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Feeding Ecology and Ecomorphology of Cichlid Assemblages in a Large Mesoamerican River Delta

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Feeding Ecology and Ecomorphology of Cichlid Assemblages in a Large Mesoamerican River Delta Environ Biol Fish (2018) 101:867–879 https://doi.org/10.1007/s10641-018-0743-1 Feeding ecology and ecomorphology of cichlid assemblages in a large Mesoamerican river delta Allison A. Pease & Manuel Mendoza-Carranza & Kirk O. Winemiller Received: 17 August 2017 /Accepted: 5 February 2018 /Published online: 13 February 2018 # Springer Science+Business Media B.V., part of Springer Nature 2018 Abstract Fish assemblages in tropical lowland rivers cichlid assemblages, perhaps due to availability of abun- are characterized by a high richness of species that feed dant resources, even in low-water conditions. Canonical on a diverse array of food resources. Although closely correspondence analysis revealed that greatest morpho- related species often have similar feeding ecology, spe- logical differences am7ong species involved functional cies within the family Cichlidae display a broad spec- traits directly associated with resource use. Relation- trum of trophic niches, and resource partitioning has ships between feeding ecology and morphology were been inferred from studies conducted in Neotropical similar to those described for other riverine cichlids. rivers. We investigated interspecific variation in food Strong ecomorphological relationships facilitate infer- resource use and its relationship to morphological vari- ences about the ecology of cichlid species, including ation among cichlid fishes within the Pantanos de Centla species that currently lack data from field studies. Biosphere Reserve, a coastal area encompassing the Knowledge of ecological relationships will be important delta of the Grijalva-Usumacinta River in Tabasco, for conservation in the Pantanos de Centla, an ecosys- Mexico. Most species consumed benthic crustaceans, tem of global significance for biodiversity and ecosys- aquatic insect larvae, and detritus, but some were more tem services. herbivorous, and one species was a specialized pisci- vore. Dietary niche overlap among species was higher Keywords Cichlidae . Morphological traits . Niche than expected for one assemblage, and similar to ran- overlap . Neotropical dom expectations for another, suggesting a lesser role for resource partitioning than has been shown for some Introduction A. A. Pease (*) Department of Natural Resources Management, Texas Tech In species-rich freshwater fish assemblages, co- University, Box 42125, Lubbock, TX 79409-2125, USA occurring species occupy a diverse suite of trophic e-mail: [email protected] niches, and some have been found to partition resources, presumably to reduce interspecific competition (Ross M. Mendoza-Carranza Sustainable Management of Basin and Coastal Zones Research 1986;WinemillerandPianka1990;Herderand Group, El Colegio de la Frontera Sur (ECOSUR) Unidad Freyhof 2006). Species within the same family are often Villahermosa, Km 15.5 s/n Carreterra a Reforma, Rancheria ecologically similar due to phylogenetic niche conser- Guineo 2a, 86280 Villahermosa, Tabasco, Mexico vatism (e.g., McNyset 2009), but closely related species K. O. Winemiller may occupy diverse trophic niches in regions where Department of Wildlife and Fisheries Sciences, Texas A&M ecological adaptive radiations have occurred (Schluter University, 2258 TAMU, College Station, TX 77843-2258, USA 2000). In the incredibly diverse cichlid communities of 868 Environ Biol Fish (2018) 101:867–879 Lake Malawi in Africa, Genner et al. (1999a, 1999b) Grijalva-Usumacinta River delta within the Pantanos found that cichlids coexisting along rocky shores de Centla Biosphere Reserve in Tabasco, México. We showed significant differences in food resource use, also sought to identify relationships between morpho- but many species had considerable dietary overlap. logical features and the feeding ecology of cichlids in They suggested that ecological segregation may not be these assemblages. We carried out the study during the necessary to support coexistence in these diverse cichlid dry season, a period when dietary niche partitioning has assemblages. In more dynamic fluvial habitats, environ- been shown for other fluvial cichlid assemblages (e.g., mental variation often causes shifts in resource avail- Winemiller 1991b; Jepsen et al. 1997). During low- ability (Grossman et al. 1998), and trophic segregation water periods, availability of food resources declines may occur during periods when fish densities are high and densities of most fish species increases, which in- while habitat and food availability are low (Winemiller creases the potential for competition. We also hypothe- 1990, 1991b; Winemiller and Kelso-Winemiller 2003). sized that morphological variation among cichlids Cichlids in the Neotropics consume a broad array of would involve traits directly related to trophic resource resources, from aquatic macrophytes to other fishes, use in a manner similar to ecomorphological relation- with some being trophic specialists while retaining an ships documented for other fluvial cichlids (e.g., ability to exploit a variety of food items (Lowe- Winemiller et al. 1995;Cochran-Biedermanand McConnell 1991; Winemiller et al. 1995; Montaña and Winemiller 2010; Montaña and Winemiller 2013). In- Winemiller 2013). Differences in dietary preferences are formation on the trophic ecology of cichlid fishes in the likely to be reflected in morphological traits related to Lower Grijalva-Usumacinta should enhance manage- foraging and consuming food. Prior studies of riverine ment of fisheries resources and conservation of this cichlids have revealed strong relationships between region’suniquebiodiversity. morphological traits and diet (e.g., Winemiller et al. 1995;López-Fernándezetal.2012;Montañaand Winemiller 2013). For example, Winemiller et al. Materials and methods (1995) found that across continents, riverine cichlids showed convergent relationships between traits such as Study sites and data collection gut length, gape size, and head length and the proportion of fishes, invertebrates, vegetation, and detritus The Grijalva and Usumacinta join together in Tabasco, consumed. Mexico to form the largest river of Mesoamerica and a Cichlids comprise a major component of the fresh- major center of fish diversity (Myers 1966; Bussing water fish fauna of Mesoamerica, and the evolution and 1985; Miller 2005). Approximately 115 fish species zoogeography of the group (subfamily Cichlinae) has from 31 families have been documented in the been studied extensively (e.g., Martin and Bermingham Grijalva-Usumacinta region, and an estimated 36% of 1998;Hulseyetal.2004; López-Fernández et al. 2013). these species are endemic (Miller 2005). Cichlids are the Centers of Mesoamerican cichlid diversity, such as the most species-rich family in this region, with many spe- Río Grijalva-Usumacinta region of southern México, cies occurring together in local assemblages (Rodiles- contain many coexisting cichlid species that display Hernández et al. 1999;Miller2005; Soria-Barreto and considerable morphological diversity that likely reflects Rodiles-Hernández 2008). niche diversification (Myers 1966; Bussing 1985; Pease Sampling locations were within the Pantanos de et al. 2012). Studies in other Mesoamerican regions Centla Biosphere Reserve (Fig. 1), a vast protected area (e.g., Winemiller et al. 1995;Cochran-Biedermanand within the Lower Grijalva-Usumacinta Basin which was Winemiller 2010) suggest that coexisting cichlid species designated by the RAMSAR convention of 1995 as a partition trophic niches. Relatively little ecological in- wetland of international significance. San Pedrito La- formation is available for cichlids of the Lower Grijalva- goon (18°20′36^N 92°33′50^W) is a permanent fresh- Usumacinta (Miller 2005) despite the high diversity and water lagoon with expansive beds of aquatic macro- importance of many species for small-scale fisheries phytes (chiefly Vallisneria americana) and shorelines (Mendoza-Carranza et al. 2013). dominated by reeds (Phragmites australis). Polo Stream In this study we examined diets, niche breadth, and (18°29′21^N92°38′23^W) is a slow-moving, niche overlap of coexisting cichlid species in the mangrove-dominated freshwater tributary of the Río Environ Biol Fish (2018) 101:867–879 869 Fig. 1 Map of the study area within Pantanos de Centla Biosphere Reserve, Tabasco, Mexico Grijalva below its confluence with the Río Usumacinta. (2005) and deposited in the fish collection at El Colegio Along the shorelines of the stream, structural cover is de la Frontera Sur in Villahermosa, Tabasco. provided by riparian plants (mangrove trees and terres- Given that the objective of this study was to evaluate trial grasses), and the sandy substrate is covered with interspecific patterns of morphological and dietary diver- coarse particulate organic matter. A detailed description sity within local species assemblages, juvenile size clas- of these study sites is given in Mendoza-Carranza et al. ses, which tend to show relatively low interspecific di- (2010) and Sepúlveda-Lozada et al. (2015). Intensive vergence in morphology and diet (i.e., small size classes surveys were carried out in February and March 2007 of all species consume mostly microcrustacea and other during the dry season (Espinal et al. 2007;Yáñez- small aquatic invertebrates), were excluded from analy- Arancibia et al. 2009). In Pantanos de Centla, the dry ses. Species that were rarely captured during surveys of season is characterized by low water levels (< 2 m in the mesohabitats (N <5)alsowereexcludedfromanaly- Polo Stream and <1 m in San Pedrito Lagoon),
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