Lake trophic food webs: structure, function, interactions and spatio-temporal variations
Comparative analysis of lake food webs is a focal point of research in contemporary limnology and lake management. The study of ecological processes determining foodweb structure and function lead to the emergence of constrasting hypotheses and intense debates on the relative role of nutrients and food web structure in regulating temperate lake ecosystems. In contrast, studies in tropical lakes are in general descriptive and the search for integrate concepts and models is yet in development. This review paper presents an overview and a critical analysis of actual knowledge, methodological approaches, regulation models and controversies on foodweb structure and function in temperate and tropical lakes. Our synthesis suggests that the apparent diversity in models of lake foodwebs could reflect a gradient (or contiuum) of intermediate foodweb situations, regulated by various environmental factors. The differences among lakes could be related to three main biotic factors, independently of the climatic, geographical and trophic conditions: 1. the important cascading effect of strictly piscivorous fish in temperate lakes compared to the weak cascades induced by opportunistic omnivorous fish in tropical lakes, 2. the primacy of omnivory and opportunistic feeding behaviour of tropical fish, 3. the key role of herbivorous macrozooplankton (cladocerans, mostly Daphnia spp.) in temperate lakes where they are both selective preys of planktivorous fish and efficient grazers of nanophytoplankton, and 4. the synchronous reproduction of fish with seasonal plankton succession in temperate lakes, compared to continuous reproduction of fish and lack of seasonal coupling in tropical lakes. Consequently, food webs regulation ranges along a gradient of situations with two extreme models: 1. a model with intense cascading (top-down) regulation and attenuation of bottom-up effects typical of oligo-mesotrophic temperate lakes, characterized by the dominance of piscivorous fish and large herbivorous zooplankton (Daphnia spp.), and 2. a model with intermediate regulation encountered in eutrophic temperate lakes and most of tropical lakes, characterized par the dominance of filter omnivorous fish and small size zooplankton. Our synthesis also emphasizes the importance of coupling experimental approches in mesocosms or whole-lake biomanipulation with long-term monitoring and modelisation to fully understand and predict the functionning of lake ecosystems over different spatial and temporal scales