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BCB 322: Landscape Ecology. Lecture 6: Emerging patterns I: Heterogeneity. Heterogeneity. Patterns arise in any landscape as a result of the underlying processes Disturbance and fragmentation are closely allied, and have significant impacts on the environment

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BCB 322: Landscape Ecology

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bcb 322 landscape ecology

BCB 322:Landscape Ecology

Lecture 6:

Emerging patterns I:


  • Patterns arise in any landscape as a result of the underlying processes
  • Disturbance and fragmentation are closely allied, and have significant impacts on the environment
  • Heterogeneity is the main pattern in any landscape, & is inherent at all scales
  • “The uneven, non-random distribution of objects” (Forman, 1995)
  • Since every pattern in a landscape results from, and produces new processes, heterogeneity is important for landscape function
  • The analysis of heterogeneity is fundamental to the understanding of landscape functions and spatial ecological processes


  • Three types to be considered:
    • Spatial: variation in space, either horizontally (as under human disturbance regimes) or vertical (uneven vegetation distribution above ground – generally natural)
    • Temporal: similar to spatial, but implies variation for a


single point over time. Two areas may have the same spatial variation, but differ in time.

    • Functional: variation in distribution of communities/ populations. Linked to life history of organisms
  • Soil composition is a strong driver of heterogeneity, & it varies strongly from individual plots (1-10m) to the full landscape (Becher, 1995). It can also vary vertically in the soil profile
  • The effect of geological heterogeneity is unpredictable, and a probabilistic approach is used when modelling it
  • By creating borders and edge effects, additional processes are set up in the landscape, influencing the flux of materials
  • Since plant and animal species respond fairly rapidly to changes in mosaic heterogeneity, minor variations can often be observed with remote sensing techniques.
  • Effectively, heterogeneity can exaggerate biological-environment interactions (eg: skylarks avoid small prairie spaces, even though they are functionally similar to larger patches) (Farina, 1998)
  • Heterogeneous landscapes can show several different stable configurations, and can shift between them rapidly given sufficient incitement.
  • This is termed a polyclimactic state, and it is determined by variations in deterministic & stochastic factors (wind, climate extremes, climate change, edaphic conditions) and internal factors (disease, predation, invasion)
  • Hence disturbance can play a role in maintaining heterogeneity and preventing stable equilibria developing.
  • A moderate disturbance regime can increase heterogeneity, but it depends on the initial conditions
  • Homogeneous system: heterogeneity increases to the midpoint, then falls
  • Highly heterogeneous: after an initial gain in heterogeneity, there is a rapid dropoff (Kolasa & Rollo, 1991)
  • Obviously, some disturbance can be useful for maintaining heterogeneity.
  • Likewise, heterogeneity can impact on the disturbance regime:
    • fires spread less readily in a mixed woodland than in a pure coniferous matrix (reduced disturbance)
    • predators (eg: foxes) in woodlots & parklands in an agricultural environment can impact on livestock (increased disturbance)
  • Species distribution depend on community heterogeneity: large sample areas include more species, and thus should be more similar than smaller ones with a limited selection of species.


  • Three main types of heterogeneity structure (Addicot et al., 1987):
    • Divided homogeneous (suitable patches in an unsuitable matrix)
    • Undivided heterogeneous (patches of varying suitability)
    • Divided heterogeneous (varying suitability patches in an unsuitable matrix
  • Fire, grazing & the two in combination were tested in grasslands (Glenn et al, 1992)
  • Locally, burning seemed to have higher heterogeneity than grazing, whilst the corollary was true at a regional scale.
  • Overall, untreated local plots had the most heterogeneity, but regional responses varied to a large degree, depending on season of burning (spring burning then grazing increased heterogeneity, autumn burns reduced it)
  • This variation makes studies complicated – some processes depend on patchiness, but not all.
  • General, organismal responses are a good measure by which to assess relevant scale. (neighbourhood scale)
  • Neighbourhood for vagile species is obviously their territory or resource area
  • For sessile species it is more complex, but it can be estimated according to the areas from which food, predators and other foragers come.


  • Spatial heterogeneity is one of the main factors determining bird species diversity in the landscape (MacArthur et al., 1962)
  • Bird diversity was also higher in shrubby areas because shrubs have higher heterogeneity than tree areas (even though trees have more variation in canopy structure)
  • Animals also drive heterogeneity:

grazing animals at high densities can permanently alter vegetation cover by compacting soil and removing vegetation (eg: rural goat grazing in Zimbabwe)

  • Animals can even alter geomorphology, & all sizes can impact heterogeneity:
    • Insects (ants & termites shift soil, dung beetles enrich soil)
    • Small mammals (moles & rabbits digging soil, other rodents distributing seed)
    • Medium mammals (beaver dams, pigs rooting)
    • Large mammals (elephants/buffalo opening up cover)
    • All sizes of birds affect seed distribution & enrich soil with droppings
  • Species-level heterogeneity requires individuals to modify their foraging habits – in larger patches they can afford to spend less time foraging
  • Time spent within a patch varies as the square of the linear dimension of the patch, whilst travel time between them varies linearly (ie): large patches are preferable and used in a more specialised way than small patches. (MacArthur & Pianka, 1966)
  • For large species, it has been found that they respond to landscape-level heterogeneity, but not locally (bison, Wallace et al., 1995)
  • Thus, they move in a determined manner (non-randomly) between patches, but within patches move randomly, minimising energy use (optimal foraging)
  • Similar observations of sheep grazing showed that they generally moved directly to the nearest plant, whilst occasionally moving between plants
  • It is possible that heterogeneity therefore plays a role in determining optimal foraging strategies for species, and may even enhance the efficiency
animal movements
Animal movements
  • Certainly, movement within a matrix is determined by suitable/unsuitable patches, preventing animals from moving in straight lines (Johnson et al., 1992)
  • It may also play a role in the recollection of routes in a home range (homing ability)
  • Bumble bees’ foraging routes are generally longer in uniform stands, whilst in a varied stand they tend to backtrack more often
  • Certainly digger wasps (small range) use landmarks to find their way about, and are prepared to fly further in a varied landscape
animal movements1
Animal movements
  • Bees exhibit a similar behaviour
  • Homing ability drops off fairly quickly in flat or uniform landscapes (Plowright & Galen, 1985)
  • In comparison, in mountainous areas they can return from as far off as 9km, due to increased landmarks
metrics of heterogeneity
Metrics of heterogeneity
  • We can measure the heterogeneity of a landscape in terms of any resource (soil structure, plant diversity, biomass, thicket structure, animal distributions…)
  • The variation in structure hence reflects changing functions and processes in the landscape
  • In order to assess this, we use several different metrics, each of which considers different aspects of the structure.
    • Fractal dimension (measures the complexity of edges)
    • Contagion (the extent of aggregation of patches)
    • Evenness (measures number of different patch types & their proportions in the landscape)
    • Patchiness (contrasts neighbouring patches in a matrix)
  • Li & Reynolds (1994) measured these different metrics against four components of heterogeneity
  • Heterogeneity is the principal characteristic of any landscape
  • It varies due to underlying processes, and affects these processes in turn, by initiating or amplifying biological interactions
  • Can be considered in terms of temporal, spatial or functional components
  • Affects disturbance, although in both a positive & a negative manner
  • Heterogeneity affects animal processes (eg, grazing efficiency of ungulates) and is likewise affected by the interactions of animals with landscape functional components
  • Measured using different indices, including contagion, fractal dimension, evenness & patchiness
  • Addicot, J.F., Aho, J.M. & Antolin, M.F. (1987) Ecological neighbourhoods: scaling environmental patterns. Oikos49: 340-346
  • Becher, H.H. (1995) On the importance of soil homogeneity when evaluating field trials. Journal of Agronomy & Crop Science74: 33-40
  • Farina, A. (1998) Principles and Methods in Landscape Ecology. Chapman and Hall, London, UK
  • Forman, R.T.T. (1995) Land Mosaics. The ecology of landscapes and regions. Cambridge University Press, Cambridge.
  • Glenn, S.M., Collins, S.L. & Gibson, D.J. (1992) Disturbance in tallgrass prairies: local and regional effects on community heterogeneity. Landscape Ecology. 7: 243-251.
  • Johnson, A.R., Wiens, J.A., Milne, B.T. & Crist, T.O. (1992)Animal movements and population dynamics in heterogeneous landscapes. Landscape Ecology 7: 47-58
  • Kolasa, J. & Rollo, C.D. (1991) Introduction: the heterogeneity of heterogeneity: a glossary. In: Kolasa, J. and Pickett, S.T.A. Ecological heterogeneity. Springer-Verlag, New York, pp 1-23
  • Li, H. & Reynolds, J.F. (1994) A simualtion experiment to quantify spatial heterogeneity in categorical maps. Ecology75:2446-2455.
  • MacArthur, R.H., MacArthur, J.W. & Preer, J. (1962) On bird species diversity. II. Prediction of bird census from habitat measurements, American Naturalist 96: 167-174
  • MacArthur, R.H. & Pianka, E.R. (1966) On optimal use of patchy environment. American Naturalist100: 603-610
  • Plowright, R.C. & Galen, C/ (1985) Landmarks or obstacles: the effect of spatial heterogeneity on bumblebee foraging behaviour. Oikos44: 459-464