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Feeding Strategies Differentiate Four Detritivorous Curimatids in the Amazon

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Feeding Strategies Differentiate Four Detritivorous Curimatids in the Amazon Web Ecol., 20, 133–141, 2020 https://doi.org/10.5194/we-20-133-2020 © Author(s) 2020. This work is distributed under the Creative Commons Attribution 4.0 License. Feeding strategies differentiate four detritivorous curimatids in the Amazon Thatyla Farago1, Gabriel Borba1, Sidineia Amadio1, Joicyeny Oliveira2, Geraldo Santos1, Adalberto Val1, and Efrem Ferreira1 1Brazilian National Institute for Research of the Amazon (INPA), Av. André Araújo, 2936, Petrópolis, 69067-375, Manaus, AM, Brazil 2Amazonas State University (UEA), Av. Djalma Batista, 2470, Pq Dez de Novembro, 69050-010, Manaus, AM, Brazil Correspondence: Thatyla Farago ([email protected]) Received: 13 June 2019 – Revised: 30 July 2020 – Accepted: 13 August 2020 – Published: 2 October 2020 Abstract. Differences in food intake and morphological attributes may facilitate the coexistence of detritivorous fish. The present study investigated the possible differences in the feeding strategies of four species of curimatids that inhabit the floodplain of the central Amazon. For this, we determined the diet, daily food cycle, and whether characteristics of the intestine were related to the length of the fish and the amount of detritus consumed. The detritivory was confirmed, and we observed a difference in the foraging time between species. We found differ- ences in the length and weight of the intestine and the relationship of these variables with the length of the fish and the amount of detritus consumed. Our study suggests that despite belonging to the same family and food group, curimatids have characteristics that allow them to consume the detritus in different ways. 1 Introduction by detritus (Pereira and Resende, 1998; Vaz et al., 1999; Aranguren, 2002; Giora and Fialho, 2003; Alvarenga et al., One of the ecological questions still unanswered about 2006; Mérona et al., 2008). aquatic environments concerns the factors that determine Detritus is a highly available food resource, not limited to species richness in detritivorous communities (Moore et al., seasonal and spatial factors, and due to its structural charac- 2004). How is it possible for the detritus to sustain a high teristics, it may have been in the environment for millennia quantity and diversity of fish in neotropical environments? (Moore et al., 2004). It is defined as partially decomposed In the Amazon basin, one of the most representative detri- organic matter from plant and animal tissues, in addition to tivore groups is from the Curimatidae family, accounting for microorganisms and minerals (Gneri and Angeluscu, 1951; approximately half of the total fish biomass in South Amer- Gerking, 1994; Moore et al., 2004; Santana et al., 2015; Far- ican rivers (Bowen, 1983; Lowe-McConnell, 1999; Vari and rel et al., 2018; Zimmer, 2019). However, the chemistry of Röpke, 2013). This family has approximately 70 species. water and the spatial and seasonal availability of sources The species of the genera Psectrogaster and Potamorhina that form detritus is what will define its final composition are the most abundant and widely distributed (Albert and (Bowen, 1983; Goulding et al., 1988). Reis, 2011; Vari and Röpke, 2013; Van der Sleen and Albert, The composition of the detritus (origin, quantity, and 2018). However, coexistence mechanisms are still poorly un- quality) can affect the feeding rate, population density, and derstood for this group. The species, in general, share the trophic niche of detritivores (Rossi et al., 2015). However, same environment and co-occur throughout the year, regard- niche overlap appears to be common for this food group. The less of the hydrological cycle (Batista et al., 1998; Correia fact that many species consume detritus and occupy the same et al., 2015; Röpke et al., 2016). Besides, studies of feed- environment, with an apparent lack of competition, seems to ing ecology have not encountered significant differences in be promoted precisely by the abundance of the total resource the diets of these species, which are invariably dominated Published by Copernicus Publications on behalf of the European Ecological Federation (EEF). 134 T. Farago et al.: Feeding strategies differentiate four detritivorous curimatids in the Amazon and its high availability (Gerking, 1994; Pianka, 2000; Sid- tiate the consumption of detritus by the four species and fa- lauskas, 2007). However, the coexistence of detritivores can cilitate their occurrence in the same ecological niche in Ama- also be favored by differences in the use of this resource. The zonian freshwater ecosystems. possible variety of detritus composition allows detritivores to specialize in food aggregates with different combinations of 2 Materials and methods substrates (Delariva and Agostinho, 2001; Oliveira and Isaac, 2013; Rossi et al. 2015; Bayley et al., 2018). In addition, dif- 2.1 Fish sampling ferences in detritus consumption can also occur in ontoge- netic development (Semaprochilodus spp., Winemiller and The curimatid species selected for the present study are the Jepsen, 2004; Sarotherodon mossambicus, Bowen, 1979), most abundant and frequently found in the Catalão region, during the reproductive period (Curimatella lepidura, Al- a seasonally flooded area on the opposite margin of the varenga et al., 2006), associated with foraging space or sea- Negro River from the city of Manaus (3◦080–3◦140 S and sonality (Loricarichthys platymetopon, Lopes et al., 2009; 59◦530–59◦580 W). The four species were Potamorhina alta- Hypostomus spp., Oliveira and Isaac, 2013; Prochilodus spp., mazonica (Cope, 1878), Potamorhina latior (Spix and Agas- Bowen, 1983), by competition (Hypostomus spp., Oliveira siz, 1829), Psectrogaster amazonica (Eigenmann and Eigen- and Isaac, 2013), or due to other physiological and/or eco- mann, 1889), and Psectrogaster rutiloides (Kner, 1858). We logical demands (Loricariidae, Lujan et al., 2011). obtained the specimens analyzed in the present study from Independent of these conditions, all detritivorous fish have the Catalão/INPA Project (INPA – CEUA Ethics Committee, adaptations in the digestive process to extract large amounts protocol 051/2015; IBAMA collecting license no. 52392-2). of nutrients, since the detritus provides less energy and pro- This project was initiated in 1999 and involves the collection tein than other types of food (Sazima and Caramaschi, 1989; of standardized monthly samples of specimens using a set of Bowen et al., 1995; Yossa-Pérdomo and Araújo-Lima, 1996; gill nets (mesh sizes ranging from 30 to 120 mm), which are Castro and Vari, 2004; German and Bitong, 2009; Faria and positioned in the water for 24 h, with the fish being removed Benedito, 2011). To enable a high rate of absorption and as- every 6 h (6, 12, 18, and 24 h). The specimens were taken similation of nutrients, the digestive tract of these species to the Laboratory of Fish Population Dynamics (LDPP) at is characterized by an extremely long intestine when com- INPA in Manaus for identification, biometrics, and biologi- pared to other feeding categories (Zihler, 1982; Smith, 1989; cal analyses. Kramer and Bryant 1995a; Karachle and Stergiou, 2010; Becker et al., 2010; Griffen and Mosblack, 2011; German 2.2 Diet and the daily feeding cycle et al., 2015). This characteristic, although general, can vary from 3 to 10 times the body size among detritivorous species We determined the composition of the diets of the specimens (Moraes et al., 1997; Giora and Fialho, 2003; Alvarenga et collected between June 2010 and July 2011 by two meth- al., 2006; German, 2009; Silva, 2016). ods: (i) the frequency of occurrence (the percentage of the The differences found in the digestive tract, the type of number of stomachs containing food that included the item) detritus consumed, and foraging location is what defines the and (ii) the relative volume, based on a visual estimate of feeding strategy of each species. And this set of factors is the percentage of the volume of each stomach taken up by closely aligned with feeding habits and phylogeny (Hidalgo the item (Hyslop, 1980). These values were multiplied by et al., 1999; Guisande et al., 2012). Comparisons of the feed- the estimated repletion of each stomach (0 %, 10 %, 25 %, ing strategies of closely related and sympatric species are still 50 %, 75 %, or 100 %) to correct for errors resulting from lacking, as is the case with curimatids. These species could the analysis of stomachs with different degrees of repletion present similar morphological characteristics, and they may (see Goulding et al., 1988; Ferreira, 1993). The values ob- nevertheless respond to the environment in different ways. tained by the two methods described above (frequency of They may present subtle shifts in certain traits, in partic- occurrence and relative volume) were used to calculate the ular those related to metabolism and digestive morphology alimentary index (IAi) proposed by Kawakami and Vazzoler (Ward-Campbell et al., 2005; Hilton et al., 2008; Mérona et (1980): IAi D Fi × Vi=6(Fi × Vi), where IAi is the food in- al., 2008; Wagner et al., 2009; Griffen and Mosblack, 2011; dex of item i, Fi the frequency of occurrence of item i, and Porreca et al., 2017). Thus, we investigated variations in the Vi the relative volume of item i. feeding strategy, specifically the relationship between mor- We used the data collected between January 2013 and phological attributes and intake of detritus, of four abundant June 2018 to determine the daily feeding cycle. The time at curimatid species in an aquatic environment in the central
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