Body size in freshwater planktonic crustaceans: An overview of extrinsic determinants and modifying influences of biotic interactions

  • Hart R
  • Bychek E
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In view of its fundamental and pervasive influences and impacts on organism physiology and ecology, body size is recognized as a key component of evolutionary fitness and serves as the cornerstone of a seminal contribution in freshwater zooplankton ecology-the Size Efficiency Hypothesis (SEH) of Brooks & Dodson (Science 150:28-35, 1965). While the roles and implications of body size in predation and competition-central tenets of the SEH-have been widely considered and reviewed, no broader integrated synthesis exists of the collective array of body size determinants and their implications in the ecology in crustacean zooplankton-a numerically and functionally dominant group of aquatic organisms. Focusing on planktonic Cladocera and Copepoda in inland waters, in particular, we provide a wide-ranging overview of the direct and/or indirect effects of environmental conditions, consumable resources and biotic interactions that independently and/or collectively influence the phenotypic expression of body size (particularly as length), both within and between species. Some indirect ultimate evolutionary consequences of body size are considered, and we identify some controversies and unresolved issues related to this biologically crucial trait. While by no means exhaustive, our overview reveals a complex nexus of extrinsic proximate abiotic and biotic factors and interactions that influence body size, the phenotypic expression of which in natural systems commonly reflects contrasting outcomes related to conflicting direct and/or indirect selective pressures. In general, however, body size (both inter- and intra specifically) declines with rising temperature and increases with rising food supply (depending on its quality), although both temperature and food supply exert contrary influences on particular taxa (or life history stages) under certain environmental circumstances. Predation undoubtedly has an overriding influence on body size selection. Depending on its mechanistic basis (visual, tactile or both in tandem), it selectively favours either small or large body size, both within (adults vs. juveniles) and between prey species, which are accordingly often 'size-trapped' between contrasting selective pressures, with consequent indirect effects. The bioenergetics of fundamental physiological processes undoubtedly set constraints on body size and serve as the primary determinant. However, within such constraints, the phenotypic expression of body size reflects its adaptive modification in response to the prevailing abiotic and biotic environment. As such, body size represents an emergent ecological property, reflecting the outcome of specific circumstances and conditions, which vary both temporally within and spatially between different ecosystems, and are accordingly context dependent. Nevertheless, underlying physiological advantages of larger size (within and between species) among crustacean zooplankters-lower mass-specific metabolic rates (although recently challenged), higher individual feeding rates (at least among cladocerans), potentially wider food size-ranges, better starvation tolerances, higher potential fecundity, etc.-collectively favour the selection of increased body size, as predicted by the SEH. Although competitive superiority of large size (measured in terms of minimal food requirements) has been confirmed experimentally, this cannot be generalized to natural conditions, where conflicting and temporally variable pressures apply, and contribute to generally mixed, and temporally variable body size compositions. Complex undrlying ecological interactions and influences ultimately determine the phenotypic expression of body size in directions consistent with fitness optimization under prevailing circumstances. Certain specific and general deficiencies in information are identified. In particular, the overwhelming emphasis on daphniid cladocerans as model study taxa in freshwater ecosystems has marginalized the acquisition of a comparably broad and penetrating understanding of specific features both of non-daphniid cladoceran and copepod life histories and body size selection. Among daphniid cladocerans, contemporary definitive understanding devolves largely from reductionist laboratory approaches. Holistic re-integration of these mechanistic findings into natural system circumstances presents a difficult challenge that is attracting increasingly attention. With regard to copepods, synthetic integration of the expansive marine knowledge base appears crucial to inform and direct future investigations on freshwater taxa. The question of intrinsic body size regulation in copepods and cladocerans, especially in regard to final phenotypic plasticity in body size expression, awaits resolution. Overall, body size remains a multi-facetted and complex topic, offering promising challenges for further investigation.

Author-supplied keywords

  • Body size
  • Cladocera
  • Copepoda
  • Ecophysiological bioenergetics
  • Extrinsic determinants
  • Intrinsic controls

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  • R. C. Hart

  • E. A. Bychek

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