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A review of planktivorous fishes: Their evolution, feeding behaviours, selectivities, and impacts

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Résumé

La vision classique des limnologistes fut de considérer les interactions entre les composants des écosystèmes lacustres comme un flux d'influence unidirectionnel des sels nutritifs vers le phytoplancton, le zooplancton, et finalement les poissons, par l'intermédiaire de processus de contrôle successivement physiques, chimiques, puis biologiques (Straškraba, 1967). L'effet exercé par les poissons planctophages sur les communautés zoo- et phytoplanctoniques ne fut reconnu qu'à partir des travaux de Hrbáček et al. (1961), Hrbáček (1962), Brooks & Dodson (1965), et Straškraba (1965). Ces auteurs montrèrent (1) que dans les étangs et lacs en présence de poissons planctophages prédateurs visuels, les communautés zooplanctoniques étaient composées d'espèces de plus petites tailles que celles présentes dans les milieux dépourvus de planctophages et, (2) que les communautés zooplanctoniques résultantes, composées d'espèces de petites tailles, influençaient les communautés phytoplanctoniques. Bien que la variabilité de la réponse du phytoplancton à la prédation par les poissons révèle l'importance d'autres facteurs (tels que la limitation en sels nutritifs et la compétition interspécifique des algues), ces travaux démontrèrent que les communautés zoo- et phytoplanctoniques pouvaient effectivement être affectées par l'alimentation sélective des poissons planctophages. Pendant les deux dernières décennies, de nombreux travaux en limnologie se sont concentrés sur cet impact radical des poissons sur les communautés planctoniques. La réponse directe des communautés zooplanctoniques à la prédation visuelle des poissons planctophages (appelés en anglais ‘particulate feeders’) a suscité un interérêt tout particulier, alors que les effets multiniveaux causés par les poissons planctophages filtreurs (prédation sur le zooplancton plus broutage du phytoplancton) ont été plus rarement abordés. Les objectifs de cette révision sont de documenter les inter-relations poissons-plancton, afin (1) d'obtenir des éléments d'appréciation de l'impact des poissons sur les communautés planctoniques, et (2) d'établir des modèles mécanistiques d'alimentation planctophage tenant compte du répertoire alimentaire et de la sélectivité du poisson, des répones adaptatives du plancton, et des conditions du milieu.

L'approche utilisée ici est basée sur des résultats expérimentaux de terrain et de laboratoire provenant de la littérature concernant les systèmes tropicaux et tempérés d'eau douce (parfois marine). Quatre groupes de poissons planctophages sont distingués: les prédateurs visuels limités par la taille de leur bouche (c'est-à-dire les larves et les espèces de petites tailles: ‘gape-limited predators’), les prédateurs visuels proprement dit (‘particulate feeders’), les filtreurs par pompage (‘pump filter feeders’), et les filtreurs par dĺacement (‘tow-net filter feeders’). Pour chaque groupe, les mécanismes de sélection des proies sont analysés, aussi bien du point de vue du prédateur que de la proie. Afin de rechercher les mécanismes déterminant la sélectivité alimentaire du prédateur et de discuter ses effets potentiels sur les commurlautés de proies, l'acte de prédation est décompóse en une séquence d'évènements successifs (Holling, 1966): la détection, la poursuite, la capture, la rétention et la digestion pour les prédateurs visuels; et la capture, la rétention et la digestion pour les filtreurs. Les avantages et les inconvénients de plusieurs mesures de sélectivité (appélées indices d'électivité), aussi bien que leur utilisation appropriée sont discutés. Les modèles de sélection de proies et les théories de recherche optimale sont analysés pour les différents modes d'alimentation des planctophages. Des modèles mécanistiques basés sur l'approche de Holling (loc. cit.) sont proposés pour chaque mode d'alimentation afin de déterminer les vulnerabilités différentielles des proies et l'amplitude optimale de la diète.

Cette révision concerne les domaines de l'écologie générale, de la limnologie, de l'aménagement des pêcheries (tel que, par exemple, l'utilisation des ressources planctoniques, le repeuplement, l'introduction, ou le maintien de populations naturelles de poissons), et du contrôle biologique des processes d'eutrophisation (approches par biomanipulation). Elle insiste sur le réel besoin de connaissances sur la sélectivité alimentaire et l'utilisation de la nourriture par les poissons planctophages. Elle permet de conclure que prédateurs et proies sont intimement et mutuellement adaptés. Aussi, dans la plupart des cas, il apparait peu approprié d'aborder la dynamique du plankton et la qualité des eaux sans tenir compte de l'estimation des pressions de prédation et de broutage exercées par les poissons planctophages.

Abstract

The classical approach of limnologists has been to consider the interactions between lake ecosystem components as an unidirectional flow of influence from nutrients to the phytoplankton, to the zooplankton, and finally to the fish, through successive controls by physical, chemical, and biological processes (Straškraba, 1967). The effect of planktivorous fishes on zooplankton and phytoplankton communities was not recognized until the studies of Hrbáček et al. (1961), Hrbáček (1962), Brooks & Dodson (1965) and Straškraba (1965). They showed that (1) in ponds and lakes in the presence of planktivorous fishes the zooplankton communities were composed of smaller bodied species than in those lacking planktivores, and (2) the resulting small-bodied zooplankton communities affected the phytoplankton communities. Although the variability of the phytoplankton response to fish predation showed the importance of other factors (such as nutrient limitation and interspecific competition of algae), these studies emphasized that zooplankton and phytoplankton communities can be affected by the feeding selectivity of planktivorous fishes. During the last two decades, many limnological studies have focused on this dramatic impact of fish on plankton communities. The direct response of zooplankton communities to visual fish predation (i.e. particulate feeding) has been of major interest, whereas the multilevel effects of filter-feeding fish (predation on zooplankton plus grazing on phytoplankton) have been neglected. The objectives of this review are to document fish-plankton interrelationships in order to (1) provide insights into the impact of fish on plankton communities, and (2) outline mechanistic models of planktivory according to the feeding repertory and the selectivity of the fish, the adaptive responses of the plankton, and the environmental conditions.

The approach adopted here is based on field and laboratory experimental results derived from the literature on tropical and temperate freshwater (occasionally marine) systems. Four types of planktivorous fish are distinguished: the gape-limited larvae and small fish species, the particulate feeders, the pump filter feeders, and the tow-net filter feeders. For each type of planktivore, the mechanisms of prey selection are analyzed from the point of view of both the predator and the prey. To investigate the main determinants of the predator feeding selectivity, and to discuss its potential effects on prey communities, the predation-act is divided into a sequence of successive events (Holling, 1966): detection, pursuit, capture, retention, and digestion for particulate feeders; and capture, retention, and digestion for filter feeders. The strengths and weaknesses of various measures of selectivity (i.e. electivity indices), as well as their appropriate usages are considered. Available prey selection models and optimal foraging theories are analyzed for the different planktivore feeding modes. Mechanistic models based on Holling's (loc. cit.) approach are proposed for each feeding mode to determine differential prey vulnerabilities and optimal diet breadth.

This review has application to several fields, including general ecology, limnology, fisheries management (for example, utilization of planktonic resources, stocking, introduction, or maintenance of natural fish populations), and biological control of the eutrophication processes (biomanipulation approaches). It emphasizes the real need for more knowledge of the feeding selectivity and food utilization of planktivores. It concludes that predator and prey are mutually adapted. Thus, in most cases, study of plankton dynamics and water quality should include the assessment of fish predation and grazing pressures.

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  1. Xavier Lazzaro

    Present address: Laboratorio de Limnologia, Centro de Recursos Hidricos e Ecologia Aplicada, Departamento de Hidráulica e Saneamento, Universidade de São Paulo, Av. Dr. Carlos Botelho, 1465, SP. 13560, São Carlos, Brazil

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  1. ORSTOM (Institut Français de Recherche Scientifique pour le Développement en Coopération), 213, rue La Fayette, 75480, Paris Cedex 10, France

    Xavier Lazzaro

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Lazzaro, X. A review of planktivorous fishes: Their evolution, feeding behaviours, selectivities, and impacts. Hydrobiologia 146, 97–167 (1987). https://doi.org/10.1007/BF00008764

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