Spatial heterogeneity as a multiscale characteristic of zooplankton community

Zooplankton spatial heterogeneity has profound effects on understanding and modelling of zooplankton population dynamics and interactions with other planktonic compartments, and consequently, on the structure and function of planktonic ecosystems. On the one hand, zooplankton heterogeneity at spatial and temporal scales of ecological interest is an important focus of aquatic ecology research because of its implications in models of productivity, herbivory, nutrient cycling and trophic interactions in planktonic ecosystems. On the other hand, estimating zooplankton spatial variation at the scale of an ecosystem, is a powerful tool to achieve accurate sampling design. This review concentrates on the spatial heterogeneity of marine and freshwater zooplankton with respect to scale. First to be examined are the concept of spatial heterogeneity, the sampling and statistical methods used to estimate zooplankton heterogeneity, and the scales at which marine and freshwater zooplankton heterogeneity occurs. Then, the most important abiotic and biotic processes driving zooplankton heterogeneity over a range of spatial scales are presented and illustrated by studies conducted over large and fine scales in both oceans and lakes. Coupling between abiotic and biotic processes is finally discussed in the context of the multiple driving forces hypothesis.Studies of zooplankton spatial heterogeneity refer both to the quantification of the degree of heterogeneity (measured heterogeneity) and to the estimation of the heterogeneity resulting from the interactions between the organisms and their environment (functional heterogeneity) (Kolasa & Rollo, 1991). To resolve the problem of measuring zooplankton patchiness on a wide range of spatial scales, advanced technologies (acoustic devices, the Optical Plankton Counter (OPC), and video systems) have been developed and tested in marine and freshwater ecosystems. A comparison of their potential applications and limitations is presented. Furthermore, many statistical tools have been developed to estimate the degree of measured heterogeneity; the three types most commonly used are indices of spatial aggregation, variance: mean ratio, and spatial analysis methods. The variance partitioning method proposed by Borcard et al. (1992) is presented as a promising tool to assess zooplankton functional heterogeneity.Nested patchiness is a common feature of zooplankton communities and spatial heterogeneity occurs on a hierarchical continuum of scales in both marine and freshwater environments. Zooplankton patchiness is the product of many physical processes interacting with many biological processes. In marine systems, patterns of zooplankton patchiness at mega- to macro-scales are mostly linked to large advective vectorial processes whereas at coarse-, fine- and micro-scales, physical turbulence and migratory, reproductive and swarm behaviors act together to structure zooplankton distribution patterns. In freshwater environments, physical advective forces related to currents of various energy levels, and vertical stratification of lake interact with biological processes, especially with vertical migration, to structure zooplankton community over large to fine- and micro-scales. Henceforth, the zooplankton community must be perceived as a spatially well-structured and dynamic system that requires a combination of both abiotic and biotic explanatory factors for a better comprehension and more realistic and reliable predictions of its ecology.