How Do Grazers Achieve Their Distribution? A Continuum of Models from Random Diffusion to the Ideal Free Distribution Using Biased Random Walks

A conceptual model is described for generating distributions of grazing animals, according to their searching behavior, to investigate the mechanisms animals may use to achieve their distributions. The model simulates behaviors ranging from random diffusion, through taxis and cognitively aided navigation (i.e., using memory), to the optimization extreme of the Ideal Free Distribution. These behaviors are generated from simulation of biased diffusion that operates at multiple scales simultaneously, formalizing ideas of multiple‐scale foraging behavior. It uses probabilistic bias to represent decisions, allowing multiple search goals to be combined (e.g., foraging and social goals) and the representation of suboptimal behavior. By allowing bias to arise at multiple scales within the environment, each weighted relative to the others, the model can represent different scales of simultaneous decision‐making and scale‐dependent behavior. The model also allows different constraints to be applied to the animal's ability (e.g., applying food‐patch accessibility and information limits). Simulations show that foraging‐decision randomness and spatial scale of decision bias have potentially profound effects on both animal intake rate and the distribution of resources in the environment. Spatial variograms show that foraging strategies can differentially change the spatial pattern of resource abundance in the environment to one characteristic of the foraging strategy.

[1]  James Smith,et al.  The Food Searching Behaviour of Two European Thrushes. Ii: the Adaptiveness of the Search Patterns , 1974 .

[2]  William E. Grant,et al.  AN ARTIFICIAL INTELLIGENCE MODELLING APPROACH TO SIMULATING ANIMAL/HABITAT INTERACTIONS , 1988 .

[3]  G. Pickup,et al.  Modelling patterns of defoliation by grazing animals in rangelands , 1994 .

[4]  Simon Benhamou,et al.  A comparative analysis of spatial memory processes , 1995, Behavioural Processes.

[5]  G. A. Parker,et al.  Ideal free distributions when individuals differ in competitive ability: phenotype-limited ideal free models , 1986, Animal Behaviour.

[6]  Aníbal Ollero,et al.  Fuzzy Methodologies for Interactive Multicriteria Optimization , 1980, IEEE Transactions on Systems, Man, and Cybernetics.

[7]  Source-sink dynamics and the coexistence of species on a single resource , 1997 .

[8]  Spatial heterogeneity, pollinator behaviour and pollinator-mediated gene flow: bumblebee movements in variously aggregated rows of oil-seed rape , 1997 .

[9]  K. Warburton,et al.  Tendency-distance models of social cohesion in animal groups. , 1991, Journal of Theoretical Biology.

[10]  William J. Sutherland,et al.  Aggregation and the `ideal free ` distribution , 1983 .

[11]  D. L. GUNN,et al.  Classification of Taxes and Kineses , 1937, Nature.

[12]  R. W. Richards,et al.  Characteristics of spatial memory in cattle , 1989 .

[13]  John M. Fryxell,et al.  Forage Quality and Aggregation by Large Herbivores , 1991, The American Naturalist.

[14]  John R. Krebs,et al.  INDIVIDUAL DECISIONS AND THE DISTRIBUTION OF PREDATORS IN A PATCHY ENVIRONMENT. II. THE INFLUENCE OF TRAVEL COSTS AND STRUCTURE OF THE ENVIRONMENT , 1991 .

[15]  D Grünbaum,et al.  Using Spatially Explicit Models to Characterize Foraging Performance in Heterogeneous Landscapes , 1998, The American Naturalist.

[16]  Audrey E. Cramer,et al.  Vervet monkeys as travelling salesmen , 1997, Nature.

[18]  S. Benhamou,et al.  Spatial analysis of animals' movements using a correlated random walk model* , 1988 .

[19]  Peter Turchin,et al.  Translating Foraging Movements in Heterogeneous Environments into the Spatial Distribution of Foragers , 1991 .

[20]  E. Charnov Optimal foraging, the marginal value theorem. , 1976, Theoretical population biology.

[21]  Hugh P. Possingham,et al.  The Distribution and Abundance of Resources Encountered by a Forager , 1989, The American Naturalist.

[22]  J. Bolhuis,et al.  Exponential decay of spatial memory of rats in a radial maze. , 1986, Behavioral and neural biology.

[23]  David Saltz,et al.  Forging at Different Spatial Scales: Dorcas Gazelles Foraging for Lilies in the Negev Desert , 1994 .

[24]  Manfred Milinski,et al.  LONG-TERM MEMORY FOR FOOD PATCHES AND IMPLICATIONS FOR IDEAL FREE DISTRIBUTIONS IN STICKLEBACKS' , 1993 .

[25]  G. Pierce,et al.  Eight Reasons Why Optimal Foraging Theory Is a Complete Waste of Time , 1987 .

[26]  S. L. Lima,et al.  Landscape-level perceptual abilities in white-footed mice : perceptual range and the detection of forested habitat , 1997 .

[27]  Lotfi A. Zadeh,et al.  Optimality and non-scalar-valued performance criteria , 1963 .

[28]  S. Benhamou,et al.  How animals use their environment: a new look at kinesis , 1989, Animal Behaviour.

[29]  J M Smith,et al.  Evolution and the theory of games , 1976 .

[30]  W. Morris,et al.  Population Consequences of Constitutive and Inducible Plant Resistance: Herbivore Spatial Spread , 1997, The American Naturalist.

[31]  E. Menzel Chimpanzee Spatial Memory Organization , 1973, Science.

[32]  T. Collett,et al.  Insect navigation en route to the goal: multiple strategies for the use of landmarks , 1996, The Journal of experimental biology.

[33]  S. Fretwell,et al.  On territorial behavior and other factors influencing habitat distribution in birds , 1969 .

[34]  P. Day The Organisation of Learning , 1977 .

[35]  Yosef Cohen,et al.  Spatial Heterogeneities, Carrying Capacity, and Feedbacks in Animal-Landscape Interactions , 1997 .

[36]  Derek W. Bailey,et al.  Large Herbivore Foraging and Ecological HierarchiesLandscape ecology can enhance traditional foraging theory , 1987 .

[37]  A. Houston,et al.  The distribution of animals between resources: a compromise between equal numbers and equal intake rates , 1995, Animal Behaviour.

[38]  A. Etienne,et al.  The effect of a single light cue on homing behaviour of the golden hamster , 1990, Animal Behaviour.

[39]  A. Bennett,et al.  Do animals have cognitive maps? , 1996, The Journal of experimental biology.

[40]  Anthony J. Parsons,et al.  The use of spatial memory by grazing animals to locate food patches in spatially heterogeneous environments: an example with sheep , 1996 .

[41]  Simon Benhamou,et al.  Efficiency of area-concentrated searching behaviour in a continuous patchy environment* , 1992 .

[42]  J. O'Keefe,et al.  The hippocampus as a spatial map. Preliminary evidence from unit activity in the freely-moving rat. , 1971, Brain research.

[43]  Association of relative food availabilities and locations by cattle. , 1989 .

[44]  S. Al-Moghrabi,et al.  Inorganic carbon uptake for photosynthesis by the symbiotic coral-dinoflagellate association II. Mechanisms for bicarbonate uptake , 1996 .

[45]  N. Andrew Spatial Heterogeneity, Sea Urchin Grazing, and Habitat Structure on Reefs in Temperate Australia , 1993 .

[46]  Yosef Cohen,et al.  Linking Moose Population and Plant Growth Models with a Moose Energetics Model , 1998, Ecosystems.

[47]  S. Focardi,et al.  A mathematical framework for optimal foraging of herbivores , 1995, Journal of mathematical biology.

[48]  J. Krebs,et al.  Memory for the location of stored food in marsh tits , 1981, Animal Behaviour.

[49]  C. Patlak Random walk with persistence and external bias , 1953 .

[50]  W. Alt Biased random walk models for chemotaxis and related diffusion approximations , 1980, Journal of mathematical biology.

[51]  E. Laca,et al.  Mechanisms that result in large herbivore grazing distribution patterns. , 1996 .

[52]  Monica G. Turner,et al.  A landscape simulation model of winter foraging by large ungulates , 1993 .

[53]  S. L. Lima,et al.  Towards a behavioral ecology of ecological landscapes. , 1996, Trends in ecology & evolution.

[54]  R. Morris Spatial Localization Does Not Require the Presence of Local Cues , 1981 .

[55]  Yosef Cohen,et al.  A SPATIALLY EXPLICIT MODEL OF MOOSE FORAGINGAND ENERGETICS , 1997 .

[56]  J. G. Skellam Random dispersal in theoretical populations , 1951, Biometrika.

[57]  明 大久保,et al.  Diffusion and ecological problems : mathematical models , 1980 .

[58]  SIMON BENHAMOU,et al.  No evidence for cognitive mapping in rats , 1996, Animal Behaviour.

[59]  George Sugihara,et al.  Fractals: A User's Guide for the Natural Sciences , 1993 .

[60]  J. Krebs,et al.  INDIVIDUAL DECISIONS AND THE DISTRIBUTION OF PREDATORS IN A PATCHY ENVIRONMENT , 1988 .

[61]  D. J. Anderson,et al.  Optimal foraging and the traveling salesman , 1983 .

[62]  Stuart A. Altmann,et al.  Baboons, space, time and energy , 1974 .

[63]  N. B. Kotliar,et al.  Multiple scales of patchiness and patch structure: a hierarchical framework for the study of heterogeneity , 1990 .

[64]  C. Clark,et al.  Dynamic Modeling in Behavioral Ecology , 2019 .

[65]  Keith D. Farnsworth,et al.  Beyond the Ideal Free Distribution: More General Models of Predator Distribution , 1997 .

[66]  Innchyn Her A symmetrical coordinate frame on the hexagonal grid for computer graphics and vision , 1993 .