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Dietary´–Morphological Relationships in a Fish Assemblage of the Bolivian Amazonian Floodplain

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Dietary´–Morphological Relationships in a Fish Assemblage of the Bolivian Amazonian Floodplain Journal of Fish Biology (2003) 62, 1137´–1158 doi:10.1046/j.1095-8649.2003.00108.x,availableonlineathttp://www.blackwell-synergy.com Dietary´–morphological relationships in a fish assemblage of the Bolivian Amazonian floodplain M. POUILLY*†‡, F. LINO†, J.-G. B RETENOUX* AND C. ROSALES† *Institut de Recherche pour le De´veloppement (IRD), Universite´ Lyon 1, Laboratoire d’Ecologie des Hydrosyste`mes Fluviaux, 43, Bd du 11 Novembre 1918, F-69622 Villeurbanne Cedex, France and †Instituto de Ecologı´a, Universidad Mayor de San Andre´s, Bolivia, CP 10077, La Paz, Bolivia (Received 24 January 2002, Accepted 7 April 2003) Morphological correlates of diet were examined in 48 species of freshwater fishes from floodplain lakes in the central part of the Mamore´River (Bolivian Amazon). The species were classified, according to the percentage occurrence of seven food items, into eight broad trophic categories: mud feeders, algivores, herbivores, terrestrial invertivores and omnivores, carnivores, zooplanktivores, aquatic invertivores and piscivores. There were significant rela- tionships between the diet and morphology of the fishes even when the effect of taxonomical relatedness between species was eliminated. Relative gut length was the main morphological variable used to order species on a carnivore to mud feeder gradient. Standard length and head and mouth size were the morphological variables most closely associated with prey size. Mud feeder, algivore and piscivore species appeared as the most dietary and morphologically specialized. These results support both the hypotheses that species morphology influences the diet and that morphological similarity is conserved even in comparison with taxonomically unrelated species. # 2003 The Fisheries Society of the British Isles Key words: Bolivia; diet; ecomorphology; river floodplain; tropical freshwater fishes. INTRODUCTION A correlation between morphological attributes and diet has been sought in testing ecomorphological hypotheses in fish assemblages. The results of studies on diet and morphological relationships in temperate and tropical river fish assemblages, however, have excluded the idea of a general pattern. Some studies have revealed strong relationships between diet and morphology (Gatz, 1979; Wikramanayake, 1990; Winemiller et al., 1995; Piet, 1998; Hugueny & Pouilly, 1999; Xie et al., 2001) while others found relatively weak and indistinct relation- ships (Grossman, 1986; Douglas & Matthews, 1992; Motta et al., 1995; Winemiller & Adite, 1997). Some relationships between morphological attributes and diet have been encountered repeatedly, e.g. gut length is frequently positively ‡Author to whom correspondence should be addressed. Tel.: þ33 4 72446299; fax: þ33 4 72431141; email: [email protected] 1137 # 2003 The Fisheries Society of the British Isles 1138 M. POUILLY ET AL. correlated with herbivory, gill raker length is often associated with the con- sumption of plankton and fish size (and related head and mouth size) is com- monly associated with prey size. Relationships also exist between morphological attributes and feeding behaviour, e.g. the orientation of the mouth and the position of the eyes are linked with the position of the fish relative to its food (Gatz, 1979). At present, even though many studies have reported strong relationships between species ecology and morphology, no powerful predictive model exists for fish assemblages. This discrepancy may be partially due to the importance of fish behaviour in determining the type of prey that can be used (Grossman, 1986), the relatively high level of opportunism in freshwater fish species (Hugueny & Pouilly, 1999), the influence of phylogeny on limiting morpho- logical adaptation (Douglas & Matthews, 1992; Motta et al., 1995) and the selection of morphological and dietary variables. Ecomorphological hypotheses include two main considerations: (1) species morphology is likely to be similar within an ecological group and to differ between ecological groups depending on the nature of the resource used and the strategy developed by the species to use it and (2) the morphological variations correspond to a response to selective pressure and result in the phenomenon of convergence: ‘morphological similarity of phylogenetically unrelated species’ (Winemiller, 1991). The present study investigated the trophic ecology and morphology of 48 dominant fish species inhabiting lakes of the floodplain of the Mamore´ River, one of the principal drainages of the Bolivian Amazon. MATERIALS AND METHODS STUDY AREA The Bolivian Amazonian plain is situated in the upper watershed of the Madera River, one of the primary tributaries of the Amazon River (Fig. 1). The majority of this territory (c. 550 000 km2) comprises the ‘Llanos de los Moxos’ (province of Beni), a landscape dominated by savannah with some patches of forest confined to the higher part of the plain, and forest galleries (bands of vegetation) along the rivers. The Mamore´River drains the south Bolivian Andes and >85% of the Moxos savannah area, and corres- ponds to a large floodplain system with a potential flood extension of c. 150 000 km2 (Denevan, 1980). Local climatic conditions are marked by the alternation of a wet season (October to March) and a dry season (April to September). A substantial annual flood (maximum flow in the main channel in February 1987 was 8010 m3 sÀ1) generally occurs at the end of the wet season (December to April) and can last as long as 3 or 4 months (Loubens et al., 1992). The study area is situated in the central part of the Mamore´River (between 14300 and 14520 S and 64510 and 65010 W) near the city of Trinidad (capital of Beni province). Loubens et al. (1992) described the aquatic habitats of the Mamore´River floodplain. The study site included eight lakes in the central Mamore´River floodplain that corresponded to four different ecological habitats: oxbow lakes at three locations (near the river, in the middle of the forest ‘band’ and at the limit between forest and savannah) and savannah lakes. Pouilly et al. (1999) presented a preliminary description of limnological para- meters, plankton and fish communities. # 2003 The Fisheries Society of the British Isles, Journal of Fish Biology 2003, 62, 1137´–1158 DIET AND MORPHOLOGY OF AMAZONIAN FISHES 1139 82° 78° 74° 70° 66° 62° 58° 54° 50° 12° 8° 4° 0° Amazon Basin 4° 8° Rio Madera 12° Beni Study area Rio Mamoré 16° 0 500 1000 km Bolivia 20° Projection Lambert’s Azimuthal FIG. 1. Map of the Amazon watershed showing the Bolivian part of the upper Madera basin (Beni plain and Andes) and the study area in the central Mamore´River. FISH SAMPLING Fishes were sampled using 13 gillnets (25 m long and 2 m depth) with a wide range of mesh sizes (10, 15, 20, 25, 30, 35, 40, 50, 60, 70, 80, 90 and 110 mm). Sampling was conducted during eight periods between March 1998 and March 2000. For each sampling (lake and period), gillnets were left in place for 2 h in the evening (1700´–1900 hours) and for 2 h in the morning (0500´–0700 hours). They were placed near the shore and their locations were approximately the same throughout the study. Captured fishes were preserved in buffered formaldehyde (4%), transported to the laboratory and then placed in buffered alcohol (75%). They were identified at the species level (or at the genus level if systematic knowledge was inadequate). Identification was based on voucher specimens left by a previous systematic research programme (Lauzanne & Loubens, 1985; Lauzanne et al., 1991) at the Trinidad fish collection (CIRA-UTB), the Museo Nacional de Historia Natural, La Paz and at the Muse´e National d’Histoire Naturelle, Paris. The 48 species analysed represented 89% of the total number of individuals captured during the study (M. Pouilly, unpubl. data). Species were selected in order to present a broad range of systematic groups. The 48 species studied belonged to five main orders (Characiformes, Siluriformes, Gymnotiformes, Perciformes and Clupeiformes) and represented 18 families out of the 26 registered during the study (six families of # 2003 The Fisheries Society of the British Isles, Journal of Fish Biology 2003, 62, 1137´–1158 1140 M. POUILLY ET AL. Characiformes out of eight collected; seven families of Siluriformes out of nine; two families of Gymnotiformes out of five; one family of Perciformes out of two; two families of Clupeiformes out of two collected). Changes in fish diet and morphology depend on the stage of development and the size of the fish (Me´rigoux & Ponton, 1998). Consequently, for each species, a range of sizes that were assumed to correspond to the adult stage was studied. All the fishes included in the morphological analysis were also used for the analysis of diet. In order to optimize the estimation of diet in a given species, however, extra fishes captured were included in the diet analysis. FISH DIET ANALYSIS Estimation of diet was based on the analysis of stomach contents. After identification of the fish, the stomach was extracted by dissection. Empty stomachs or stomachs with almost fully digested contents were eliminated. The contents of the remaining stomachs were examined under a microscope and items were separated into seven categories: soft substratum (MUD); algae or periphyton (ALG); aquatic or terrestrial vegetation, fruits or seeds (VEG); zooplankton (cladocerans, rotifers or copepods, ZOO); aquatic inverte- brates (AIN); terrestrial invertebrates (TIN); fishes (FISH). The invertebrate categories (terrestrial and aquatic) mainly
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