Adaptive models for large herbivore movements in heterogeneous landscapes Research article First Online: 01 April 2005 DOI:
10.1007/s10980-005-0061-9 Cite this article as: Morales, J., Fortin, D., Frair, J.L. et al. Landscape Ecol (2005) 20: 301. doi:10.1007/s10980-005-0061-9 Abstract
It is usually assumed that landscape heterogeneity influences animal movements, but understanding of such
processes is limited. Understanding the effects of landscape heterogeneity on the movements of large
herbivores such as North American elk is considered very important for their management. Most simulation
studies on movements of large herbivores use predetermined behavioral rules based on empirical
observations, or simply on what seems reasonable for animals to do. Here we did not impose movement
rules but instead we considered that animals had higher fitness (hence better performance) when they
managed to avoid predators, and when they acquired important fat reserves before winter. Individual
decision-making was modeled with neural networks that received as input those variables suspected to be
important in determining movement efficiency. Energetic gains and losses were tracked based on known
physiological characteristics of ruminants. A genetic algorithm was used to improve the overall performance
of the decision processes in different landscapes and ultimately to select certain movement behaviors.
We found more variability in movement patterns in heterogeneous landscapes. Emergent properties of
movement paths were concentration of activities in well-defined areas and an alternation between small,
localized movement with larger, exploratory movements. Even though our simulated individuals moved
shorter distances that actual elk, we found similarities in several aspects of their movement patterns such as
in the distributions of distance moved and turning angles, and a tendency to return to previously visited
References Adler, P.B., Raff, D.A., Lawenroth, W.K. 2001 The effect of grazing on the spatial heterogeneity of vegetation Oecologia 128 465 479 CrossRef Google Scholar Anderson, J.A. 1995An introduction to neural networks MIT Press CambridgeMassachusetts, USA Google Scholar Bailey, D.W., Gross, J.E., Laca, E.A., Rittenhouse, L.R., Coughenour, M.B., Swift, D.M., Sims, P.L. 1996 Mechanisms that result in large herbivore grazing distribution patterns Journal of Range Management 49 386 400 CrossRef Google Scholar Beecham, J.A., Farnsworth, K.D. 1998 Animal foraging from an individual perspective: an object orientated model Ecological Modelling 113 141 156 CrossRef Google Scholar Belovsky, G.E. 1984 Herbivore optimal foraging: a comparative test of three models American Naturalist 124 97 115 CrossRef Google Scholar Bergman, C.M., Fryxell, J.M., Gates, C.C. 2000 The effect of tissue complexity and sward height on the functional response of Wood Bison Functional Ecology 14 61 69 CrossRef Google Scholar Bolker, B., Pacala, S.W. 1997 Using moment equations to understand stochastically driven spatial pattern formation in ecological systems Theoretical Population Biology 52 179 197 PubMed CrossRef Google Scholar Cain, M.L. 1991 When do treatment differences in movement behaviors produce observable differences in long-term displacements? Ecology 72 2137 2142 CrossRef Google Scholar Cook, J.G. 2002 Nutrition and food Toweill, D.E. Thomas, J.W. eds. North american elk: Ecology and management Smithsonian Institution Press Washington, USA 259 349 Google Scholar Cook, R.C., Cook, J.G., Mech, L.D. 2004 Nutritional condition of northern Yellowstone elk Journal of Mammalogy 85 714 722 CrossRef Google Scholar Delgiudice, G.D., Moen, R.A., Singer, F.J., Riggs, M.R. 2001 Winter nutritional restriction and simulated body condition of Yellowstone elk and bison before and after the fires of 1988 Wildlife Monographs 147 1 60 Google Scholar Dieckmann, U., Law, R., Metz, J.A.J. 2000The geometry of ecological interactions: simplifying spatial complexity Cambridge University Press CambridgeUK CrossRef Google Scholar Farnsworth, K.D., Beechman, J.A. 1999 How do grazers achieve their distribution? a continuum of models from diffusion to the ideal free distribution using biased random walks American Naturalist 153 509 526 CrossRef Google Scholar Farnsworth, K.D., Illius, A.W. 1996 Large grazers back in the fold: Generalizing the prey model to incorporate mammalian herbivores Functional Ecology 10 678 680 Google Scholar Fortin, D. 2002 Optimal searching behaviour: the value of sampling information Ecological Modelling 153 279 290 CrossRef Google Scholar Fortin, D. 2003 Searching behavior and use of sampling information by free-ranging bison ( Bos bison) Behavioral Ecology and Sociobiology 54 194 203 Google Scholar Fortin, D., Boyce, M.S., Merrill, E.H. 2004 Multi-tasking by mammalian herbivores: overlapping processes during foraging Ecology 85 2312 2322 CrossRef Google Scholar Fortin, D., Fryxell, J.M., Pilote, R. 2002 The temporal scale of foraging decisions in bison Ecology 83 970 982 CrossRef Google Scholar
Frair J.L., Merrill E.H., Beyer H.L., Morales J.M., Visscher D.R. and Fortin D. 2004. Determining scales of movement by elk (Cervus elaphus) and their responses to heterogeneity in forage resources and predation risk. Landscape Ecology.
Fryxell, J.M. 1991 Forage quality and aggregation by large herbivores American Naturalist 138 478 498 CrossRef Google Scholar Fryxell, J.M., Greever, J., Sinclair, A.R.E. 1988 Why are migratory ungulates so abundant American Naturalist 131 781 798 CrossRef Google Scholar Goldberg, D.E. 1989Genetic algorithms in searchoptimization and machine learning Addison-Wesley Pub. Co. Reading, Massachusetts, USA Google Scholar Grimm, V., Frank, K., Jeltsch, F., Brandl, R., Uchmanski, J., Wissel, C. 1996 Pattern-oriented modelling in population ecology Science of the Total Environment 183 151 166 CrossRef Google Scholar Gross, J.E., Shipley, L.A., Hobbs, N.T., Spalinger, D.E., Wunder, B.A. 1993 Functional response of herbivores in food-concentrated patches: Tests of a mechanistic model Ecology 74 778 791 CrossRef Google Scholar Hudson, R.J., Frank, S. 1987 Foraging ecology of bison in Aspen boreal habitats Journal of Range Management 40 71 75 CrossRef Google Scholar Hudson, R.J., White, R.G. 1985Bioenergetics of wild herbivores CRC Press Boca Raton, FloridaUSA Google Scholar Huse, G., Giske, J. 1998 Ecology in Mare Pentium: an individual-based spatio-temporal model for fish with adapted behaviour Fisheries Research 37 163 178 CrossRef Google Scholar Huse, G., Strand, E., Giske, J. 1999 Implementing behaviour in individual-based models using neural networks and genetic algorithms Evolutionary Ecology 13 469 483 CrossRef Google Scholar Illius, A.W., Gordon, I.J. 1987 The allometry of food Intake in grazing ruminants Journal of Animal Ecology 56 989 999 CrossRef Google Scholar Illius, A.W., Gordon, I.J. 1992 Modelling the nutritional ecology of ungulate herbivores: evolution of body size and competitive interactions Oecologia 89 428 434 Google Scholar Jiang, Z., Hudson, R.J. 1992 Estimating forage intake and energy requirements of free-ranging wapiti Cervus elaphus Canadian Journal of Zoology 70 675 679 CrossRef Google Scholar Johnson, C.J., Parker, K.L., Heard, D.C., Gillingham, M.P. 2002 Movement parameters of ungulates and scale-specific responses to the environment Journal of Animal Ecology 71 225 235 CrossRef Google Scholar Jung, H.G., Koong, L.J. 1985 Effects of hunger satiation on diet quality by grazing sheep Journal of Range Management 38 302 305 CrossRef Google Scholar Kareiva, P., Odell, G. 1987 Swarms of predators exhibit “preytaxis” if individual predators use area-restricted search American Naturalist 130 233 270 CrossRef Google Scholar Keeling, M.J., Wilson, H.B., Pacala, S.W. 2000 Reinterpreting spacetime lags, and functional responses in ecological models Science 290 1758 1761 PubMed CrossRef Google Scholar Kot, M., Lewis, M., den Driessche, P. 1996 Dispersal data and the spread of invading organisms Ecology 77 2027 2042 CrossRef Google Scholar Law, R., Murrell, D.J., Dieckmann, U. 2003 Population growth in space and time: spatial logistic equations Ecology 84 252 262 CrossRef Google Scholar Lewis, M.A. 1997 Variability, patchiness, and jump dispersal in the spread of an invading population Tilman, D. Kareiva, P. eds. Spatial Ecology: the Role of Space in Population Dynamics and Interespecific Interactions Princeton University Press Princeton, New Jersey, USA 46 69 Google Scholar Lewis, M.A., Murray, J.D. 1993 Modelling territoriality and wolf-deer interactions Nature 366 738 740 CrossRef Google Scholar Lima, S.L., Zollner, P.A. 1996 Towards a behavioral ecology of ecological landscapes Trends in Ecology and Evolution 11 131 135 PubMed CrossRef Google Scholar McNaughton, S.J. 1984 Grazing lawns: animals in herds, plant formand coevolution American Naturalist 124 863 886 CrossRef Google Scholar Mitchell, M., Forrest, S. 1995 Genetic algorithms and artificial life Langton, C.G. eds. Artificial Life: An Overview MIT Press CambridgeMassachusetts, USA 267 289 Google Scholar Mitchell, W.A., Lima, S.L. 2002 Predator-prey shell games: large-scale movement and its implications for decision-making by prey Oikos 99 249 259 CrossRef Google Scholar Moen, R., Pastor, J., Cohen, Y. 1997 A spatially explicit model of moose foraging and energetics Ecology 78 505 521 Google Scholar Morales, J.M. 2002 Behavior at habitat boundaries can produce leptokurtic movement distributions American Naturalist 160 531 538 PubMed CrossRef Google Scholar Morales, J.M., Ellner, S.P. 2002 Scaling up movement in heterogeneous landscapes: the importance of behavior Ecology 83 2240 2247 CrossRef Google Scholar Morales, J.M., Haydon, D.T., Frair, J., Holsinger, K.E., Fryxell, J.M. 2004 Extracting more out of relocation data: building movement models as mixtures of random walks Ecology 89 2436 2445 CrossRef Google Scholar Murrell, D.J., Law, R. 2000 Beetles in fragmented woodlands: a formal framework fro dynamics of movement in ecological landscapes Journal of Animal Ecology 69 471 483 CrossRef Google Scholar Parker, K.L., Robbins, C.T., Hanley, T.A. 1984 Energy expenditures for locomotion by mule deer and elk Journal of Wildlife Management 48 474 488 CrossRef Google Scholar Pastor, J., Moen, R., Cohen, Y. 1997 Spatial heterogeneities, carrying capacity, and feedbacks in animal-landscape interactions Journal of Mammalogy 78 1040 1052 CrossRef Google Scholar Robbins, C.T. 1993Wildlife feeding and nutrition, second edition Academic Press San DiegoCaliforniaUSA Google Scholar Schmidt, K.A., Ostfeld, R.S. 2003 Mice in space: Space use predicts the interaction between mice and songbirds Ecology 84 3276 3283 CrossRef Google Scholar Schmitz, O.J., Beckerman, A.P., O'Brien, K.M. 1997 Behaviorally mediated trophic cascades: Effects of predation risk on food web interactions Ecology 78 1388 1399 CrossRef Google Scholar Shipley, L.A., Spalinger, D.E., Gross, J.E., Hobbs, N.T., Wunder, B.A. 1996 The dynamics and scaling of foraging velocity and encounter rate in mammalian herbivores Functional Ecology 10 234 244 CrossRef Google Scholar Spalinger, D.E., Hobbs, N.T. 1992 Mechanisms of foraging in mammalian herbivores: new models of functional response American Naturalist 140 325 348 PubMed CrossRef Google Scholar Strand, E., Huse, G., Giske, J. 2002 Artificial evolution of life history and Behavior American Naturalist 159 624 644 PubMed CrossRef Google Scholar Tilman, D. Kareiva, P. eds. 1997Spatial Ecology: The Role of Space in Population Dynamics and Interespecific Interactions Princeton University Press Princeton, New Jersey, USA Google Scholar Turchin, P. 1998Quantitative Analysis of Movement: Measuring and Modeling Population Redistribution in Animals and Plants Sinauer Associates SunderlandMassachusetts, USA Google Scholar Turchin, P., Batzli, G. 2001 Availability of food and the population dynamics of arvicoline rodents Ecology 82 1521 1534 CrossRef Google Scholar Turner, M.G., Wu, Y., Romme, W.H., Wallace, L.L. 1993 A landscape simulation model of winter foraging by large ungulates Ecological Modelling 69 163 184 CrossRef Google Scholar Turner, M.G., Wu, Y., Wallace, L.L., Romme, W.H., Brenkert, A. 1994 Simulating winter interactions among ungulates, vegetation, and fire in northern Yellowstone Park Ecological Applications 4 472 496 CrossRef Google Scholar Whitley, D. 2001 An overview of evolutionary algorithms: practical issues and common pitfalls Information and Software Technology 43 817 831 CrossRef Google Scholar Wiens, J.A., Stenseth, N.C., Van Horne, B., Ims, R.A. 1993 Ecological mechanisms and landscape ecology Oikos 66 369 380 CrossRef Google Scholar Wilmshurst, J., Fryxell, J., Hudson, R. 1995 Forage quality and patch choice by wapiti ( Cervus elaphus) Behavioral Ecology 6 209 217 CrossRef Google Scholar
Wilmshurst J.F., Fryxell J.M. and Bergman C.M. 2000. The allometry of patch selection in ruminants. Proc. R. Soc. Lon, Ser B 267: 345–349.
Google Scholar Zollner, P.A., Lima, S.L. 1999 Search strategies for landscape-level interpatch movements Ecology 80 1019 1030 CrossRef Google Scholar