Scale‐Dependence in Ecological Systems


Scale has a profound influence on how we conduct ecological studies, interpret results and understand the links between processes operating at different rates. All of these factors profoundly influence our ability to predict responses to change. The ecological patterns and variability we observe range from millimetres to across ocean basins and from seconds to the expanse of evolutionary history. Patterns apparent at one scale can collapse to noise when viewed from other scales, indicating that perceptions of the importance of different processes vary in a scale‐dependent manner. Moreover, rather than the environment simply providing an arena within which organisms are born grow and die, many organisms interact with the environment, altering it for both for themselves and for other species. Because of these factors, studying ecological systems is far from simple and scale needs to be considered in study design and analysis.

Key Concepts

  • The pattern you see depends on the scale at which it is studied.
  • Threats from human activities occur at varying scales, from small point sources to large diffuse threats and changes in disturbance regimes across landscapes.
  • How organisms interact with the environment depends on how they perceive it and how patchy it is.
  • How an organism moves and how far it can move is crucial to how an organism perceives and responds to their environment.
  • Many organisms can alter their environment, both for themselves and for other species.
  • Connectivity between locations and habitats does occur not only for organisms but also for ecosystem processes and services.
  • Because most processes are scale dependent, studies must explicitly consider scale in their design.
  • Understanding both the scale of threats and the scale over which species live are essential for successful management and conservation.

Keywords: scale; mobility; meta‐populations; meta‐communities; meta‐ecosystems; study design; conservation; protected area networks

Figure 1. Three categories of species mobility and their implications for integrations of spatial variability over time. Species that migrate seasonally will exhibit home ranges (initial flat section of curve) much smaller than their migration distance and most of the increase in spatial scale will be incorporated within a year. Species that widely disperse within one generation as larvae, juveniles or adults (e.g. some marine invertebrates and terrestrial insects) will still exhibit home ranges on short temporal scales and dispersal over multiple generations is still likely to change geographical distributions. Species with limited dispersal at all life stages will have defined home ranges and their spatial distributions will change only slowly over multiple generations. NB actual intercepts, slopes and inflexion points will depend on individual species characteristics and the media through which they move (air, land and water).
Figure 2. In hierarchy theory, environmental processes set the background for small‐scale biotic processes. In multiscale theory, both environmental and biotic processes operate across the potential range of scales with interactions occurring between them.


Allen TFH and Starr TB (1982) Hierarchy Perspectives for Ecological Complexity. Chicago, IL: University of Chicago Press.

Arrhenius O (1921) Species and area. Journal of Ecology 9: 95–99.

Barry JP and Dayton PK (1991) Physical heterogeneity and the organisation of marine communities. In: Kolasa K and Pickett STA (eds) Ecological Heterogeneity, pp. 270–320. New York, NY: Springer.

Baselga A (2010) Partitioning the turnover and nestedness components of beta diversity. Global Ecology and Biogeography 19: 134–143.

Botsford LW, Hastings A and Gaines SD (2001) Dependence of sustainability on the configuration of marine reserves and larval dispersal distance. Ecology Letters 4: 144–150.

Bruno JF, Stachowicz JJ and Bertness MD (2003) Inclusion of facilitation into ecological theory. Trends in Ecology & Evolution 18: 119–125.

Coco G, Thrush SF, Green MO and Hewitt JE (2006) The role of feedbacks between bivalve (Atrina zelandica) density, flow and suspended sediment concentration on patch stable states. Ecology 87: 2862–2870.

Connell JH (1961) The influence of interspecific competition and other factors on the distribution of the barnacle Chthamalus stellatus. Ecology 42: 710–723.

Diamond JM (1975) The island dilemma: lessons of modern biogeographic studies for the design of nature reserves. Biological Conservation 7: 129–146.

Drolet D, Bringloe TT, Coffin MR, Barbeau MA and Hamilton DJ (2012) Potential for between‐mudflat movement and metapopulation dynamics in an intertidal burrowing amphipod. Marine Ecology Progress Series 449: 197–209.

Englund G and Cooper SD (2003) Scale effects and extrapolation in ecological experiments. Advances in Ecological Research 33: 161–213.

Fluharty D (2000) Habitat protection, ecological issues, and implementation of the Sustainable Fisheries Act. Ecological Applications 10: 325–337.

Gilpin ME and Hanski IA (1991) Metapopulation Dynamics – Empirical and Theoretical Investigations. London, UK: Academic Press.

Goss‐Custard JD (1980) Competition for food and interference among waders. Ardea 68: 31–52.

Grant J (1983) The relative magnitude of biological and physical sediment reworking in an intertidal community. Journal of Marine Research 41: 673–689.

Hanson CA, Fuhrman JA, Horner‐Devine MC and Martiny JBH (2012) Beyond biogeographic patterns: processes shaping the microbial landscape. Nature Reviews. Microbiology 10: 497–506.

Hanski I and Simberloff D (1997) The metapopulation approach; its history, conceptual domian and application to conservation. In: Hanski I and Gilpin ME (eds) Metapopulation Biology: Ecology, Genetics, and Evolution, pp. 5–26. San Deigo, CA: Academic Press.

Hassell MP (1978) Arthropod Predator–Prey Systems. Princeton, NJ: Princeton University Press.

Haury LR (1978) Patterns and processes in the time‐scales of plankton distributions. In: Steele J (ed) Spatial Pattern in Plankton Communities, pp. 277–327. New York, NY: Plenum Press.

Hewitt JE, Thrush SF, Dayton PK and Bonsdorf E (2007) The effect of spatial and temporal heterogeneity on the design and analysis of empirical studies of scale‐dependent systems. American Naturalist 169: 398–408.

Hines AH, Whitlatch RB, Thrush SF, et al. (1997) Nonlinear foraging response of a large marine predator to benthic prey: eagle ray pits and bivalves in a New Zealand sandflat. Journal of Experimental Marine Biology and Ecology 216: 211–228.

Hui C, Veldtman R and McGeoch MA (2010) Measures, perceptions and scaling patterns of aggregated species distributions. Ecography 33: 95–102.

van de Koppel J, Bouma TJ and Herman P (2012) The influence of local‐ and landscape‐scale processes on spatial self‐organization in estuarine ecosystems. Journal of Experimental Ecology 215: 962–967.

Kotliar NB and Weins JA (1990) Multiple scales of patchiness and patch structure: a hierarchical framework for the study of heterogeneity. Oikos 59: 253–260.

Leibold MA, Holyoak M, Moquet N, et al. (2004) The metacommunity concept: a framework for multi‐scale community ecology. Ecology Letters 7: 601–613.

Levins R (1969) Some demographic and genetic consequences of environmental heterogeneity for biological control. Bulletin of the Entomological Society of America 15: 237–240.

Loreau M, Mouquet N and Holt RD (2003) Meta‐ecosystems: a theoretical framework for a spatial ecosystem ecology. Ecology Letters 6: 673–679.

MacArthur RH and Wilson EO (1967) The Theory of Island Biogeography. Princeton, NJ: Princeton University Press.

Moilanen A (2005) Reserve selection using non‐linear species distribution models. American Naturalist 165: 695–706.

Murphy DD, Freas KE and Weiss SB (1990) An environment‐metapopulation approach to population viability analysis for a threatened invertebrate. Conservation Biology 4: 41–51.

Nachman G (2006) The effects of prey patchiness, predator aggregation, and mutual interference on the functional response of Phytoseiulus persimilis feeding on Tetranychus urticae (Acari: Phytoseiidae, Tetranychidae). Experimental and Applied Acarology 38: 87–111.

Nanninga GB and Berumen ML (2014) The role of individual variation in marine larval dispersal. Frontiers in Marine Science 1: 71.

O'Neill RV, DeAngelis DL, Waide JB and Allen TFH (1986) A Hierarchical Concept of Ecosystems. Princeton, NJ: Princeton University Press.

Olds AD, Connolly RM, Pitt KA, et al. (2016) Quantifying the conservation value of seascape connectivity: a global synthesis. Global Ecology and Biogeography 25: 3–15.

Pikitch EK, Santora C, Babcock EA, et al. (2004) Ecosystem‐based fishery management. Science 305: 346–347.

Polis GA, Power ME and Huxel GR (2004) Food Webs at the Landscape Level. Chicago, IL: University of Chicago Press.

Puckett BJ and Eggleston DB (2016) Metapopulation dynamics guide marine reserve design: importance of connectivity demographics and stock enhancement. Ecosphere 7: 6.

Rietkerk M and Van de Koppel J (2008) Regular pattern formation in real ecosystems. Trends in Ecology & Evolution 23: 169–175. DOI: 10.1016/j.tree.2007.10.013.

Sale PF, Hanski I and Kritzer JP (2006) The merging of metapopulation theory and marine ecology: establishing the historical context. In: Kritzer JP and Sale PF (eds) Marine Metapopulations, pp. 3–28. Amsterdam: Elsevier Academic Press.

Schneider DC (1991) Role of fluid dynamics in the ecology of marine birds. Oceanography and Marine Biology Annual Review 29: 487–521.

Smallegange IM, van der Meer J and Kurvers RHJM (2006) Disentangling interference competition from exploitative competition in a crab‐bivalve system using a novel experimental approach. Oikos 113: 157–167.

Snelgrove PVR, Thrush SF, Wall DH and Norkko A (2014) Real world biodiversity‐ecosystem functioning: a seafloor perspective. Trends in Ecology & Evolution 29: 398–405.

Soranno PA, Cheruvelil KS, Bissell EG, et al. (2014) Cross‐scale interactions: quantifying multi‐scaled cause‐effect relationships in macrosystems. Frontiers in Ecology and the Environment 12: 65–73.

Thomas CD and Kunin WE (1999) The spatial structure of populations. Journal of Animal Ecology 68: 647–657.

Thrush SF, Pridmore RD, Hewitt JE and Cummings VJ (1994) The importance of predators on a sandflat: interplay between seasonal changes in prey densities and predator effects. Marine Ecology Progress Series 107: 211–222.

Thrush SF, Hewitt JE, Cummings VJ, et al. (2000) The generality of field experiments: interactions between local and broad‐scale processes. Ecology 81: 399–415.

Thrush SF, Hewitt JE, Hickey CW and Kelly S (2008) Multiple stressor effects identified from species abundance distributions: interactions between urban contaminants and species habitat relationships. Journal of Experimental Marine Biology and Ecology 366: 160–168.

Thrush SF, Hewitt JE, Lohrer A and Chiaroni LD (2013) When small changes matter: the role of cross‐scale interactions between habitat and ecological connectivity in recovery. Ecological Applications 23: 226–238.

Wilson DS (1992) Complex interactions in metacommunities with implications for biodiversity and higher levels of selection. Ecology 73: 1984–2000.

Wu JC, Jelinski DE, Luck M and Tueller PT (2000) Multiscale analysis of landscape heterogeneity: scale variance and pattern metrics. Geographic Information Sciences 6: 6–19.

Further Reading

Belovsky GE, Botkin DB, Crowl TA, et al. (2004) Ten suggestions to strengthen the science of ecology. Bioscience 54: 345–348.

Dayton PK and Tegner MJ (1984) The importance of scale in community ecology: a kelp forest example with terrestrial analogs. In: Price PW, Slobodchikoff CN and Gaud WS (eds) A New Ecology: Novel Approaches to Interactive Systems, pp. 457–483. New York, NY: Wiley.

Hildrew AG, Giller PS and Raffaelli D (1994) Aquatic Ecology: Scale, Pattern and Processes. Oxford, UK: Blackwell Scientific.

Levin SA (1992) The problem of pattern and scale in ecology. Ecology 73: 1943–1967.

Contact Editor close
Submit a note to the editor about this article by filling in the form below.

* Required Field

How to Cite close
Hewitt, Judi E, Thrush, Simon F, and Lundquist, Carolyn(Jan 2017) Scale‐Dependence in Ecological Systems. In: eLS. John Wiley & Sons Ltd, Chichester. [doi: 10.1002/9780470015902.a0021903.pub2]