gap in the dividers by which we measure distance and/or time. For example. In a fractal. In building a fractal object. there are rapid fluctuations due to winds and arthropods (Nakamura 1989). herbs and trees can hardly be studied together because of differences in scale (Hoekstra et al. The structure of the world has scales of many different sizes. like transpiration for example. reflecting a complex cascading structure. Conflict of interpretation arises from conflict of scale.mMeasurements for computing Example Coastline Leaf Plant Landscape . Because of scale differences. how long a time are karge.0 Fowler et aL 1989 Argoui et aL 1989 Object/Process Crooked line Outline Outline Surface Dispersed material Movement Solids Time series Measurements Unit lengths Automated unit length (pixel lengths) Grid sizes Pixel areas Circles Box Box Wavelet 4. all scales are equivalent and undiscernible from the form itself (Frontier 1987). soft-litter dynamics change in space and time.Table 1. Plant physiologists work with individual leaves. these scientists may disagree about mechanisms of function.I. following a "cascade. The gap is normally the prerogative of the investigator. 1991). and sphere size in measuring dispersed biomass.I_ Fractal geometry is the first mathematical theory that explicitly uses the concept of observation scale. For example. what is the relation between the two? Both are required in models designed for predictability." However. Results of one group do not apply to the problems studied by another group. 1989). who deal with intuitively familiar phenomena. there is slow accumulation with change due to succession. Combining time and space scales is a challenge for many scientists. like pixel length in measuring leaf perimeters. Even on one small site.

Hierarchy theory emphasizes linkages among scales. Allvariables and parameters are written down and then separated into dimensionless groups. Predictions of spatially broad-scale systems cannot be made with temporally small-scale samples because the time-scale grain is too Free to catch the major movements in the dynamics of the system. similitude defines scale-invariant relationshipsfor water in porous media. For example. but these kinds of classical similitude scaling techniques assume an underlying uniformity of the processor structure. interconversion of units is the goal. 1985). Power laws lack natural scale. f(x) is still proportional to xa. a uniformity that is usually missing (West and Goldberger 1987). f(x) = cx a where c and a are constants. along with the attached boundary conditions of a problem.ions among species. They are called "scale free" or "true on all scales. In allometric relationshipsthere is usually a problem deciding which of two variables is the dependent and which is the independent variable (Zhirmunsky and Kuzmin 1988). Arbitrary scales (e. Allometry is a power law specifically relatedto biological growth. a is an empirical parameter. in soils. Although some correlation exists between scale and level of organization {May 1989). Several scaling techniques have been developed that should be compared and contrasted with fractal methods (see box). the solution can be applied to many other systems that may be physically different. The technique of similitude analysis aims for equations with the least number of variablesby combining variables and making them dimensionless (Hillel and Elrick 1990). Similitude uses this length scale to produce scaled transport coefficients and water potentials in porous media (Sposito and Jury 1990). When an organism develops in stages." or "happening on all scales" (Schroeder 1991). and renormalization are scientifictechniques developed to deal with scale. That is. quadrat size) may limit the detection of interaction (Wiens 1989). A characteristic length scale emerges that reflects the sizes of solid particles and the sizes of pores in a particular geometric arrangement. Physiologists have long expressed metabolic rate as a function of body size with the allometric power law 13 . Some allometric relationshipslink organism size with other parameters. The Fibonacci series has been used for scaling in the lung. Scaling is a common problem in science. but are "scale models" of each other (Miller 1980). where large-scale polities flow from short-term studies (Wiens 1989). In dimensionalanalysis. Scaling issues may be particularly important in the study of interact. They resemble fractal analysis in some ways and differ from it in others. Nor can similitude analysis be used for systems in which some of the variables are unknown (Snyder 1990). W is body weight. Scales may contract or expand from one hierarct_cal level to another (Zhinnunsky and Kuzmin 1988). This is a severe problem in resource management. An infinity of possible physicalsystems is "collapsed" into one reduced solution that describes them all (Miller 1980). Butfor fractals the variables are size and number of units of measurement (Zeide and Pfeifer 1991). Power laws are self-similar. there are no apriori meanmgful levels of scale (Cherbit 1990). Power lawsand aliometryare techniques for describing the relationshipbetween variables as the scale changes. Dimensional analysisis different from similitude analysis because it directly reduces the appropriate differential equations themselves. there may be scale changes at specific points (Zhirmunsky and Kuzmin 1988). AIIometricdevelopment is a relationship between any two properties of a process in which the relationshipis defined by a power function. Fractal analysis may resemble allometry because both approaches use power functions.g. Ifx is re-scaled (multiplied by a constant). dimensional analysis.. We need standardized ways to define and detect scales.Growth rates may increase and decrease over time. ratios between parameters are substituted for actual values (including the units) of those parameters (Gunther and Morgado 1987). allometry. After a solution is found to a reduced formula. but the two are different. like metabolic rates (Sernetz et al. and b is an exponent (Gunther and Morgado 1987). Other Scaling Techniques and Their Relation to Fractais Similitude. In Huxley's original equation Y = a VIP Y is any biological function. AIIometry'spower function relates linear rates of growth between body parts.

resulting in different forms. (If the fractal dimension is 1. and b = 3/4. a tenfold decrease in the size of the insect is like a threefold increase in the apparently available vegetation. "indexes the scale-dependency of the pattern. RenormaF izability may be a fundamental property of nature. Domains are separated by sharp transitions. Variability increases with the approach of transition as has been shown for increasing quadrat size. But there is disagreement about the value of b (Calder 1987).3 to 1. For fractals. The goal is to define the algorithms that will translate across scales (Frontier 1987. similar to phase transitions in physical systems. each on a single length of scale. but the pattern remains self-similar.the system is divided into many equal sections. and this feature of fractals makes them useful in dealing with problems of scale in research. M the body mass. Attempts to modelthe -3/2 rule geometrically have had to assume that plants maintain the same average shape." The fractal dimension is constant ff the processes at fine scale are responsible for the processes at larger scale. Therefore. and that one average replaces all the other variables in the section. the original spacing is restored by reducing back to the dimensions of the original system (Wilson 1979). A change in fractal dimension signals a change in process.with a scale of roughlythree times the original spacing. Renormalization divides a problem with multiple scales into smaller problems. Renormalization is a technique for preserving information while changing scale. For example. and large-scale currents.5. and point samples (Wiens 1989). At the same time. like the principle of special relativity is a fundamental property of nature (Pagels 1985). pattern may differ in detail at different scales. Organisms adapt to these different scales. can be seen more clearly. The fractal dimension.8. Fractal geometry may be useful for identifying domains of scale. Renormalizationblurs out the smaller features of the system but preserves the larger ones. D. the spacing in the new system is larger than the old system--the variables are farther apart. or fractal dimensions (Frontier 1987). even-aged stand will have a slope of -3/2 on a double-logarithmic graph. Domains of scale may be regions of scale change over which the pattern for the phenomenon of interest does not change. Three steps are repeated many times. "part of a large scale element is equal to the whole of the small scale element" (Nakamura 1989). they will look almost identical. But average plant shape is not independentof mass across the plant kingdom. behaviors. The resulting system reflects only the long-range properties of the original system. local currents. turbulence is bounded between the scale of the planet and the scale of the molecule. it is self-similar. First. continuous linear transects. lapping. but those slightly larger. Comparisons across the plant kingdom (shrubs vs. that is. Fractal dimensions have been calculated for various vegetational habitats.5. For insects that eat the edges of leaves (a one-dimensional approach to the vegetation). Renormalization recognizes self-similarity at scalechanges and thereby ties directly into fractal theory. For the third step. according to the self-thinning "rule. that is. the pattern changes or at least changes linearly with change in scale. with all finer scale fluctuations eliminated (Wilson 1979). regions of fractal self-similarity may represent scaling domains. with an average of about 1. But in between are viscosity. Then the variable is averaged for each section. the fluctuations at the smallest scale have been eliminated. for example) are hampered by scale problems (Weller 1989). it provides information about the behavior of distinct but related systems in which the fundamental scale of length gets larger with each iteration. Renormalization functions like a microscopethat zooms in on self-similar structure (Stewart 1989b). and they range from 1. To use a growth model from forestry. then a part will look like the whole. After this. herbs. Scale disappears. Fractal dimensions of the physical environment may also change by scale. 14 Scale-free Fractals In a fractal situation. if something is renormalizable. waves." growth and mortality over time in a crowded. grow isometrically. Wiens 1989).P = aM b with P the rate of energy transformation. How fractals bridge (quantify) the scale differences between small scale and large scale can be seen in the following example of small insects on large vegetation. ifa process or object is self-similar. After the first transformation. a tenfold reduction in the measurement scale increases the perceived . Magnifications of smaller and smaller parts will stabilize. Outside the domain.

a simple harmonic oscillator has a spectrum of one frequency. Foliage tissue is organized as a sponge. The richness of structure shows the "underlying fractal networks" in physiology {West and Shlesinger 1990). and winding from Richardson's study of turbulence through Kolmogorov's study of mechanics. less frequent cycle (Pimm and Redfearn 1988). but is only self-similar over some range of scale. rather than cycles per second. The lung branches 24 times. each of which has the same structure as the whole. 15 The Parts The mathematical fractals introduced in section 2 are self-identicalwthe figures remain the same as the scale changes. such as in electrocardiograms. The number of branching steps optimizes the transfer of matter and energy (Frontier 1987). Self-alTme means selfsimilar. it. They are called 1/fphenomena {West and Shlesinger 1990). with limits. The multiplicity of scales can sometimes be decoded by time series analyses. Self-similarity means that a part of a line. can represent. self-similarity has come into its own with the investigation of fractals. A real tree branches for eight binary steps. And at the other end of the scale. but the parts resemble the Fractals allow us to quantify process change over scales (De Cola 1991). a fractal. FRAC'I'AI_ AND SNI. when combined. but on a smaller scale. having no characteristic scale. the effect is squared: a tenfold decrease in the insect's size is like a tenfold increase in apparent habitat (May 1988). another fractal (Frontier 1987). the average of population sizes over a few years (an "equilibrium") may really be a stage of cycling over a long time period. and gill branches 4 times. In population biology. for most efficient air and sap contact. for example. Beyond that. and a random time series has a broad spectrum. For comparison.length by 10 °. where fis a frequency {West and Shlesinger 1990). Fractals in nature they are self-similar: whole. If insects eat surfaces instead of edges (a two-dimensional approach). but with more than one scaling factor (Schroeder 1991). an individual tree belongs to a forest. Its frequencies make up an inverse power law spectrum. It cannot grow indefinitely or sap could not be supplied to all the leaves. on the idea that an underlying process is continuous. It cannot branch indefinitely because air circulation is required. Many natural phenomena exhibit this spectrum and thereby this scale invariance. statistically speaking. the inverse power law spectrum shows the fractal or scaleinvariant nature of the underlying process. "A set is self-similar if there exists some finite system of transformations such that the images of the set under these transformations. But a fractal process or fractal time series is very different. The various time scales or frequencies of a process can be arrayed in a spectrum." The work of D'Arcy Thompson was based on the scaling concept of similitude.le-S_TY and the Whole Although self-similarity is an old idea dating from Leibniz' study of the line in the early 1700's. It is lack of scale that leads to self-similarity (Schroeder 1991). In other words. It also means that two lines with the same fractal dimension will be like each other and perhaps have the same generating mechanism (Vlcek and Cheung 1986). the set can be split into a finite number of pieces. the whole line. yield the original set. A real biological object is not infinitely selfsimilar. are not self-identical. the twigs bear leaves with another fractal structure. Frequencies in space correspond to numbers of cycles per meter. For both space and time frequencies. But the concept of self-similarity does not require underlying continuity (West 1987). The whole of the figure can be reproduced by magnifying some part of . That cycle may be moving on an even larger. A tree is self-similar. Controversies about whether ecological systems are equilibrium or non-equilibrium systems probably can be resolved by the appropriate choice of time scale and the application of fractal techniques (Wiens 1989). 1/f. Stewart (1989a) described self-similarity more formally in the language of set theory.s = 3). 5.

This may reflect a basic principle of phase transitions. Diffusion through a solid is an example of local changes making global results. High D values indicate short-range factors like soil chemistry (Phillips 1985). a sufficient contact surface) is achieved after a few generative steps." Landforms have a range of D = I.75. If the structure's function (for example. and fractal geometry is a way to approach complex systems 6Vest and Goldberger 1987). indicating the influence of large-scale factors like climate and geology. Fractals provide a measure of short-range vs. a physical impossibility (Tyler and Wheatcraft 1990b). Long time periods of little activity were suddenly punctuated by remarkable events. the smaller sample will have a smaller correlation (Schroeder 1991). Biological systems may require the complex zones may boundaries provided by fractals. they are likely to be "jagged on every scale" (May 1989). may be addressed using notions of fractal structure (De Cola 1991). Low D means "domination by long-range controls. Dimension is a measure of cornplexity (Mendes-France 1990). Renormalization theory in physics is closely related to self-similarity. I to 1. then adjacent samples are correlated and self-similarity is missing. all involve formation or destruction of long-range order (Peitgen and Richter 1986). The accumulation of numbers of points at the diffusion front obeyed a power law characteristic of fractional Brownian motion (Rosso et aL 1990). By zooming in on the path. The length of the actual path depends on the unit of measurement. The number of points was sampled at various times. If the probabilities are not equal. tracing a fractal line. the dynamics of the diffusion front revealed macroscopic results (undiffused areas become diffused) from microscopic changes (single particle movements) over very short time periods. and complexity characterize many biological systems. FRACTAI_ Jaggedness AND COMPLEXITY and Boundary Because a fractal structure exhibits some degree of self-shnilaritymthe structure is the same at several or many scales--fractal structure has bearing on the relationship between the local scale and the global scale. Contact between interacting parts of an ecosystem be enhanced by fractal geometry. When Rosso et aL (1990) simulated the diffusion of a substance through a solid. Energy. solid to liquid or from magnetic . Local vs. Systems undergoing self-similar. relating what is happening here to what is happening there. Fluctuating phenomena in nature are often statistically self-similar structures. mean fluctuations of numbers of points stabilized at the frequency characteristic for fractal phenomena. Global states. The physics of phase transitions is like the mathematics of Julia sets. then selfsimilarity is curtailed. trees and lungs have structure with limited steps (Frontier 1987). Strike out every other event. the "diffusion front" had a fractal dimension of 1. and the statistics of the first set will look like the statistics of the second set. When studied in detail. whether from to non-magnetic Many distribution patterns in time and space are not Gaussian. This f'mite length has to exist or the particle would travel an infinite length. Another example of the physical limitations of self-similarity is a particle moving through soil. The major problem of irregularity in spatial data. there is equal probability of two events happening. which are or may be fractal structures (Peitgen and Richter 1986). Complex fractals produced by the repetition of a simple geometric transformation can turn out completely different by some slight change in the formula (Briggs and Peat 1989). with well-dei'med means and variances. Rather. Phase 16 phase transition are highly transitions. 1/f (Rosso et aL 1990). heterogeneity. When the sampling time was greater than a2/4D (a is the inter-site distance and D is the coefficient of diffusion). Discontinuity. 6.3. but there is no value of D that marks the point on the continuum between complex and simple systems (Phillips 1985). With statistical self-similarity. long-range variation. Where adjacent events are correlated.Biological fractals are also truncated because morphogenesis is expensive in energy and information. a f'mite length becomes visible.

The events leading to the formation of a coastline are impossible to reconstruct. energy. The geometry of plankton patches is a function of water turbulence (Frontier 1987). the fractal root system forms the contact between the tree and nutrients in the soft. Intersects with the linear sample." Chance enters. The fractal object. Frontier (1987) suggests sampling plankton along a ship trajectory. Because the intersection of two fractal objects is a fractal. the landscape is the appropriate spatial unit. spatial pattern is the driving force determining system function. interfaces that are dimensionally constrained. not only on the microscopic level. Energy flow couples with transport mechanisms between points of different energy levels. like in the Brownian molecular motion of molecules. the dimension of the intersection is equal to the sum of the dimensions of the two original fractals. and information may flow more easily from one segment to another (Frontier 1987). then D = d + 2. Noise Randonmess. The littoral zone brings together the primary producers and the decomposers. complex ecosystem boundary. and information through interfaces. for example. A fractal wall configuration and transport system accelerates the energy flow and the cycling of matter through the system. minus the dimension of the space in which the two fractals interact.2. Noise is the uncertainty In complex systems. accelerating the cycIing of matter. The same rule applies to the intersection of a fractal with a non-fractal object. is a component of fractal formation In nature. 5. Patregardand Garlic Biomass is often distributed in hierarchical patches. Here is where the physical and biological Interact. noise. but it is actually shoreline fractal dimension that should be correlated with other properties. If D and d are the fractal dimensions of. less of the system or scale (Ben-Jacob 1990). the fractal canopy forms the contact between the atmosphere and the chlorophyll web. Independently of the underlying 17 How does pattern form from a disordered environment? Is pattern formation In nature the result of unique causes and effects governing each phenomenon. respectively. Interfaces that are fractals. This contact zone between two ecosystems increases as the fractal dimension of the shoreline increases (Frontier 1987). Lake shape has been described in terms of a ratio between shoreline length and water volume. An example of the self-organization complex patterns are systems with a diffusion of . the plankton in three dimensions {the swarm). Pond shape and lake shape are related to biological properties like productivity. For trees. The lungs and g_s may be other examples of fractal structures that exchange energy and matter between the organism and its environment {Frontier 1987). Noise contributes to the complexity of fractals. Pattern is studied at large scale to relate it to ecological processes studied at smaller scale. the swarm and the'linear sample. where population Interacts with population and with the abiotic to form pattern. Geomorphological processes happen "through many ill-explored Intermediates. For large-scale spatial heterogeneity. d = D + 1 . the science of interaction between populations and environment (Frontier 1987). Mandelbrot (1983) highlighted this in his coastline example. Some unifying principles are emerging from growth patterns. Parameters have to be developed that defme spatial pattern in a way that hooks up with the underlying biophysical process (Krummel 1986). but also on the macroscopic level (Mandelbrot 1983). For landscape ecology. Fractal organization is especially visible in ecology. Life may be det'med by interaction and the flow of matter.3 = D . or do unifying principles underly the formation of pattern (Ben-dacob and Garik 1990)? Pattern may form from the Interaction of microdynamics with macrodynamics. The probability distribution function describes complexity. Complex Pattern and Self-org_!zatlon field like the one discussed in section terns group into typical morphologies. Webs of organisms connect through exchange of matter and energy. and patterns like these can be studied using the fractal approach. The littoral zone is an important example of a fractal.matter.

. until it reaches distributed or lognorincrease in complex1/f systems (diathe curve of lognoras complexity a 1/f distribution. The 1/f and the lognormal distributions will overlap more and more as complexity increases. species distribution) or general biology {e. 78: 40-45. Several researchers in forestry-related disciplines have used the concept of fractional dimension (table 2). TO FORF_T SYSTEMS 7. M. and will be discussed in the sections that follow. and melody in music (Voss and Clarke 1975). B. Fractal dimensions have been computed for a diverse array of things.mechanisms. As simple systems with little variability become more complex. taxonomy). The power spectrum is related autocorrelation function. it is not surprising that initial appLications to problems in forestry should emphasize shape and size. Besides the fractal dimension. (Reprinted with permissionfrorru" West.5) over many degrees of frequency." with the form f4 (0. The area under mal distributions increases increases. straight line with negative slope. More complex leaves had higher D and greater standard deviation among leaves. from tree rings to taxonomic lists. There is a sloped time scales (Voss and Clarke The spectrum called fractal to the 1/fpower specby a single correrelationship over 1975). Shlesinger.) 18 traditionally linear and/or areal measures.g.:c_ General ariable(logscale) v Leaf shapes Diagram 11 . APPLICATIONS As with most young fields.g. the overlap of the lognormal and 1/f distributions will be large and the distributions will be indistinguishable. However. Vlcek and Cheung (1986) calculated the fractal dimensions of the perimeters of leaves (diagram 12). 1990.. The trum cannot be characterized lation time. the applications of fractal geometry to the biological sciences have been primarily descriptive.. the techniques and approaches may be useful to the forest scientist. When many mechanisms are working at many scales and the main process is accomplished by many subprocesses. Nevertheless. They belong more formally to the realms of ecology (e. but these measures do not represent natural shapes very well. and of fractals and self-similarity. Noise in natural phenomena. many of the studies do not deal strictly with forestry. Leaf Shape have been described by -e" > e- ® _ "6 C o r_ "5 .. These include loudness fluctuations in music and speech. The behavior of 1/f phenomena is due to complexity rather than content (West and Shlesinger 1989). a physical variables are "1 f-like. Given fractal geometry's predilection for re-analysis of shape and size. there has been some use of fractals and scale. As normally really distributed systems ity. they come to resemble gram 11). whose aim is to describe and understand the forest's complexity. The nine species they measured were significantly different. American Scientist.5 < 1 < 1. The power spectra of many of noise distributed as 1/fis noise because it is scale-free. the area under the curve increases until the distribution is described by a straight line with negative slope: a i/fsystem.uVariabiIity of lognorrreal and I/f systems. This is why the inverse power law can be so universal (West and Shlesinger 1990).

1-2. Tree Branching (Crawford and Young Data from two species of oak were analyzed with two different models: a Markov model where branch length depends on the length in the previous order. Power law and exponential distributions were indistinguishable for small values of the exponent.0 1.05 1.44 1.04 1.02-1. 1.19 1.8 1. So.28 2. below 60 ha Deciduous patch.Table 2.22-1.4 1. two different species that differ in the size and number of lobes of their leaves can have identical fractal dimensions. But at higher values.1-1.1 Mandelbrot 1983 Mandelbrot 1983 Krummel 1986 Krumme11986 Morse et aL 1985.36 1.45 1.11.13-1.75 1.21 1. In the fractal model.19.1-2.3-1. dimensions for their leaf perimeters. unpublished Lorimer. 1.52 1.18. of segments within the 1990).0 1. Crawford 19 Crawford and Young (1990) studied tree branching by characterizing and then classifylng the kinds of branching in actual trees.5-2. but branching measured in the lower and upper crowns of one species was the same.3 2.81 1. May 1988 Lorimer. and each was assigned an integer label. were grouped in a hierarchy of orders. the function giving the relative frequency of each scale in the structure is a fractal.64 1. The opposite is also possible. had different fractal between diameter order order number and number and average length. the same tree species can have leaves with variable fractal dimensions.6 1.27 2. and a fractal model. the branching followed the fractal model. the other without one with notched leaves and notches. Although fractal geometry can be used to study leaf shape within species.22 Reference Burrough 1985 Taylor 1988 Crawford and Young 1990 Vlcek and Cheung 1986 Vlcek and Cheung 1986 Vlcek Vlcek Vlcek Vlcek Vlcek Vlcek VIcek Zeide Zeide and Cheung and Cheung and Cheung and Cheung and Cheung and Cheung and Cheung 1990 1990 1986 1986 1986 1986 1986 1986 1986 Sugar maple leaf Silver maple leaf Gingko leaf English elm leaf American elm leaf American basswood leaf White oak leaf Ponderosa pine crowns Hemlock crowns Cypress patch (Florida) 8roadleaf patch (Florida) Deciduous patch.55 1. unpublished Burrough 1985 Phillips 1985 Burrough 1985 Burrough 1985 Burrough 1985 Burlando 1990 Two ginkgo trees. 1.77 0.05 1.07-1. _The fractal dimension of the distribution of scales will have a bearing on the overall appearance of the tree (Crawford and Young 1990).--Fractal Feature measured Tree ring indices Tree ring widths Oak branching Red oak leaf White oak leaf dimensions from the forest Dimension 1. section Landforms River flows Annual precipitation Taxonomy 1. As in the case of the oaks shown in the diagram. The authors then noted relationships .46-1. the points where branches emerge.18 1. Segments between nodes.15 1. Wyoming Soil properties Landforms. Because branching in the two species of oaks was different. above 60 ha Vegetational habitats Fire scars.47-1.02 1. Minnesota Fire scars. it may not be useful for differentiatlng species (Vlcek and Cheung 1986).

45. 16: 124-12 7. Fractal analysis of leaf shapes. (Reprinted with permissionfrorrr" Vlcek. Zeide (199 l a) cornputed fractal dimensions of crowns on three different plantations. In a study of loblolly pine. A. 1986. Zeide (1990) developed the two-surface method. foresters did not have the tools to measure complex objects like tree (Loehle difference 1983).63. J. all the needles appearing on the crown surface (Zeide 1990). Cheung.. The internal branching angle (identical for all angles within a tree) of at Another forest characteristic. The leaves with the most complex shapes have the largest fractat dimensions. and other properties. A dimension of two indicates trees very intolerant of shading. there will be a 2O . and 2.74.mFractal dimensions of several tree leaves. Between the volumes of the largest and smallest crowns. Tree Crowns The regression of the logarithm of leaf mass (assumed to be proportional to leaf area) plotted on the logarithm of the cubic root of crown volume (representing the linear size of the crown) for 10 conifer species in the Rocky Mountains indicates self-similarity for crown surface. can be modeled as a fractal packing problem. it Is difficult to adequately d_ e_ _ merge. E. The fractal dimensions. Tree Models Mandelbrot (1983) modeled "trees" of different fractional dimensions and noted the effect on form (diagram 13). The fractal dimension may be sensitive to thinning (Zeide 199 la). This relationship is A (a is a constant) (Zeide 1990). There are practical problems in calculating the fractal dimension of tree crowns.81 for very tolerant hemlocks. arrangement of tree crowns. In the forest. and Young conclude that branching pattern is genetically rather than environmentally determined.27 for intolerant ponderosa pines to 2. the branching structure showed large intraorder variability because of contributions many scales (Crawford and Young 1990). with mass proportional to volume (meaning needles tolerant to low light). packing of smaller and smaller sizes stops at the scale of the crown size of the smallest plant existing at a given height above the ground. productivity. The slope ranged from 2. increased with site quality. D.b_ _A Before fractal geometry. then fill in the remaining spaces with the next size and so forth. The usual method of box-counting is difficult to apply because three-dimensional o eecroo O = aE D/2 1_ _ _ i Diagram 12. 2. A value of three indicates uniform needle distribution.two or three orders measures of of magnitudeof surfaces. tree surface area. Yet without adequate estimate tree respiration per unit area. E. 2.) obtain. Fill a volume of space with as many large irregular objects as possible. tree surface measurement depends on the unit of measurement (Zeide 1990). has a parameter identispatial data on tree crowns are difficult to fied as a fractal d_nension of crown surface. Canadian Journal of Forest Research. and surface area of a convex hull that encloses the crown. Like all fractal structures. however. The fractal model reproduced the power law behavior of mean branch length as a function order. The relationship between leaf area of a tree.

Freeman and Co." Varying the branch angle over a wide range while keeping branch length and D the same results in quite a variety of tree shapes (Mandelbrot 1983). a fractional dimension. What of thick-stemmed tree forms In nature? Real trees illustrate the idea of a shape having many Besides the D of the branch tips. the fractal nature is more apparent. When D and A are not Integers. the dimension of the tips is D. 1983. the trees in the diagram are not strictly self-similar. D increases from slightly above 1 (top left) to 2 (bottom left). but there may be architectural and physiological constraints. Each branch is still dimension 1. From a fractal poInt of view. The fractal geometry of nature. and that it is equal From a topological (that is. and d2. with a branching angle greater than 180 degrees. With D greater than the Euclidean dimension.H. The internal branching angle changes as thefractal dimension changes. A. allowed to overlap. Mandelbrot extended fractal cano- Trees are both branch tips and branches.emerged from the restriction that no branches overlap. satisfy the relation For botanical Diagram 13. Without thin trunks. The branches dominate. whose dimensions "clash." than 1983). a tree's branch diameters before and after bifurcations d. even though the branches seem to dominate In an intuitive and topological sense. To put this formally. When tree branches are set of tips is no longer a dimension 0. the dust of Euclidean of dimension 1 (with dimension). __ Da Vinci claimed that "All the branches of a tree at every stage of its height when put together are equal in thickness trunk (below Otherwise A is less to the D (Mandelbrot them). A corollary to D = 3 and A = 2 is that the branch's leaf area is proportional both to the volume of the branch's outline and to the crosssectional area of the branch. Euclidean) point of view. dl. but the exceptions to this may be more Interesting than the rule. another parameter. (Reprinted with permissionfrorru MandeIbrot. but a curve no change in the fractal (1983) calls these curves pies. A have D = A. trees have exponent. dA = dl a + d2 A with the exponent 1983). A second corollary is that the ratio (tree height) 3 (trunk diameter) 2 is constant for each to the ratio species. but the whole tree dimension is D. the diameter self-similar tree with a trunk will linear scales.) trees. That is. NY: W. 21 . D measures the abundance of branching (Mandelbrot 1983). they are fractally negligible. B. New York. has a D less than 2 (Mandelbrot 1983).reModels of trees with non-overlapping branches. The tree on the lower right. D equal to 3 would give the most surface to the sun. The data for the measurement of A are sparse (Mandelbrot 1983). the branch tips would be dimension 0 (points) and the branches would be dimension 1 (lines). being A = 2 (Mandelbrot ___ D = 3 and A = 2.

vegetation is a candidate for fractal analysis (Palmer 1988). Branch shedding gives long sections of clear bole (Loehle 1983). Parameters for curves and twists of branches and the ratio of the size of the trunk to the branches are added. Before the fractal concept. species patches are irregular. trunks and large branches on hardwoods do not fit the log-log relationship. Conserving the cross-sectional area after each forking episode gives a slope of-2: the figure will till space. small patches had lower fractal dimension than large patches. natural forests have boundaries simple algorithms by adding increments to a cylinder and creating forks at each increment. When the fractal dimension of deciduous forest patches in Mississippi was computed from digitized aerial photographs. the "natural" patterns were more irregular (Krummel 1986). 1991). can now be quantified (Loehle 1983). Vegei_on Palmer (1988) suggests Patches that vegetation Is a This ratio may not vary among (Mandelbrot 1983). having detail at many spatial scales. For real trees. Below 60 ha. Loehle (1983) derived a formula for stem and root surface areas. the slope is between . . Size and Density of Organisms Related to Habitat A very realistic recursive algorithm in which the main tree trunk extends to smaller and smaller branches. the length of branches in a radius class has to be measured. Shorrocks et al. Tree crown models can be constructed with prime example of a fractal. High elevation between meadow are fractal (Loehle 1983).6) were more irregular than the perimeters of broadleaf and mixed broadleaf tree patches (D = I). tree can be generated with a with few measuresampling. However. The corresponding fractal dimension will thus range from 2 to 3. species There are engineering limitations to a tree's height: a fixed percent of the critical buckling height of a uniform cylinder of the same base diameter loaded under its own weight. If the cross-sectional area is not conserved.(linear scale of a branch's (branch's drainage volume) 3 formula in the computer program is like a over diameter) 2 metaphor for increased genetic information evolutionary time (LaBrecque 1987). whether the habitat is soil (Frontier 1987) or foliage (Morse et al. can be known. The perimeters of cypress patches (D = 1. plus the trunk and large branches. an intuitively important but quantitatively vague concept. 1985. including the fractal dimension. The extra information added to the simple recursive 22 Sizes and densities of individuals are related to the fractal dimensions of their habitats. quantification of vegetative pattems had not been possible despite its usefulness for habitat typing. forest patches had regular shapes: the smaller patches were generated by survey and township divisions. to the larger scale of a landscape's vegetation patches. This tree looks !ike it has evolved a long way from Mandelbrot's trees. Patchiness. Forest biomass estimates can then be made ments and without destructive bark thickness as a function. Size as well as species can influence the fractal dimension of a patch perimeter. to the yet larger scale of vegetation patterns influenced by geomorphological features. large vegetation patches are joined by satellite patches of different sizes. tree utilization similar to boundaries (Mandelbrot forest and islands 1983). This requirement yields the same results as fractals with D = 3 and A = 2 (Mandelbrot 1983). volumes. From the small scale of individual plants.1 and -2: the figure will have a finite volume. LANDSAT analysis. Once basic values have been obtained. Above 60 ha. With of branch radius. Plotting log frequency of branches of a given radius vs. Mandelbrot (1983) cites an unpublished report on patch irregularity in the Okefenokee Swamp. and other applications. log of that radius gives the straight line with negative slope that is characteristic of fractals. and numbers of tips. In the forest. the amount of bark on a tree which is important for whole (Loehle 1983). analysis of root disease centers. From the air. more attributes can be estimated after measuring diameter at the base of the crown and top of the root system.

and smaller insects represent smaller units of measurement.44.5. _--_---_ _ / _ = 10 Diagram 14. then population density scales as (L3)-°-75 (L is body length).. 23 measured for lichen transects ranged from 1.. If a fractal surface has a linear transect with dimension 1. with a mean of 1.. .75 of body weight. Order of magnitude decreases in body length predict increases of between 178 x 3. increasing the available space for small animals (Morse et at 1985). creases the distance from point to point inby a factor of 10 = 3.R. from the fractal dimension of the .. Shorrocks eta/.*"0 _ \. Larger insects represent larger units of measurement.16 . 1985. The body size vs.780-fold in animal number (Morse et al. Reprinted with permission frorm Morris. A 10-fold decrease in body length results in a (103) °-Ts = 178-fold increase in the density of individuals. The fractal dimension of vegetation surfaces. The fractal dimension of a tree. variety. and if population density is proportional to the reciprocal of metabolic rate (this being related to resource utilization). given the numbers and body lengths of collected Insects (Morse et aL 1985).28 to 1. measurement sticks. The fractal dimensions for _. Sweep net data for numbers of insects agree with the predictions (diagram 14). Species Diversity and Possible applications of fractals in community ecology Include analyses of geographical distributions of species and resource partitioning among species. D.560-fold and 178 x 10 = 1. If rectabolic rate scales as 0. I X _ I _l _ .. We may reverse the calculations to obtain fractal dimensions of vegetation. et al. Lawton (1986) also suggested the formulation of a general theory of plant architecture that relates the size.. Wiens and Mikne 1989).Morse et ak (1985) computed fractal dimensions for plant surfaces ranging from 1. If so. Fractal dimension of arthropod body lengths. • _ _. _ " * * . On vegetation with a fractal dimension of 1. The authors further speculate that species-area distributions and the allocation of environmental resources between species are fractal processes. Wflliamson and Lawton (1991) note that when a species distribution is mapped at different scales. Nature. This can be carried one step further. pairs of maps appear selfsimilar. So the relative numbers of animals of different body lengths living on vegetation surfaces can be predicted.79. may be an important "ecological indicator" (Wflliamson and Lawton 1991). Arthropods living on the surfaces of plants are like rulers.: . Vegetation surfaces can change over time as well as from place to place. can change by season (Wflliamson and Lawton 1991). There are 10.. (1991) obtained similar data insects on lichen. the area perceived by a 3-ram animal will be an order of magnitude larger than the same area perceived by an animal 30 mm long..5..._The negative log-log relationship between numbers of arthropods and body length collected by sweep nets from understory foliage in a primary forest in Costa Rica (left) and by pyrethrum knockdown from a forest canopy (Betula pubescens) in Eng/and (r/ght).78. for example. 314: 731-733. which may differ in different locations. with 1. .000° XCX_ '_ 100. 1985).. for every order of magnitude increase in measurement length.37 to 1. insect frequency graphs for collembola and mites collected from the lichen matched the predictions vegetation.. a pattern at one scale can be used to predict patterns at larger or finer scales. X _ _ _ _ _ l " " _ _ / _ X • _ . I • o implications for the animals that live on these plants (Morse et aL 1985.58 the average..16. More data are needed on fractal dimensions of habitats and the body sizes of animals that live In those habitats. fractal nature of plant structures.

a pattern shown by plants and animals at several taxonomic levels. until the smallest grains (monotypic taxa) fill the smallest spaces (Burlando 1990). The smaller grains leave spaces taken by even smaller grains.013 genera had fractal dimension of 1. which had the lowest. b. the number of conditions required for species occurrence). The parameters define the shape of the curve: g is a negatively sloped asymptote. . the probability of species occurrence when the conditions that it requires are present). small insects require fewer resources to survive than large insects. Frontier (1987) proposes ecological interpretations of the parameters: b is linked to the diversity of the environment (i. Frontier (1987) states.66. Large grains packed together leave empty spaces among themselves where smaller grains can fit. For plantfeeding insects. The greater habitat area and smaller resource requirement combine to create a larger habitat carrying capacity for small insects. Animal Movement In a fractal framework. Observations of insect communities support this prediction: the number of small insects increases exponentially with decreasing body size.5. Minelli et at 1991) (diagram 15). and 1/g is linked to the predictability of the community (i.e.. The shape--convex. Another way to show distribution of individuals among species is a rarlk-frequency distribution. Some examples: 70. 777 families of world plants in 145 orders had a fractal dimension of 1. K a constant. Arthropods and higher plants show D estimates near 1. Many taxa have only one or two subtaxa.258 species of world mammals in 1. Further. Because plant surface is a fractal. the argument accounts for less than half of the observed increase in the number of small relative to large species. and b is the direction of approach. where Po. Such distributions can be described with a curve predicting the probability of species occurrence Pr(Sr) as a function of its rank r. Although this argument may explain the large number of small individuals. taxa with different numbers of subtaxa are like different-sized grains of sand. after log transformation. Why should taxonomic diversity be fractal? In an analogy similar to the concept of fractal tiling. Pr(Sr) = Po(r + b)s. The equation for these plots is N = KS -D where N is the frequency of the genera. Finally. Fractal dimensions computed from 44 taxonomic checklists and catalogs ranged from 1. and so on. and g are parameters to be estimated. Rank-frequency distributions may have a fractal origin (Frontier 1987).439 genera had fractal dimension of 1.e.029 species of South American Coleoptera in 6. Similar values occur for the same groups from different geographical areas. Log-log plots of frequencies of genera against numbers of species fit straight lines with negarive slope. 4.46. few taxa have many subtaxa. except for amphibians and reptiles. S the number of species per genus.1 to 2.21 (Burlando 1990). and -D the exponent corresponding to the log-log regression line slope b. Vertebrates had the highest values. the explanation may depend in part on the observation that plant surfaces are fractals. more absolute habitat surface area is available to small insects than to large insects.Lawton (1986) addresses the question of why there are so many more little species than big species in animal communities. then each species is given a point on the graph. that 1/g is the fractal dimension of the distribution of individuals among species. The unascribed increase can be due to increasing resource variety at smaller habitat scales. The value for families within orders was similar to the value for species within genera. with rank on the abscissa and frequency on the ordinate. surface distances depend on the unit of measurement. 24 Fractal taxonomy gives us more information about the organization of species diversity. Species are ranked by frequency. For example. or stepped--gives information about species distribution. a relationship common to fractals (Burlando 1990. The absolute value of D is the fract_ dimension (Burlando 1990). so a fractal approach may be useful in the study of animal mobility (Weiss and Murphy 1988). concave.14. collected by different taxonomists. without proof..

--Fracta/taxonomy: log-log plots of frequencies of genera with numi_rs of species. The graph on the lower right shows the frequencies of orders with numbers of families. . Caterpillars 3 cm long travel 1. to cover a linear meter. HYMENOPTERA NOrth America_ . B.N POLYCHAETES World "'" k COLEOPTERA South America i .5... 1-mm caterpillars travel 2 m to cover a linear meter.42 m... Diagram 16 shows actual distances traveled by caterpillars of different sizes over fractal paths of 1. JA i Js&J &A i l& Diagram 15. 25 . ....) Caterpillar dispersal is a critical mechanism in the population dynamics of many Lepidoptera.. the less the mortality and the faster the development time (Weiss and Murphy 1988}.TRACHEOPHYTES North America" SPONGES Mediterranean MOLLUSCS North America A . Large larvae can move farther. (Reprinted with permission frorrL" Burlando. 1_ . A straight line with a negative slope is common in fractal relationships.1 to 1. | A l t :. FISHES North America BIRDS World ANIMALS A I . Caterpillar paths can be computed fractal dimensions. 146:99-114. Journal of Theoretical Biology.1. The fractal dimensions of taxonomic systems. 1990. distance traveled increases (Weiss and Murphy 1988}. The sooner caterpillars fred food. first-instar larvae have to be close to food. representing only slight roughness. As D increases. Even at D = 1. | A 1 1 I1 1 l 11 1 II. Completely smooth surfaces would have D = 1. with the length for given of the caterpillar as the step-length.

These search patterns are probably fractal. but also hypotheses about causes of pattern. The fire clusters of burned vs.rs} twigs or down a whole forest. damage at various spatial scales: it can break The effect of wind turbulence on breakage in the forest can be studied by quantifying canopy roughness. At the critical point.t ¢ :::_:_$ _ I I t l . and fire create openings in the forest.3 12 _. broad hunting behavior is replaced by local hunting behavior-straight scanning movements become BrownJan-type movements. nonburned trees resulting from model parameters had fractal dimensionality of 1. Lorimer (unpublished) computed the fractal dimension of the perimeters of spatial fire patterns in Minnesota (Heinselman 1973) and Wyoming (Romme 1982). Hence. A caterpillar 1 cm long must travel 251 body lengths to move 1 m along a surface with a fractal dimension of 1.2. 1988. An example is forest disturbance by wind. Loehle (1983) postulated that wind damage may be seKsimilar.75.o3 log CATERPILLAR LENGTH (m. Some areas are used extensively. wind may cause 8 1. 1.' 32 . there was statistical self-similarity. Patterns of wind damage in different forests can be explained in terms of interaction between a fractal. 2: 116-118.D. home ranges are usually represented by bell curves or polygons. The fractal nature of these openings. Forest Applying fractal Disturbance to spatial patterns 24 concepts _. meaning the point of fire propagation. some areas are never used.4 enables not only more precise descriptions. Nor is home range merely a bounded space within which an animal may be found. (Reprinted with permission from: Weiss.__ 0 1. MacKay and Jan (1984) modeled forest fire dynamics with the techniques of percolation theory. small eddies whirl within larger eddies. but they are actually more fragmented and irregular. not much data are available on the fractal nature of these kinds of forest disturbance. trees are either burned or not burned (Schroeder 1991). Instead of up-spin and down-spin. caterpillars of different lengths travel different distances. 26 . In their protocol.ool . Besides wind.) Dispersal distances can be expressed in terms of body length. called the percolation threshold... may contribute to vegetational patchiness. Animal ranges have fractal characteristics (Loehle 1990). Fractal geometry and caterpillar dispersal Functional Ecology.0 __:: :l. agents such as insects..46 to 1. S. correlated with the fractal pattern of the prey. the irregularity of the boundary between damaged and undamaged areas. Diagram 16..t.oos .s ' ' When predators encounter prey.B.64. Murphy. Therefore.16 J. the distance an animal travels depends both upon its size and the texture of the substrate (Weiss and Murphy 1988). a burning tree in a densely packed lattice was able to ignite its nearest neighbors according to an assigned probability. disease. A third example of fractal ramifications of animal movement is Frontier's (1987) discussion of complex and hierarchical predator behavior. self-similar process (wind turbulence) and a fractal surface (the forest canopy) (Loehle 1983).ol . A collection of trees can therefore be modeled like a ferromagnet. For larger animals. The same insect would have to crawl 1.5.--Over rough surfaces.000 body lengths to move 1 m along a surface with a fractal dimension of 1. D ranged from 1. Wind turbulence is self-similar.. However. D.D=1.

....! ! ii_'_i".: :-: :-: :.." -. and 1... 26: 1047-1054.........: :-: :-: :.........46.: :-: :-: :. !ii_iiiii!!i_!!iiii!iliiiiilJ]iH!!i!_ii!! Diagram 17.... (Reprinted with permission frorrr" Tyler. The fractal dimensions of 1..._.......: ......89..... the water retention curve is like the curve for a coarse sand........ :. Before the fractal concept was developed. iiiililiiiiiiiii!iiiiiiliiiiiii!iiiliiliii!iii!iiiiliiiiliiii!iiiiliiiilii! .-. and (d) is 1.._J_z_ ------------._..96.. -:In:.._. Much information is available on particle size distribution. .....'..: :.99... 1..._-.: :-: :-: "... !-!-!-! -!li!-!-! -! !-!-! "-""-"! ! "-'... Near a D of 1. laractal processes in soil water retention. : : :::::::::::: * !!! !!!it! :. the curve is like the curve for a clay soil (T_$1erand Wheatcraft 1990a). Water Resources Research.: :....99 measure the ratio between characteristic pore size and pore area... ltrheatcraft.......:-::-:i:::-::::'L::_..... :.":'£HIH£H££_£_HHHHHH_£H_H££1ilH£_! iiii_iiiiiiiJiiiiiiiIiiiiiiiiiiiiiiJiii !ii!Niiiiif!!Nii!fi!!iNiiii!N!i!iilN :::::!!:!!!!!!!!!!!11!!........... (b) is 1.._ _i!!iiiL......... Near a fractal dimension of 2.. :._i--_-.." . ...-m[] [] [] [] [] [] ::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::: !ij!!!!!!!!!!i!i!!i!i!_i_iil_.... ........" -"-" "-'..iiii .89...!!!_i... the Sierpinstd carpet.... The diagram shows how the fractal dimension affects the shape of the curve for water retention.-..." ..:!'... :..'-" ! ! ! "... but measures of water retention are difficult to obtain. Tyler and Wheatcraft (1990a) model the relationship with a classic fractal form. ::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::: "::-".5.: :.....i'_i_'! .... !'!...._i--_-----.: :.._. The nearer the value is to 1...."-""-" ".. the more the soft is dominated by large pores: the carpet will be sparse at large scales of measurement..Modeling Soils with Siext)ixxski Carpets {diagram 18). .._:::i:-:.... !'!.." -. i!ii!ii!iiiiiiliiiiiiiiiiiiiiiiiiiiiliiii!iiliiiiiliiiilililiiiiliiliiiilii:.. it was difficult to relate particle-size distribution in soils to water retention data." :.._...." "-" ! "-" .::"! "'" : !'!!'!!'!!'!: "'" . the authors calculate water retention data from the models a b [] [] [] [] [] [] _ii [] [] [] [] [] [] [] [] [] [] [] []_ll [] [] [] [] [] [] [] H_'E_[] [] [] [] [] [] IR M [] [] [] [] [] -l[] [] Ill -il. The dimension of carpet (a) is 1......." . _ .--Sferpinski carpets modeling soils of different fractional dimensions..._ _...: :. The fractal dimensions are measures of soil texture..: :........--.46. Using different Sierpinski carpets as models of different soils (diagram 17). :"!-!'!-!'!!'!!'" "'"-:..... 1990o. W... ._t..: .96.... 1. (c) is 1.. -r_.. S. ..._--i---_!'!ii!-!!-! ..W. ...) 27 ...................." .:. -_ .: :.. S.

" 1989 October 18. The spatial organization of soft can change over very short time scales because of weathering. continuous. It can be compared with other soil properties like surface charge and aggregate stability to examine processes of soft creation and destruction (Bartoli et al..---. at that scale the second forest is homogeneous (Palmer 1988). S. " . Soils with lowerfractal dimensions are dominated by large pores.I In this section. Soil A returns water like coarse sand. Fractal processes soil water retnetioru Water Resources Research. Wheatcrafl." Consider two forests of 20 ha each. . i i i l I Fractals may make it possible to quantify soil heterogeneity and to relate structure to specific soil processes (Young and Crawford Spatial 1991). Heterogeneity and Sarnpllng v uJ o.. It can also mean "remaining similar upon subdivision.mRelationship between water retention andfractal dimension in soils. ON 016 -1 0...0 Diagram 18. Wheatcraft. Traditional be applied to this S A_r_#_tg 28 Diagram 19..W. or consistent.. 26: 1047-I054. o-1.. = Ill = O_ _< " _ " '. At low magnification (field scale).. the sample on the left is homogeneous. the species have the same mix in each hectare. But this concept of homogeneity is linked to scale. ^..mHeterogeneity can depend on scale. (Reprinted with permission frorrc Tyler. 1990b..• • \ __ ' '. Scaling in soil physics: p_les and applications. D. . D. Madison. each species occurs only within its own 1-ha unit. ".I _ "¢ • cultivation. 1991). o ".W. In. In the first forest. root growth.W. In the first forest. S. heterogeneity looks the same over a range of scales Wheatcraft 1990b) (diagram scaling techniques cannot kind of heterogeneity.) . Elrick. lo"".. well mixed. depending on the "averaging window" or plot size (diagram 19). In a fractal case. WI: Soil Science Society of America: 109-122.4 i i 1. Spec.) in The fractal geometry of soil structure can be carried further. Dml. Soil B returns water like clay.2 o8 •R_AL SATURm. 1990o. 0 "l 0 . In the second forest. t •• I . Wbsdow I 10" l 0. Las Vegas.L o." Hillel. the sample on the right is heterogeneous. In the second forest. heterogeneity emerges on a scale slightly larger than individual trees. The second forest is heterogeneous. proceedings of a symposium. and soil management practices like (Tyler and 20). S. Most of the examples are from soil science and geological research. o=199 : . (Reprinted with permissionfrorru Tyler. only one species is likely. at the scale of several individual trees.. The systems we study can "look" homogeneous or heterogeneous..89 D-148 SOIL B. individual trees of one species will be neighbors of other species. The forest is homogeneous. NV. At high magnification (laboratory scale). so.. • .98" so. 1 What is homogeneity? It can mean lacking variation. so_Lc. eels. each with 20 species of trees. we will investigate the applicability of fractal techniques to issues regarding the sampling of heterogeneity..W.. . Fractal geometry gives us a new way to view heterogeneity. PubL 25. S.. The consequences of fractal scaling in heterogeneous soils and porous media.

Soft-forming factors (interaction of parent materials. 29 are . and other factors. and within each factor may be many scales of interaction (Burrough 1983a). The REV approach sees softs as building blocks of homogeneous material Above the critical size. short-range sources of variation. for soils. NV... Spec. For sampling. for other environmental variables. As applied to softs. climate. When large-scale effects dominate. not abrupt." This implies that.W.iii:_: For soils..5) are compared to/Ys of other environmental variables (less than 1. (Reprinted with permission from: "IiAler. that is. suggested finding the major present in a landscape before at any scale. At the landscape level. proceedings of a symposium. Madison. and degree of correlation among soil properties and properties of the landscape. increments along the sample series tend to be positively correlated. 25. Large D means the variation is irregular and uncorrelated. This presents a problem for statistical analysis because an important underlying assumption. A changing D over closely related scales would require sample spacing with the smallest value of D.. increments along the sampled series are negatively correlated (Burrough 1983a). relief.. WI: Soil Science Society of America: 109-122. interpolation to may be risky because D large (Burrough 1983a)." 1989 October 18. The consequences offractal scaling in heterogeneous soils and porous media. Zones of distinct dimensions could be connected by transition zones. hydrology. biological action. and improving interpolation (Burrough 1981). Two nearby quadrats will have more similarity than two separated quadrats. Large D values in softs may be due to smallscale variations caused by rock weathering. Wheatcraft. However. will be constant. S. A good soft survey recognizes scale of change. representative area volume (REV) is a sample size below which a sample will show wide variation as a function of its location. Elrick. abruptness of change. 1990b. D. eds. rather than quantitative tools (Webster 1985).. Sampling could then be tailored to a particular scale range. non-sampled sites values are usually Burrough (1983a) scales of variation begirming studies for example. is violated.) All vegetation surveys are subject to spatial dependence. surveyors brought intuition to this job. D. Fktbl. Fractals and geostatistics combine to describe the complexity of the spatial variation of soft. the fractal dimension D is useful for showing how a given sampling design could resolve the variations present with the appropriate scales.--Se/f-s/m//ar heterogeneity.." HilIel.W. D values smaller. biological acitivity) operate over different spatial scales. Diagram 20. That kind of sample spacing would resolve long-range variations without the confusion of unresolved short-range effects (Burrough 1983a). they have levels of variability clustered at particular scales. Scaling in soil physics: principles and applications. improving efficiency. The heterogene_ will look the same regardless of the sampling scale. So.. D values can also range widely. The fractal dimension is a useful indicator of autocorrelation over many scales. Identifying these scales would have great practical value. but other natural phenomena are structured. Some natural phenomena do display statistical selfsimilarity over many spatial scales. Las Vegas. softs appear to be "noisier. When/Ts of soft (greater than 1.5). saving money. 1990a). means important. they are not excluded from the fractal concept because of this. erosion. variation is less erratic (BulTough 1983a). In. Fractal geometry may suggest alternative sampling schemes (Palmer 1988). and that. the variable of interest i _iiiiiii_ above some critical scale (Tyler and Wheatcraft . A large D after the resolution of the survey has been adjusted.S.. that replicates are independent. Small D means that the samples found the change in the variable to be smooth. Previously. D values can be used to sort out scales of variation linked to particular natural processes.

One study (Phillips1986) identified29 variables involved in shore erosion of the Delaware Bay. To determinethe structureof a process. corresponding to a fractal dimension of 2. In block analysis. corresponding to a fractal dimension of 1. For shoreline erosion. variance is plotted as a function of doubling quadrat or block area. variance is plotted against scale. The slope of the double logarithmic plot of a parabola is 2. so D's can be calculated relative to the sampling interval (Burrough 1981). geomorphologistsfind the scales of variation and then identifycontrols operating at those scales (Phillips1986). Block techniques for ecological pattem analysis are related to the semivariogram. or local controls.the important scale of variation is the local scale (100 m) (Phillips 1986). If the variable changes as a linear function of distance. Fractal dimensions can be calculated from these diagrams. meters. they have little influence. shorelinewidth. maximum elevation above mean low water. large-scale influences in a natural system. Shoreline erosion is an example of a complex processoperating at many scales. Semivariograms are better because they plot variance as a continuous function of scale. Many semivariograms are available in the literature. 30 . Geostattstical methods use spatial autocorrelation.Fracta] techniques can be combined with geostatistics to fine-tune sampling in heterogeneous environments. Block analysis has few replicates at large sizes. The fractal dimension can be computedfrom this double-log plot and in this case is 1. as shown by the semivariogram(diagram 21). In a semivariogram.and that if any longrange trends exist. Any long-range trends that may exist are obscured by local effects (Phillips1986). A fractal dimension of nearly2 means that the pattern of erosion is verycomplex and irregular. the slope of the semivariogram will be zero.and phi units (to measure sand grains). the theory of regionalized variables (Meentemeyer and Box 1987). and only few scales are considered. that wide variation occurs over short distances. the semivariogram will be a parabola. The shoreline erosion can be studied with the fractal dimension and the variogram used together (see box). D and Shoreline Erosion An examplefrom geomorphology illustrates how the semivariogram and fractal dimension together can give informationabout natural phenomena. and mean sand grain size.a pattern resulting from short-range. So the fractal dimension indexes the variable's spatial dependence (Palmer 1988). The semivariogram and fractal dimension show that the distribution of erosionrates is highly complex and variable. Shoreline erosion is an example of small-scale influences vs. Controlsover relationshipsin geomorphologyvary with spatial scale. Commercial geologists developed geostatistics to predict the amount of metal ore in unsampled locations (Palmer 1988). The semivariogram summarizes the variance in a dependent variable as a function of scale. If the values for the variable in two near samples are as different as they are in two distant samples.91. Calculation is computer intensive because each quadrat is compared to each other quadrat (Palmer 1988). Wave attack was a predominant variable acting over many scales. The variables representedseveral scales: kilometers. Some of these variables were tidal range. It is a statistically random pattern indicatingnegative correlation between adjacent sites.

. Highly complex enviromnental gradients may be common in nature. An implicit assumption is continuous spatial variation. Geostatistics and fractal dimensions are the quantitative tools for finding gradients (Phillips and analyzing these 1985).91) indicates a complexpattem of erosion.. The 31 . a fractal approach can make data storage more efficient (Goodchild and Mark 1987). and scale the data according to specified probability rules (Stevens 1989). Iterated function systems have been applied to data compression of digitized pictures.000:1. In a semivari_ram. Peitgen et al. The fractal dimension expresses the complexity of the data. (Reprinted with permission frorru" Phillips. 1986. and data from the contraction process help to determine the fractal dimension (Cipra 1989).CE Diagram 21.D. But two or more contraction mappings will become very complex. Annals of the Association of American Geographers. In the forest context.. Delaware Bay. gradient analysis is part of a method that includes ordination and classification of vegetation.. Iterated function systems consist of several sets of equations that rotate. _. 1 2 S 4 Loa O.. °.3 0 . A single contraction mapping iterated on a bounded set will bring the set to a single point.. These systems are also called contraction mappings--mappings that bring points closer together. Fractals from contraction mappings can be described by the small set of numbers that make up the matrices of the mappings and the time that each mapping is applied. The techniques of fractal analysis can be applied to environmental data to quantify the scales over which one can extrapolate from the sampiing site to the larger area (Krummel 1986)._ 1-2 _ semivariance function helps in this context by o >. New Jersey. 76: 50-62. or a sample from a site may not represent the site's heterogeneity.ST^. variance is plotted against scale.. o 0. A measure of the spatial heterogeneity of a variable would be useful {Phillips 1985). An iterated function system is a data-compression method (Bamsley 1989. Is variability regular enough so that spatial position can be substituted for position along an environmental gradient? Does the measured gradient show deviations from the regular trend that might affect ecological interpretation (Phillips 1985)? A linear series approaching for most gradient analysis D = 2 is too complex (Phillips 1985).0 the environmental variable (McBratney and giving the sampling density needed to represent Webster 1983). Thefractal dimension (1. ! ! ! 0. However. The representation can be exact or approximate (Cipra 1989). so any fractal can be reduced to small data sets. there may be significant deviation from the trend at one sampling site. translate.--Semivariogram of wave attack as an erosion variable acting over many scales.. Spatial analysis of shoreline erosion.. like geographic information systems. 1992). The attractor is usually a fractal. J. Data-compression ratios range from 10:1 to 10. and can be used for checking variability along a gradient. .. Data Storage and Compression For large data sets.

Franklin and Forman 1987). large areas of presettlement vegetation do not exist in most landscapes. To close this review. Fractal geometry may improve the prediction of species-habitat interactions because of its ability to unify patterns of habitat heterogeneity at a variety of scales. FRACTAL APPLICATIONS IN MANAGING FOREST LANDSCAPES section. These lands encompass one-third of the Nation. The maintenance of old-growth forests is one goal of ecosystem management (Crow 1990). Each old-growth patch is surrounded by a buffer zone in which humanrelated activities are restricted. old-growth forests may hold the key to developing practices for sustainable forestry in managed portions of the landscape. Among the many methods of describing landscape structure (see Turner (1989) for review). structural. An alternative involves constructing a network of old-growth patches connected with corridors. area. and functional features. The best management for old-growth patches and buffer zones.g. In addition to old-growth management. 32 . Old-growth forests have unique structural and compositional features that provide critical habitat for associated plant and animal species. Studies of the demography of an animal species often focus on the effects of the spatial pattern of vegetation on distribution. best tools for conducting scales (Milne 1992). Fractal geometry may improve the management of old-growth areas and increase the understanding of specieshabitat interactions. population density) with changes in scale. (1989)). For example. For example. movement. "Fne inference (although not explic 2 ifly shown) is that beetle movement is not related to cover at any one level of resolution. conserving the full complement of compositional. a second area of concern is the maintenance of species diversity. Because the scale at which organisms interact with their environment is not known. locating and designing timber harvests in multiple-use areas can lessen the ecological impact in relation to old-growth areas (e. Results can be used as a basis for mimicking natural disturbance. Habitat suitability models relate vegetation attributes to the presence of the animal of interest (Verner et aL 1986). Movement could be predicted only by integrating cover across scale using the fractal dimension. 1987. Consequently. fractal geometry provides several techniques for measuring the heterogeneity of vegetation. All the methods represent exponential changes in measured quantities (e. may involve a fractal description of heterogeneous organization. Fractals capture much of the intuitive character Old-growth forests have been described as mosaics of groups of trees of various age classes and species (Bonnicksen and Stone 1982). and juxtaposition of habitat. There is a great deal of literature on population response to patchy environments (see Wiens (1976) and Turner (1989) for reviews). O'Neill et al. and persistence. a focus on one scale may yield equivocal results. and fractal geometry has been used to quantify the size-shape relationship of vegetation patches (Krummel et al. and they are one of the analyses at multiple In the previous applications of fractal geometry in many areas of forestry research. Wiens and Mflne (1989) found that beetle movement through a grassland is related to the fractal dimension of grass cover. Noss and Harris 1986).g. Turner et al. we focus on the potential of fractal geometry in resource management. Maintenance of biological diversity has emerged as a significant issue in the management of federal lands. Because of their longevity. Although old-growth. and they are rare. Fractal geometry may help explain the effect of landscape pattern on the spread of disturbances such as fire or insect damage (see. 1988) and the shapes of boundaries between patches (Loehle 1983). we described current of natural heterogeneity. for example. Studies of species-habitat interactions usually measure vegetation at one scale. and they have the potential to support the nationwide diversity of species and ecosystems while providing forest-based tommodities and services. quality.8. with measures of fractal dimension as a basis. fractal descriptions of the juxtaposition of vegetation patches may be used as a basis for reconstructing old-growth forests... Research focuses on the relationship between population persistence and the amount. presettlement ecosystems are often the model for landscape management (Noss 1983. perimeter length.

g. or social organization? Generalities about the relationship between species persistence and vegetation can be used as a basis for better management plans. To summarize this chapter. the causal mechanisms underlying the relationship are not known. and corridors. fractal geometry will help in reconstructing old-growth ecosystems and in understanding and predicting species behavior and persistence. While Wiens and Milne (1989) observe and relate a beetle's movement to the fractal dimension of its habitat. food. gaps. Simulation studies can also evaluate alternative vegetation patterns.g. Palmer suggests that the persistence patterns result from basic ecological principles such as habitat area. Mflne repeats the measurement for different sizes of home range. The method was originally developed to generate landscapes with elevation varying as a function of location. Next. Along with other measures of landscape structure. An even more difficult problem is to determine actions that maintain a desired fractal dimension over time. ecological equivalency. he quantifies resource availability in surrounding ceils for a given size of home range. beetles. The variable depends on its two-dimensional location so that the fractal dimension of the landscape has a particular value. with coexistence of multiple species as the criterion. views crucial to resource managers interested in the effects of alternative management policies on a range of species. interacting species.e. Mflne {1992) provides an innovative description of resource availability for multiple species at different scales using fractal geometry. a general environmental variable is defined for the habitat value of a microsite (Palmer 1992). For each cell in a landscape. physiology. jack rabbits.. Finally. which represents the degree of spatial correlation. Does the degree of fragmentation at a large scale affect orgartism movement on a smaller scale? Do movements change depending on the objective (e.The fractal dimensions of vegetation may be used as explanatory variables for a wide range of questions about individual behavior within a species (Wiens and Mflne 1989). He assumes that species fitness (i. and that the landscape is subdivided into patches each with a particular value of the aggregate variable. The fractal distributions of the resource look much more like real landscapes. antelope) respond in the same way to changes in vegetation structure through habitat fragmentation? Can differences be explained by life history strategies. Suppose that species persistence is enhanced when vegetation has a certain fractal dimension. fractal geometry has been successfully applied to the description of heterogeneous landscapes. he computes elevational maps of the resource densities for different home range sizes. Do organisms that operate at different scales within the same vegetation type (e. given growth and disturbance functions. It will take great effort to conduct similar studies with larger organisms or multiple species. Palmer {1992) simulates the effects of a range of spatial patterns on the persistence of species with different fitness functions. Saupe 1988). search for cover. This is a landscape target for vegetation management. Methods are needed for other quantities (e. or mate)? Do population density and stability depend on the fractal dimension of the vegetation? There are also interesting questions about comparative behavior across species (Wiens 1989). Instead.. The fractal dimension describes the spatial variability of the resource. These provide multi-scale views of areas of interior habitat. body sizes.g. competitive ability and fecundity) is related to an aggregate environmental variable in a deterministic way. perimeter length or area) used to estimate fractal dimensions. The fractal dinlension of the landscape is estimated as a function of the location and density of resource availability. mass effect. Palmer (1992) simulates fractal landscapes using the midpoint displacement method with successive random additions (Feder 1988. Instead of elevation. How is a desired vegetation structure created from a given starting point? Although no one has looked at this problem directly. elements have been addressed. simulation studies can generate hypotheses about the effects of spatial patterns of resources on the demography of multiple. The variable values are spatially arranged so that the landscape has a certain fractal dimension.. Work to date has shown that fractal geometry is useful 33 . and global biological constraints. The advantage of using fractal landscapes is that the environmental variable is continuously distributed in a statistically selfsimilar way rather than being either homogeneous or random.

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St. jagged. mapping. MN: U. NC-170.. Leary. Rep.S. forest mensuration. 43 p. Because most of what we measure in the forest is discontinuous. This study reviews the literature on fractal geometry and its applications to forest measurements. Tech. vegetation geometry. The fraetal forest: fxactal geometry and applications in forest science. KEY WORDS: Fractal description. Forest Service._rtmer. RoKe A. North Central Forest Experiment Station.I. Haight. ikactal geomettT has potentiN for improving the precision of measurement and description. and tkagmented. Robect G. Department of Agriculture. Paul. wildlife habitat .. microclimatology. Gen.. Nancy D. 1994. Fractal geometry is a tool lbr descI:tbtng and amalyzing h-regularity.

EHP_RIIT_NT STRTtON . quality. As a new generation of forests emerges in our region. and ownership of the forests. and (2) Reconciling the conflicting demands of the people who use them.Our job at the North Central Forest Experiment Station is discoveling and creating new knowledge and technolog_ in the field of natural resources arid conveying this information to the people who can use it. managers are confronted with two unique challenges: (1) Dealing with the great diversity in composition. Helping the forest manager meet these challenges while protecting the environment is what research at North Central is all about.