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Comparison of Fish Assemblages in Flooded Forest versus Floating Meadows Habitats of an Upper Amazon Floodplain (Pacaya ...


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COMPARISON OF FISH ASSEMBLAGES IN FLOODED FOREST VERSUS FLOATING MEADOWS HABITATS OF AN UPPER AMAZON FLOODPLAIN (PACAYA SAMIRIA NATIONAL RESERVE, PERU) By SANDRA BIBIANA CORREA A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2005

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Copyright 2005 by Sandra Bibiana Correa

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To my parents and my family. During the course of my life their love and belief in my potential encouraged me to keep going and achieve my dreams. They supported me in all of my decisions even when they separated us for long periods of time. I also dedicate this thesis to Michael Goulding who inspired me to study Amazonian fishes and who is a pioneer in showing their beauty and fragility to the world.

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ACKNOWLEDGMENTS This thesis would not have been possible without the support and commitment of my supervisory committee chair (Dr. James S. Albert) and my committee members (Dr. Lauren J. Chapman and Dr. William G.R. Crampton). All of them put lots of energy and time into the development of different aspects of this work. I thank James and Will for the outstanding time we spent in Peru during the field work that was the basis for this thesis. They were also crucial during the writing stage, and I enjoyed great discussions on Amazonian fish natural history and ecology. Lauren was my main support during the data analysis process. Without her, many of the results presented in this thesis may not have come out. I also want to thank my field assistant; B.Sc. Mario Escobedo, who worked on this project as if it was his own; and with great dedication, made it possible to meet the goal of 18 hours of continuous sampling per day. I also thank the fishermen who collaborated on the sampling: Hitler Rodriguez, Wilson Lanza, and Marco from the village of Bretaa. I thank the Instituto Nacional de Recursos Naturales (INRENA) for the research permission (under J.S. Albert). I thank the graduate students of the Zoology Department at University of Florida (UF) for great interactions. I thank Ann Taylor from the Editorial Office at UF. Finally I want to thank N. Bynum, P. Coley, R. Prendeville, the Grunwald-Seibel family, and C. Chapman for the valuable support that finally put me in graduate school. iv

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Last, I would like to acknowledge the support of the National Science Foundation (NSF-DEB 0215388, PI: J.S. Albert) which founded Project Ucamara and made possible the expedition for this thesis. This work was also supported with equipment donated by Idea Wild. I would also like to thank Fish Base, for the incredible resource that this data base represents to ichthyologists from all around the world. v

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TABLE OF CONTENTS page ACKNOWLEDGMENTS .................................................................................................iv LIST OF TABLES ............................................................................................................vii LIST OF FIGURES .........................................................................................................viii ABSTRACT .......................................................................................................................ix INTRODUCTION ...............................................................................................................1 Characterization of Floodplains in the Amazon Basin .................................................1 Importance of Flooded Forest and Floating Meadows for Fishes ................................3 Previous Studies on Amazonian Floodplain Fishes ......................................................3 Specific Objectives .......................................................................................................6 MATERIALS AND METHODS .........................................................................................7 RESULTS ..........................................................................................................................16 DISCUSSION ....................................................................................................................37 Species Richness and Abundance ...............................................................................37 Biomass and Size Distributions ..................................................................................39 Diel Variation in Species Richness, Abundance and Biomass ...................................41 Fish Assemblages in Floodplains ...............................................................................42 Sampling Limitations ..................................................................................................46 Conclusion ..................................................................................................................47 APPENDIX: SYSTEMATIC LIST OF SPECIES...........................................................48 LIST OF REFERENCES ...................................................................................................51 BIOGRAPHICAL SKETCH .............................................................................................59 vi

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LIST OF TABLES Table page 1 Abundance of 78 species in each of 10 locations in flooded forest and floating meadows of the Cao Yarina, Pacaya Samiria National Reserve. ...........................21 2 Mean values of different assemblage characteristics for flooded forest and floating meadows .....................................................................................................25 3 Comparison of day and night captures between flooded forest and floating meadows ...................................................................................................................26 4 Principal food items for 78 species and their relative abundance as a percentage of the total abundance (n = 2793 individuals) in flooded forest and floating meadows, during May 2003 at Cao Yarina, Pacaya Samiria National Reserve ....27 5 Systematic list of species organized by order, family, genus and species based on gill-netting sampling of the Cao Yarina floodplain, Pacaya Samiria National Reserve, during May 2003 ........................................................................48 vii

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LIST OF FIGURES Figure page 1 Map of the study area in the Upper Amazon, Pacaya Samiria National Reserve, Peru...........................................................................................................................14 2 Sampling place in Location 1...................................................................................15 3 Species richness saturation curves for flooded forest and floating meadows samples.....................................................................................................................30 4 Rank-ordered abundances (absolute number of individuals per species) of 78 species of fish captured during May 2003 in Cao Yarina, Pacaya Samiria National Reserve, Peru.............................................................................................31 5 Abundance per habitat of the 10 most abundant species (absolute number of individuals per habitat) captured during May 2003 in Cao Yarina, Pacaya Samiria National Reserve, Peru...............................................................................32 6 Variation in abundance, biomass, and richness during three periods of capture (EN, LN, LD) during May 2003 in Cao Yarina, Pacaya Samiria National Reserve, Peru............................................................................................................33 7 Changes in abundance of the five most common species among three sampling periods in flooded forest and floating meadows......................................................34 8 Size frequency distributions of fishes captured per habitat during May 2003 in Cao Yarina, Pacaya Samiria National Reserve, Peru.............................................35 9 Nonmetric scaling ordination (NMS) of sampling locations in species space based on the abundances of 57 species that occurred in more than 5% of the sampling locations during May 2003 in Cao Yarina, Pacaya Samiria National Reserve, Peru............................................................................................................36 viii

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Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science COMPARISON OF FISH ASSEMBLAGES IN FLOODED FOREST VERSUS FLOATING MEADOWS HABITATS OF AN UPPER AMAZON FLOODPLAIN (PACAYA SAMIRIA NATIONAL RESERVE, PERU) By Sandra Bibiana Correa May 2005 Chair: James S. Albert Major Department: Zoology Flooded forests and floating meadows of Amazonian floodplains are important habitats for fishes and yet the distribution and abundance of fish in these habitats are poorly known. This study presents the first quantitative comparison of fish species richness and composition, abundance, biomass and body-size distributions between flooded forests and floating meadows in an Amazon floodplain. This study was conducted in the floodplain of Reserva Nacional Pacaya Samiria, in the Peruvian Amazon and sampled fish assemblages in both flooded forests and floating meadows, using matched sets of gill nets of different mesh sizes. This represents the only reliable method available of doing a quantitative assessment of fish assemblages in flooded forest. Species richness was higher in flooded forest because of to a high percentage of unique species. Species abundances followed a hollow distribution in which three species accounted for 70 and 60 % of the total abundance in flooded forest and floating ix

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meadows, respectively. Diel variation in the abundance and richness of fishes was observed in both habitats. Body-size distributions were very similar in the two habitats (in the range of 70 to 400 mm). In both flooded forests and floating meadows, most of the fishes caught were invertebrate feeders and frugivores (i.e., Dianema longibarbis and Triportheus angulatus) in flooded forest; and detritivores (i.e., Psectrogaster rutiloides) in floating meadows. Fish predators and frugivores were represented by few species. Based on species composition and abundance, multivariate analysis suggested the presence of two subtly different fish assemblages in flooded forest vs. floating meadows. Water depth accounts for part of the differentiation in fish assemblages. However, the fact that many species (including all the common ones) were shared between the two habitats suggests movement of species between habitats in the floating meadows of Cao Yarina during the flood season. x

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INTRODUCTION Amazon fishes are strongly influenced by the inundation pulse (Goulding, 1980; Goulding et al. 1988; Junk, 1997). In the Amazon basin, floodplains are periodically inundated by the overflow of rivers and lakes, or precipitation (Junk, 1997). Floodplains offer a wide variety of resources for both plants and animals, and a complex nutrient and energy cycle integrates the aquatic and terrestrial phases (Junk, 1997). The powerful dynamics driven by the flood pulse make floodplains alternately suitable for aquatic and terrestrial organisms. Therefore, floodplains are believed to have a principal role in maintaining high biodiversity of both aquatic and terrestrial organisms (Goulding, 1980; Junk, 1997; Gopal & Junk, 2000). Furthermore, the Amazon basin contains the most diverse fish fauna (3,000 species) in the world (Val & Almeida-Val, 1995), which accounts for about 10 % of the global fish fauna (Groombridge & Jenkins, 1998). Besides the importance of floodplains in maintaining diversity, floodplains provide much of the carbon consumed by several commercially important species (Araujo-Lima et at. 1986a; Benedito-Cecilio et al., 2000). And fishes represent the main sources of protein and commercial income to riverine people (Bayley, 1989; Crampton et al., 2004). Characterization of Floodplains in the Amazon Basin Floodplains are mosaics of lakes, channels (parans), and levees (restingas), where flooded forest and floating meadows (patches of aquatic macrophytes) are the principal vegetation formations (Junk, 1984). Flooded forests are seasonally inundated by either nutrient-rich rivers (called vrzea in Brazil), or by nutrient-poor rivers (called igap in 1

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2 Brazil) (Prance, 1980). Nutrient-rich rivers, originate in the Andes (Goulding et at. 2003). They have a high concentration of suspended solids, with a near-neutral pH (6 to 7 pH), and high electric conductivity (50 to 300 S cm -1 ) (Junk, 1984). Nutrient-poor rivers, originating in the Amazonian lowlands, have relatively high transparency, with water color depending on the soils that they drain, and the concentration of organic compounds in the water. The characteristic dark tea color of some waters is usually the result of local podzol-based soil geochemistry. Here there is little sequestering of organic matters, resulting in waters with a high concentration of humic substances (Leenheer, 1980). In this type of water, acidity varies between 2 and 5 pH, transparency ranges from 1.30 to 2.90 m (Sioli, 1984), and electric conductivity is low (5 to 30 S cm -1 ). In latosoils, the sequester of humic compounds on the clay matrix results in highly transparent drainage water (Leenheer, 1980) ranging from 1.10 to 4.30 cm (Sioli, 1984). Additionally, rivers that drain ancient upland regions of granite shields (i.e., Brazilian and Guiana Shields), where there is little erosion, are highly transparent (Goulding et at. 2003). Chemical composition of each water type, nutrient levels, and the length of the flood season determine the vegetation present in each forest type (Prance, 1979). Biomass and local tree-species diversity (-diversity) is higher in vrzea than in igap; however at a regional scale (-diversity), igap forests are more diverse (Kubitzki, 1989). Flooded meadows are extensive rafts of floating macrophytes growing along the margins of rivers, lakes, and channels. The specific composition of these meadows changes from area to area, depending on water fluctuation. However, the most diverse macrophyte communities are present in floodplains of nutrient-rich rivers (Junk & Howard-Williams, 1984).

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3 Importance of Flooded Forest and Floating Meadows for Fishes During the flood season, flooded forest and floating meadows are the most important habitats for a variety of Amazonian fish species. Flooded forests offer a wide variety of allochthonous food resources (e.g., terrestrial herbaceous plants, leaves, flowers, seeds and fruits, and terrestrial invertebrates) (Goulding, 1980), while floating meadows offer autochthonous food resources (e.g., phytoplankton, periphyton, aquatic herbaceous plants, and aquatic invertebrates) (Junk et al., 1997). How fish species partition resources and microhabitats in floodplains is only partially known, partly because of the paucity of quantitative studies of fish assemblages in flooded forests. In general, small fish species and juveniles are the main components of the fish assemblage occupying floating meadows because of the availability of shelter and the abundance of food among meadows roots (Saint-Paul & Bayley, 1979; Goulding, 1980; Goulding & Carvalho, 1982; Junk, 1984b; Sanchz-Botero & AraujoLima, 2001; Carvalho & Araujo-Lima, 2004). When fishes become larger, they move toward the flooded forest or into the main river channel (Junk, 1997). Previous Studies on Amazonian Floodplain Fishes Given the extremely high diversity of fishes contained in the Amazon basin, one of the most striking questions in the study of Amazonian floodplains is the origin and maintenance of species diversity. Henderson et al. (1998) summarized historical and contemporary factors that aid in resolving this question in vrzea floodplains. They speculated that the floodplain fish fauna is characterized by species with wide distributions and by low endemism. The spatial and temporal interconnection of habitats, the ephemeral character of habitats at large temporal scale, and the obligate migration during dry season, act as a group in limiting opportunities for speciation within

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4 floodplains. They suggested that speciation may be occurring in river headwaters, with subsequent colonization of whitewater floodplains. Moreover, floodplain habitats would be selected for attributes of colonizing species resulting in more-simple body trends, and wide phenotypic plasticity. Other studies on Amazonian floodplain fishes, range from life history of economically key species (e.g., Arapaima gigas (Hurtado, 1999), Colossoma macropomum (Goulding & Carvalho, 1982), Cichla sp. (Cala et al. 1996), and pimelodid catfishes (Arboleda, 1988; Rodriguez, 1991; Celis, 1994; Agudelo, 1994)) to community ecology (see below), ecophysiology (Junk et al. 1983; Val et at. 1986; Val & Almeida-Val, 1995), trophic ecology and nutrients flux (Araujo-Lima et al. 1986a; Forsberg et al. 1993; Yossa & Araujo-Lima, 1998; Benedito-Cecilio et al. 2000; Leite et al. 2002), migration (Vazzoler & Menezes, 1992; Barthem & Goulding, 1997), larval movement and recruitment (Araujo-Lima et at. 1994; Araujo-Lima & Oliveira, 1998; Carvalho & Araujo-Lima, 2004), and fisheries (Bayley & Petrere, 1989; Bayley, 1995; Bayley, 1996; Merona, 1990; Almeida et al. 2001; Crampton et al. 2004). Most of the studies on community ecology of Amazonian fishes have focused on associations between floating meadows and fishes (Bayley, 1983; Soares et al., 1986; Araujo-Lima et al., 1986b; Henderson & Hamilton, 1995; Crampton, 1996; Henderson & Crampton, 1997; Henderson & Robertson, 1999; Petry et al., 2003). In contrast, there are few studies on fishes from flooded forests. The most comprehensive study was Goulding (1980), who made a detailed account of feeding behavior (emphasizing the importance of seeds and fruits in fish diet). Later, Goulding et al. (1988) made the most-intensive survey of fish species in an Amazonian affluent (i.e., Rio Negro). They did a

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5 nonsystematic sampling of the Rio Negros flooded forest and provided a list of 184 species, and food habits of 140 species in this habitat. Henderson and Crampton (1997) conducted a comparative study of fish richness and relative density in nutrient-poor and richer floodplain habitats in the Tef region, Brazil. They sampled both floating meadows and flooded forests, at dry and flood season, and presented data on species abundance, distribution, and biomass. However, the fact that they used a different sampling technique at each habitat (seine nets in meadows, gill nets in flooded forest) did not allow direct quantitative comparisons of abundance and standing crop. Probably the most complete study of community ecology of fishes in flooded forests was conducted by Saint Paul et al. (2000). They compared fish assemblages in nutrient-poor and richer flooded forests near Manaus, Brazil, by using a wide range of gill-net mesh sizes. They reported 238 species; and contrasted species diversity, distribution, abundance, and biomass between the two habitats. In all of the studies mentioned above, gill nets were used as the only available method for quantitative sampling in flooded forests. Although fish length is related to net mesh size (Jensen, 1990) and gill-net effectiveness varies with fish behavior (Jensen, 1986), the structure of flooded forests does not allow the use of other sampling techniques. The only means to improve the efficiency of gill nets is to use a combination of nets of different mesh sizes (Jensen, 1990); usually arranged in batteries, with the largest mesh sizes downstream (if there is any flow). Data on biomass and production of Amazonian fishes are almost nonexistent (Saint-Paul et at. 2000). There is an estimate of annual fish production of 31.2 g m -2 in marginal floating meadows, and 19.2 g m -2 in drifting islands of meadows. The study was conducted in Lake Mamirau (Brazil), a nutrient-rich lake, during 8 days at the beginning

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6 of the flood season (Henderson & Hamilton, 1995). Another study in the same lake estimated a mean of 13.5 g fish m -2 based on 12 days of sampling during the flood season (Henderson & Crampton, 1997). Both of these estimates were based on seine-net sampling. In contrast, for flood forest there is an estimate of 33 g fish m -2 (of net area) per day for a nutrient-poor lake (Lago do Prato, Anavilhanas, Brazil), and 104 g fish m -2 (of net area) per day for a nutrient-rich lake (Lago do Inacio, Rio Manacapuru, Brazil) (Saint-Paul et at. 2000). These estimates were based on 48 hours of sampling (over 2 years) during the flood season, and were based on gill-net sampling; therefore the authors standardized by capture per unit effort (CPUE). The fact that all of the estimators mentioned above were obtained with different sampling protocols makes them noncomparable (e.g., seine nets give an estimator of biomass per m 2 of meadow, whereas gill nets give an estimator of biomass per m 2 of net surface area). Moreover, fish abundance and biomass differ from season to season in the same area (Henderson & Hamilton, 1995; Henderson & Crampton, 1997; Saint-Paul et at. 2000). Thus, to adequately compare fish biomass between habitats, sampling must be conducted during the same season and using the same fishing techniques. Specific Objectives Given the apparent importance of flooded forest in maintaining fish diversity in Amazonian floodplains, and the lack of knowledge on how fish species partition resources and microhabitats in floodplains, the present study used a standardized sampling technique to compare species richness and composition, abundance, biomass, and size distribution of fishes in flooded forest with those of floating meadows in a floodplain of the Peruvian Amazon.

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MATERIALS AND METHODS Study site. This study was conducted in the Cao Yarina, a tributary of the Ro Pacaya in the Pacaya Samiria National Reserve (PSNR). This reserve is located at the confluence of the Maraon and Ucayali Rivers, in the Peruvian Amazon (5 20.575 S; 74 30.117 W). All specimens where collected within 10 km from one of the guard posts in the reserve (Puesto de Vigilancia 2 PV2, Fig. 1). The PSNR is the largest continuous area of protected vrzea floodplain in the Amazon basin (21,500 km 2 ) (INRENA, 2000) and therefore provides a good opportunity to survey fish communities that have not been highly perturbed as is the case in most of the lower and central Amazonian floodplains (with exceptions such as the Mamirau Reserve, Brazil). The PSNR is located within the Ucamara Depression, an active deposit of marine and continental sediments dating back from the Late Tertiary to the present (Bayley et al., 1992). Consequently, the area is an extensive floodplain that gets inundated most of the year with a short dry season from July to September (INRENA, 2000). Indeed, 86% of the area is represented by inundated forest (51%), seasonally flooded forest (34%), and rivers and oxbow lakes (1%) (Bayley et al., 1992), making the landscape a complex mosaic of water bodies all interconnected during the inundation season. Extensive beds of floating macrophytes cover approximately 30-40% of the total open water surface area at high water (INRENA, 2000). The taxonomic composition of macrophytes varies from one patch to another. The most abundant plant species occurring in the sampled patches were Polygonium sp., 7

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8 Pistia stratiotes (L.), Eichhornia crassipes Solms., Paspalum sp., and an unidentified legume species. Other taxa include Azolla sp., Neptunia sp., Ludvigia sp., Salvinia sp., Utricularia sp., and Echinochloeta sp. While most of these macrophyte species are buoyant and drift with wind and water currents, herbaceous species such as Paspalum repens Berg, and Polygonium sp.; and shrubs and trees such as Sena sp., and Cecropia sp. are rooted to the bottom. Rooted plants made the sampling effort with gillnet very difficult and in many occasions it was impossible to cut depth enough into the meadows to clear a patch for the nets. Cao Yarina is a small affluent (no more than 100 m width) of the Rio Pacaya. Conductivity ranges from 100 to 200 siemens cm -1 in the Pacaya River, similar to that of white water, despite a dark coloration of the river which is more typical of blackwaters. Water depth varied from 0.6 to 2.5 m in the sampling places in the flooded forest and from 2.5 to 6.5 m in the floating meadows. Transparency was measured with a Secchi disk and ranged from 0.8 to 2.2 m in flooded forest and from 1.5 to 2.8 m in the floating meadows. Margins of the channel were covered by extensive beds of floating meadows. Behind these macrophytes were patches of shrubs, palms and Cecropia sp. trees which all together constitute the levee zone. The vrzea flooded forest grows behind this zone. Fish sampling. This study was designed to quantify fish assemblages in a way that would facilitate comparison between flooded forests and floating meadows. Fish collection was conducted during 9 to 21 May 2003 in two habitats: flooded forests and floating meadows. In each habitat 10 locations where chosen, separated by at least one km, with the exception of Locations 7 and 8.

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9 At each location I selected three sampling positions, no further than 100 m apart. At each of these three positions a set of four gill nets of 25, 40, 80 and 120 mm mesh size were deployed. In the flooded forest, the nets were located from the edge of the dry land towards the open water, parallel to each other and with the bigger to the smaller mesh sizes facing the current direction (Jensen, 1990). In floating meadows nets were located around patches of vegetation and the order of mesh sizes was randomly assigned (Fig. 2). Nets were 20 m long and depth varied from 3.5 to 5 m. Fishing was conducted for a period of 18 h at each location. After selecting a location and recording its location with a Global Positioning System (GPS) receiver, one team of two people went inside the flooded forest and another team into the floating meadows to select the three sampling positions. Nets were deployed around 18:00 h. At midnight all captured fishes were removed from the nets and brought to the field station to be identified and measured. The nets were visited again at 06:00 h and at noon, and the same procedure was followed each time. Nets were removed at noon and brought to the field station to be repaired if necessary and all were deployed again at 18:00 h at the next sampling positions. Standard length (SL), weight (W), time at capture, and mesh size for all captured fishes were recorded, and samples of each species were fixed in 10% formaldehyde and preserved in 70% ethanol. All preserved materials were deposited at the Florida Museum of Natural History (UF), University of Florida, USA. Data analysis. Because of the low number of captures at individual sampling positions, the specimens collected at all three sampling positions for a single location were combined to produce a meaningful representation of the location. Therefore I considered locations as sample units, representing each habitat. Statistical analyses were

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10 performed using Stat View 5.0 and JPM 5.0.1, except for Jackknife richness estimator and the multivariate analyses that were performed using PC-ORD software (McCune and Mefford, 1999). Species accumulation curves for each habitat were constructed based on number of species caught per location. Each location represented 18 h of continuous sampling. Total species per habitat were estimated with a first and second order Jackknife estimator (Heltshe and Forrester, 1983) using PC-ORD. Differences in assemblage characteristics between flooded forest and floating meadows were evaluated by conducting t-tests on abundance (total number of individuals), biomass (total weight of individuals), richness (total number of species), mean SL, maximum SL, mean weight, and maximum weight of fish from 10 locations of each habitat type. The same set of analyses was also conducted excluding individuals over 400 mm SL. Biomass was also estimated as capture per unit of effort (CPUE) by calculating the amount of grams of fish caught per meter square of net surface per day of sampling. To compare the distribution of abundance of species between flooded forest and floating meadows, I ranked all species in a habitat based on abundance (n = 72 species in flooded forest and n = 57 species in floating meadows), giving the rank = 1 to the most abundant species and continuing the ranking in descending order. If more than one species presented the same abundance I gave the same rank to all of those species and then calculate the average rank (ties). I then plotted the ranks against the number of individuals. Differences in median abundance between habitats for the 10 most abundant species were also detected using the nonparametric Mann Whitney U test.

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11 Temporal differences in abundance, biomass, and richness were explored using repeated measures ANOVA. This analysis allowed me to explore the effects of time of day, habitat, and their interaction on assemblage characters. Contrast analysis was performed to evaluate differences between time periods. Distribution of body size of all fishes captured in flooded forest versus floating meadows was compared using the Kolmogorov-Smirnov test for frequency distributions. To study the distribution of species in the two habitats, multivariate procedures were performed with a matrix of species abundance per location. The original matrix contained 20 locations x 78 species. Eighteen species present in only one location (5% of the sampling units) were deleted from the matrix resulting in a new matrix of 20 locations x 60 species. This reduces the noise of uninformative rare species and is appropriate after species richness and diversity has been analyzed from with the data (McCune and Grace, 2002). However, after deleting the rare species, the coefficient of variance and average skewness for species were still very high (319.82%, and 2.55 respectively). Therefore the abundance matrix was transformed using Log 10 (x + 1). This transformation is useful in analyzing community data since it decreases the importance of highly abundant species that would skew cluster and ordination procedures (McCune and Grace, 2002). An outliers analysis (PC-ORD) pointed out a location from floating meadows (FM5) as an outlier (standard deviation from average distance among sample units > 2) (McCune and Grace, 2002). Similarly, a location from flooded forests (FF5) showed as an isolated point in exploratory ordination analysis (near to the point representing FM5). Although the standard deviation (Sd) value did not pointed out FF5 as an outlier (Sd = 1.77) I removed the two locations, based on the fact that these locations where highly

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12 predated by piranhas, therefore the species richness and species abundances were remarkable low in both. These two locations produced an artificial clumping effect in the remaining 18 locations, over the axis correlated with species richness and diversity. Finally, I removed from the matrix three species present in only one location after the removal of locations FM5 and FF5. The new matrix of 57 species and 18 locations of flooded forest and floating meadows was analyzed with a nonmetric multi-response permutation procedure (MRPP) in order to test the null hypothesis of no difference in species composition between the two habitats. The MRPP (Mielke, 1984) is a nonparametric multivariate procedure to test for differences between a priori defined groups (e.g., habitats). The nonmetric test transforms the distance matrix in ranks prior to calculating the test statistic (T). T describes the separation between groups and is associated to a p-value. A more negative value of T means a stronger separation between groups. MRPP also provides a measure of effect size called A, which describes within group homogeneity, compared with the random expectation. A ranges from 0 to 1, therefore, when A=0 the heterogeneity within groups is not different from that expected by chance, whereas, when A = 1 all items are identical within each group (McCune & Grace, 2002). McCune et al. (2000) point out that in community ecology studies, A < 0.1 is common, even when groups are obviously differ and A > 0.3 is considered high. To represent the similarities in species composition between locations, I used nonmetric multidimensional scaling (NMS; Kruskal, 1964; Mather, 1976). NMS is a nonparametric ordination technique; starting from a matrix of species abundance per sample, this procedure calculates coefficients of dissimilarity for each species, ranks

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13 those coefficients, and maps the samples in two or more dimensions, in which distance between samples reflects similarity in species composition (Clarke, 1993). The aim of the analysis is to locate samples in positions (in the graph) that result in the lowest stress, stress been a measure of departure of sample positions in the graph from the initial dissimilarity matrix (Clarke, 1993). Therefore in PC-ORD stress is scaled from 0 to 100 where zero means perfect agreement in rank orders (McCune & Grace, 2002). NMS has advantages when analyzing community data by not assuming multivariate normality and by being robust to large numbers of zero values (Clarke, 1993; McCune & Grace, 2002). The analysis was performed through the autopilot slow-and-thorough option of NMS in PC-ORD. This option performed 40 runs with real data and 50 runs with randomized data to find a Monte Carlo test of significance for the best output. The resulting ordination was rotated 15 degrees. Rotation maximizes the percentage of variance explained by each axis (McCune & Grace, 2002). The variance explained was expressed by the coefficient of determination between distances in the ordination space and distance in the original species space. Sorensen distance was selected in both cases since this distance measure has been recommended for analyzing community data (McCune & Grace, 2002). Environmental and assemblage variables were overlaid by using a joint plot, based on the individual correlations of those variables with the axes of community ordination (see Clarke, 1993; McCune et al., 2000; and Peterson & McCune, 2001 for examples of applying this ordination technique on community data).

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14 Figure 1. Map of the study area in the Upper Amazon, Pacaya Samiria National Reserve, Peru. Number one indicates the location of guard post, Puesto de vigilancia 2 (PV2), around which the sampling was conducted. Map was made by W.G.R. Crampton based on 1998 1:300,000 Landsat TM5 images; and used with his permission.

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15 L evee Mesh sizes: 40mm 80mm 120mm 25mm Floating meadows Flooded forest Figure 2. Sampling place in Location 1. Different mesh size nets are represented by shades. Nets closer to Cao Yarina, Pacaya Samiria National Reserve, Peru, are located in floating meadows whereas nets located next to the land edge, are placed into the adjacent flooded forest. Arrows represent the direction of the water current.

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RESULTS A total of 2789 individuals representing 6 orders, 20 families, 61 genera and 78 species of fishes were captured in this study (See appendix). Characiformes (35 species) was the most diverse taxon followed by Siluriformes (27 species), Perciformes (6 species of Cichlids), Gymnotiformes (7 species), Osteoglossiformes (2 species), and Synbranchiformes (1 species). Characiformes and Siluriformes alone accounted for 80% of the total number of species. Seventy three species were caught in flooded forest, 21 of which were exclusively found in this habitat. In floating meadows, 57 species were caught and only five were exclusively found in this habitat (Table 1). Species richness per location ranged from 9 to 33 in flooded forest (average 19.3 7.6) and from 14 to 30 in floating meadows (average 19.0 5.1). Species richness approximates an asymptote in both habitats (Fig. 3), for the species with individuals ranging from 38 to 740 mm SL. Total species estimates were 102 (first order jackknife) and 117 (second order jackknife) for flooded forest. In this habitat, 32 species (44%) occurred in only one location. In floating meadows total species was estimated in 74 (first order jackknife) and 79 (second order jackknife). In this habitat, 19 species (33%) occurred in only one location. Assemblage characters, averaged across locations and including abundance (total number of individuals), biomass (total weight of individuals), richness (total number of species), mean SL, maximum SL, mean weight, and maximum weight of fish were 16

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17 similar in between flooded forest and floating meadows (Table 2). Similar results were found when individuals over 400 mm SL were excluded from the analysis. Total biomass was higher in flooded forest than in floating meadows (71.0 kg vs. 49.9 kg respectively). A total of 1000.22 m 2 gill net surface area was deployed at each habitat. Capture per unit effort was estimated as 7.10 g fish m -2 per day in flooded forest and 4.99 g fish m -2 per day in floating meadows for fishes ranging between 38 to 740 mm (n = 2789 individuals). Species abundances at each habitat followed an expected hollow distribution (Hubbell, 2001) where few species were very abundant and most of the remaining species were represented by few individuals (Fig. 4). Indeed, in flooded forest 70% of the total abundance was accounted for by three species: Dianema longibarbis, Psectrogaster rutiloides, and Triportheus angulatus, while 25 species (35% of the total richness) occurred only once. In floating meadows 60% of the total abundance was represented by D. longibarbis, P. rutiloides, and Psectrogaster amazonica (Fig. 5), and 12 species (21% of the total richness) occurred only once. Among the 10 most abundant species, only T. angulatus was significantly more abundant in flooded forest than in floating meadows (Mann Whitney, U = 6.0, P < 0.001; FF 30.9 19.0; FM 4.8 3.4), whereas Hoplias malabaricus (U = 15.5, P < 0.01; FF 0.6 1.1; FM 3.2 2.7), Cichlasoma amazonarum (U = 24.50, P < 0.05; FF 0.7 1.6; FM 3.0 2.8), and Mesonauta mirificus (U = 20.50, P < 0.05; FF 0.4 0.7; FM 3.0 2.8) were significantly more abundant in floating meadows (Fig. 5). Numbers of individuals and species richness were higher at night; however biomass did not show diel differences (Table 3). Abundance and richness were highest at the

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18 18:00 to 24:00 period and the lowest at the 6:00 to 12:00 period (Fig. 6). Moreover, there were temporal differences in abundance within each habitat, as well. In flooded forest the abundance was highest during the 18:00 to 24:00 period and lowest at the 6:00 to 12:00 period (Contrast analysis for: 18:00 to 24:00 vs. 6:00 to 12:00, F = 45.31, df = 1, P < 0.0001; 24:00 to 6:00 vs. 6:00 to 12:00, F = 9.72, df = 1, P < 0.01; Fig. 6). In floating meadows the abundance was also highest at the 18:00 24:00 period, but it was equally lower in the other two periods of time (Contrast analysis for: 18:00 24:00 vs. 6:00 to 12:00, F = 12.36, df = 1, P < 0.01; 24:00 to 6:00 vs. 6:00 to 12:00, F = 0.36, df = 1, P = NS; Fig. 6). At the species level, the five most common species (D. longibarbis, P. rutiloides, T. angulatus, P. amazonica, and Ctenobrycon spilurus) represented highest abundances during the nocturnal samples (18:00 to 24:00 and 24:00 to 06:00) in both flooded forests and floating meadows (Fig. 7). There were no temporal differences in species richness within each habitat. Biomass was similar at day and night samples with approximately 2 kg of fish per net area (2000.4 m 2 ) in a 6 h period (Fig. 6). Body size distributions were very similar in the two habitats. There were no significant differences in the frequencies of SL intervals (Kolmogorov-Smirnov, X 2 = 0.667, P = NS) (Fig. 8). Community-structure patterns. Flooded forest locations differed from floating meadows locations in species composition (MRPP A = 0.15, P < 0.001, where A describes within group homogeneity compared to the random expectation). Although the A value was small, it is significantly different from zero and indicates that the homogeneity within groups is higher than expected by chance, meaning that there is a different assemblage at each habitat type.

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19 Flooded forest locations separates from floating meadows locations along the vertical ordination axis (Fig. 9). After rotation, the vertical axis explained 32% of the variation in fish assemblages and was correlated with water depth (Pearson Correlation, r = 0.68, P < 0.01), but not significantly correlated with water transparency (r = 0.40, P = NS). The vertical axis was also positively correlated with the abundance of C. amazonarum (r = 0.65, P < 0.005), H. malabaricus (r = 0.65, P < 0.005), Crenicichla proteus (r = 0.59, P < 0.01), M. mirificus (r = 0.53, P < 0.05), and negatively correlated with the abundance of T. angulatus (r = -0.86, P < 0.001), Callichthys callichthys (r = -0.71, P < 0.001), Gymnotus ucamara (r = -0.67, P < 0.005), Ancistrus sp. PUA (r = -0.62, P < 0.01), and Hoplerythrinus unitaeniatus (r = -0.60, P < 0.01). The horizontal axis explained 30% of the variation in fish assemblages and was positively correlated with the abundance of P. rutiloides (r = 0.85, P < 0.001), P. amazonica (r = 0.82, P < 0.001), Potamorhina altamazonica (r = 0.68, P < 0.005), Curimatella meyeri (r = 0.58, P < 0.05), Doradidae sp. PUA (r = 0.51, P < 0.05), and negatively correlated with the abundance of Colossoma macropomum (r = -0.50, P < 0.05) and Liposarcus pardalis (r = -0.49, P < 0.05). A third axis and explained 20% of the variation in fish assemblages and was correlated with overall species richness (r = 0.84, P < 0.001), The coefficients of correlation with environmental and assemblage variables were obtained from a joint plot and are in accordance with the highest values of depth (t = -8.31, P < 0.0001; 144 cm 47.8 in flooded forest vs. 403 cm 85.8 in floating meadows). Species richness was similar in both habitats (t = 0.1, P = NS; 19.3 species 7.6 in flooded forest vs. 19.0 species 5.1 in floating meadows).

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20 Literature review and personal observations by the author and W.G.R. Crampton, were used to categorize the diet of caught species. In flooded forest, invertebrate feeders (30% of 73 species) was the trophic guild with higher number of species, followed by detritivores (20%), piscivores (15%), and frugivores (7%). Relative abundance, showed a similar pattern, but in each guild, there was a single species that accounted for most of the abundance in this habitat. These species were D. longibarbis (39% of the total abundance), T. angulatus (20%), and P. rutiloides (13%) (Table 4). In floating meadows, a similar pattern to the one found in flooded forest was observed. Invertebrate feeders (40% of 57 species), detritivores (28%), and piscivores (16%) were the trophic guilds with higher number of species. However the relative abundance of invertebrate feeders, detritivores and piscivores was higher. Again a single species accounted for most of the abundance in each guild. Dianema longibarbis (36% of the total abundance), was the most abundant species, P. rutiloides (17% of the total abundance), was the second most abundant species and the abundance of T. angulatus (4% of the total abundance), was much lower than in flooded forest.

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Table 1. Abundance of 78 species in each of 10 locations in flooded forest and floating meadows of the Cao Yarina, Pacaya Samiria National Reserve. Taxon FM 1 FM 2 FM 3 FM 4 FM5 FM6 FM7 FM8 FM9 FM10 FF1 FF 2 FF 3 FF 4 FF5 FF 6 FF 7 FF 8 FF9 FF10 Total Acestrorhynchus falcatus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 Acestrorhynchus falcirostris 0 0 0 0 0 0 0 0 0 0 0 0 0 6 1 0 0 0 0 0 7 Acestrorhynchus microlepis 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 Adontosternarchus sp. A 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 Agamyxis pectinifrons 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 2 Anadoras grypus 0 0 0 1 1 0 0 0 0 0 0 0 0 3 0 0 1 4 0 1 11 Ancistrus sp. PUA 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 2 Arapaima gigas 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 Astyanax bimaculatus 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 3 Auchenipterichthys longimanus 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 2 Brochis splendens 0 0 0 0 0 0 14 5 0 0 0 0 0 0 0 0 2 0 0 0 21 Brycon cephalus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 Callichthys callichthys 0 0 0 0 0 0 0 1 0 0 2 0 1 0 0 0 1 1 0 0 6 Charax gibbosus 0 0 0 1 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 3 Cichlasoma amazonarum 7 3 0 4 0 2 4 0 2 8 0 5 0 0 0 0 1 0 0 1 37 Colossoma macropomum 1 0 0 0 0 0 0 5 0 1 1 0 0 0 0 3 1 3 0 3 18 Crenicichla proteus 1 0 0 5 2 0 3 0 5 2 0 0 1 0 0 0 0 0 0 0 19 Ctenobrycon spilurus 1 0 9 9 0 3 1 0 1 1 0 23 9 20 1 12 5 3 2 0 100 Curimata vittata 0 0 0 1 0 0 0 0 0 0 1 3 0 0 0 0 0 0 0 0 5 Curimatella alburna 0 2 0 4 0 0 0 0 0 0 0 6 1 10 0 0 3 0 0 0 26 Curimatella meyeri 9 2 2 3 0 0 0 2 0 0 0 8 0 1 0 0 0 2 0 0 29 Cyphocharax cf. festivus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 Dianema longibarbis 105 46 15 29 1 56 52 46 23 63 29 124 226 57 8 50 22 22 16 42 1032 21

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Table 1. Continued Taxon FM 1 FM 2 FM 3 FM 4 FM5 FM6 FM7 FM8 FM9 FM10 FF1 FF 2 FF 3 FF 4 FF5 FF 6 FF 7 FF 8 FF9 FF10 Total Doradidae sp. PUA 2 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 5 Doradidae sp. PUB 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 Eigenmannia limbata 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 Doradidae sp. PUA 2 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 5 Doradidae sp. PUB 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 Eigenmannia limbata 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 Electrophorus electricus 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 2 Erythrinus erythrinus 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 2 Gasteropelecus sternicla 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 2 Gymnocorymbus thayeri 1 0 0 1 0 0 2 0 0 2 0 0 4 6 0 0 4 0 0 0 20 Gymnotus carapo 0 1 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0 0 4 Gymnotus ucamara 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 1 0 1 4 Gymnotus varzea 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 Heros appendiculatus 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 2 Hoplerythrinus unitaeniatus 0 0 0 0 0 0 0 0 0 0 4 0 1 1 0 1 0 3 0 1 11 Hoplias malabaricus 4 2 1 7 0 1 8 4 1 4 0 0 0 3 0 2 0 1 0 0 38 Hoplosternum littorale 3 1 0 0 0 1 0 2 1 3 1 3 4 5 0 3 3 0 0 0 30 Hypoptopoma gulare 0 0 0 3 7 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 12 Hypselacara temporalis 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 2 3 Leporinus trifasciatus 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 2 0 0 0 3 Lepthoplosternum sp. PUA (black belly) 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 Lepthoplosternum sp. PUB (pepper belly) 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 Liposarcus pardalis 0 1 0 0 0 3 1 9 8 2 9 1 2 4 0 1 1 0 0 1 43 Loricariichthys cf. acutus 0 0 1 0 0 0 5 0 0 0 0 0 0 0 0 0 2 1 0 0 9 22

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Table 1. Continued Taxon FM 1 FM 2 FM 3 FM 4 FM5 FM6 FM7 FM8 FM9 FM10 FF1 FF 2 FF 3 FF 4 FF5 FF 6 FF 7 FF 8 FF9 FF10 Total Loricariichthys cf. maculatus 0 0 0 1 0 0 2 0 0 0 0 1 0 1 0 4 1 0 0 0 10 Loricariichthys cf. nudirostris 0 0 0 0 0 0 5 0 0 0 0 0 0 0 0 0 2 0 0 0 7 Loricariichthys sp.1 0 0 0 2 0 0 0 0 0 0 0 0 0 0 1 2 0 0 0 0 5 Loricariichthys sp.2 (possibly juvenile of maculatus) 0 5 3 0 0 1 0 0 1 17 0 1 2 0 0 0 0 0 0 0 30 Loricariinae sp. Indet. TF03 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 Megalechis thoracata 0 1 0 0 0 3 1 0 0 1 0 4 4 0 0 9 0 5 1 3 32 Mesonauta mirificus 8 0 2 4 3 5 0 0 2 6 1 1 0 0 2 0 0 0 0 0 34 Moenkhausia cf. chrysagyrea 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 2 Mylossoma duriventre 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 2 Osteoglossum bicirrosum 0 0 0 1 0 0 3 1 0 2 0 2 0 1 0 2 0 0 0 1 13 Parapteronotus hasemani 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 Potamorhina altamazonica 5 3 4 2 0 4 1 0 2 1 0 18 1 5 0 1 7 1 0 0 55 Potamorhina latior 13 3 0 2 0 0 0 0 0 0 0 3 0 0 0 0 0 0 0 0 21 Prochilodus nigricans 0 0 0 0 0 4 3 0 0 0 4 0 0 0 0 0 5 0 0 0 16 Psectrogaster amazonica 7 14 39 2 2 9 1 5 3 0 0 21 0 0 1 0 12 8 7 0 131 Psectrogaster essequibensis 0 3 0 0 0 0 0 0 0 0 0 7 0 0 0 0 2 0 0 0 12 Psectrogaster rutiloides 65 56 23 17 1 20 2 5 5 15 1 140 1 3 0 2 26 4 24 1 411 Pseudorinelepis genibarbis 0 0 0 0 2 0 1 0 0 0 1 0 0 1 0 0 0 0 0 0 5 Pterodoras granulosus 0 2 0 2 0 0 0 0 0 0 0 16 0 0 0 0 0 0 0 0 20 Pterygoplichthys scrophus 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 Pygocentrus nattereri 4 0 2 4 10 0 7 0 0 0 0 0 0 1 1 0 0 0 0 0 29 23

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Table 1. Continued Taxon FM 1 FM 2 FM 3 FM 4 FM5 FM6 FM7 FM8 FM9 FM10 FF1 FF 2 FF 3 FF 4 FF5 FF 6 FF 7 FF 8 FF9 FF10 Total Rhamdia quelen 0 0 0 2 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 3 Rhytiodus microlepis 0 0 0 0 3 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 5 Roeboides biserialis 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 Satanoperca jurupari 0 0 0 4 0 0 0 0 0 0 0 0 0 2 0 1 2 0 0 0 9 Schizodon fasciatus 0 0 1 2 0 0 0 1 0 0 0 0 0 1 2 0 0 1 0 0 8 Serrasalmus rhombeus 0 0 3 1 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 6 Sorubim lima 0 0 0 0 2 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 3 Synbranchus marmoratus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 Tetragonopterus argenteus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 2 Tetragonopterus chalceus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 Trachelyopterus galeatus 2 2 0 0 0 0 1 0 0 2 1 5 2 4 0 0 1 2 0 0 22 Triportheus albus 0 1 1 0 1 1 0 0 2 0 0 0 0 0 1 0 0 1 0 0 8 Triportheus angulatus 3 10 1 10 3 5 0 7 4 5 37 43 55 22 5 20 63 33 23 8 357 Total 244 159 107 129 40 119 121 95 60 138 99 439 318 169 27 115 174 100 76 65 2789 24 Abbreviations: FM floating meadows; FF flooded forest. The numbers following the abbreviations correspond to the sampling location in each habitat (1-10).

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25 Table 2. Mean values of different assemblage characteristics for flooded forest and floating meadows. FF FM P Mean abundance (total number of fish) 157.9 128 121.4 56 0.4189 Mean biomass (g) 7099.5 4510 4994.1 4516 0.3107 Mean richness (species) 19.4 8 19.3 5 0.9731 Mean standard length (mm) 97.6 15 94.6 15 0.6474 Maximum standard length (mm) 527.8 164 366.4 196 0.0806 Mean weight (g) 62.9 50 45.7 47 0.4368 Maximum weight (g) 1717.0 1283 1156.1 1242 0.3339 Calculations are based on totals, means, or maximum values averaged across 10 locations in each habitat type. Abbreviations: FF flooded forest, FM floating meadows. A t-test was used to detect differences between the two habitats in Cao Yarina, Pacaya Samiria National Reserve.

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26 Table 3. Comparison of day and night captures between flooded forest and floating meadows. Repeated measures ANOVA was used to detect effects of time, habitat and their interaction on abundance (log10x+1 transformed), richness (log10x+1 transformed), and biomass (log10x+1 transformed) of fishes with three times of capture (18:00 24:00h, 24:00 6:00h, 6:00 12:00h) and two habitats in Cao Yarina, Pacaya Samiria National Reserve. For abundance, Mauchley's criterion indicated rejection of the compound symmetry assumption, therefore adjusted probability values (G, Greenhouse-Geisser and H-F, Huynh-Feldt) are provided. SOURCE OF VARIATION df SS F P Adj. G-G H-F Abundance Between-subjects effect Habitat 1 0.117 0.452 NS Error 18 4.641 Within-subjects effect Time 2 4.222 24.894 <0.001 Time*habitat 2 1.094 6.453 <0.005 <0.05 <0.01 Error (time) 36 3.053 Biomass Between-subjects effect Habitat 1 0.018 0.023 NS Error 18 14.092 Within-subjects effect Time 2 1.478 1.754 NS Time*habitat 2 0.987 1.172 NS Error (time) 36 15.167 Richness Between-subjects effect Habitat 1 0.169 1.963 NS Error 18 1.550 Within subjects effect Time 2 0.820 10.419 <0.001 Time*habitat 2 0.164 2.087 NS Error (time) 36 1.416

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27 Table 4. Principal food items for 78 species and their relative abundance as a percentage of the total abundance (n = 2793 individuals) in flooded forest and floating meadows, during May 2003 at Cao Yarina, Pacaya Samiria National Reserve. Food items were extracted from literature. Diet of species in which a reference is not provided was assessed by W.G.R. Crampton and S.B. Correa. Taxon Diet Reference Relative abundance Acestrorhynchus falcatus Fish Planquette et al. (1996) 0.04 Acestrorhynchus falcirostris Fish Goulding et al. (1988) 0.25 Acestrorhynchus microlepis Fish Planquette et al. (1996) 0.04 Adontosternarchus sp. A Invertebrates 0.04 Agamyxis pectinifrons Invertebrates 0.07 Anadoras grypus Invertebrates 0.39 Ancistrus sp. PUA Detritus 0.07 Arapaima gigas Fish 0.04 Astyanax bimaculatus Zooplankton Planquette et al. (1996) 0.11 Auchenipterichthys longimanus Fruits Manheimer et al. (2003) 0.07 Brochis splendens Aquatic invertebrates Burgess (1989) 0.75 Brycon cephalus Fruits Goulding (1980), Anonymous (1981) 0.04 Callichthys callichthys Zooplankton Mol (1995) 0.21 Charax gibbosus Fish Winemiller (1989) 0.11 Cichlasoma amazonarum Plants Stawikowski & Werner (1998) 1.32 Colossoma macropomum Fruits Goulding (1980) 0.64 Crenicichla proteus Fish Kullander (1986) 0.68 Ctenobrycon spilurus Zooplankton Mills & Vevers (1989) 3.59 Curimata vittata Detritus Val & de Almeida-Val (1995) 0.18 Curimatella alburna Detritus Goulding et al. (1988) 0.93 Curimatella meyeri Plants Soares et al. (1986) 1.04 Cyphocharax cf. festivus Detritus 0.04 Dianema longibarbis Invertebrates 36.95 Doradidae sp. PUA Invertebrates 0.18 Doradidae sp. PUB Invertebrates 0.04 Eigenmannia limbata Invertebrates 0.04 Electrophorus electricus Fish 0.07

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28 Table 4. Continued Taxon Diet Reference Relative abundance Erythrinus erythrinus Fish Planquette et al. (1996) 0.07 Gasteropelecus sternicla Aquatic invertebrates Mills & Vevers (1989) 0.07 Gymnocorymbus thayeri Invertebrates 0.72 Gymnotus carapo Invertebrates 0.14 Gymnotus ucamara Aquatic invertebrates Crampton et al. (2003) 0.14 Gymnotus varzea Aquatic invertebrates 0.04 Heros efasciatus Plants 0.07 Hoplerythrinus unitaeniatus Aquatic invertebrates 0.39 Hoplias malabaricus Fish Planquette et al. (1996) 1.36 Hoplosternum littorale Aquatic invertebrates Boujard et al. (1997) 1.07 Hypoptopoma gulare Detritus 0.43 Hypselacara temporalis Fish 0.11 Leporinus trifasciatus Periphyton 0.11 Lepthoplosternum sp. PUA (black belly) Invertebrates 0.04 Lepthoplosternum sp. PUB (pepper belly) Invertebrates 0.04 Liposarcus pardalis Detritus Yossa & Araujo-Lima (1998) 1.54 Loricariichthys cf. acutus Aquatic invertebrates Goulding et al. (1988) 0.29 Loricariichthys cf. maculatus Detritus 0.39 Loricariichthys cf. nudirostris Detritus 0.25 Loricariichthys sp.1 Detritus 0.18 Loricariichthys sp.2 (may be maculatus juvenile) Detritus 1.07 Loricariinae sp. Indet. TF03 Detritus 0.04 Megalechis thoracata Zooplankton Mol (1995) 1.15 Mesonauta mirificus Periphyton 1.22 Moenkhausia cf. chrysagyrea leucopomis Invertebrates 0.07 Mylossoma duriventre Fruits Goulding (1980) 0.07

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29 Table 4. Continued Taxon Diet Reference Relative abundance Osteoglossum bicirrosum Terrestrial invertebrates Goulding (1980) 0.47 Parapteronotus hasemani Invertebrates 0.07 Potamorhina altamazonica Detritus Soares et al. (1986) 1.79 Potamorhina latior Detritus Goulding et al. (1988) 0.75 Prochilodus nigricans Detritus Soares et al. (1986) 0.57 Psectrogaster amazonica Detritus 4.87 Psectrogaster essequibensis Detritus 0.43 Psectrogaster rutiloides Detritus 14.72 Pseudorinelepis genibarbis Detritus 0.18 Pterodoras granulosus Fruits 0.72 Pterygoplichthys scrophus Detritus 0.04 Pygocentrus nattereri Fish Sazima & Machado (1990) 1.04 Rhamdia quelen Fish Boujard et al. (1997) 0.11 Rhytiodus microlepis Plants Soares et al. (1986) 0.18 Roeboides biserialis Fish scales 0.04 Satanoperca jurupari Invertebrates Keith et al. (2000) 0.32 Schizodon fasciatus Plants Planquette et al. (1996) 0.29 Serrasalmus rhombeus Fish Correa (1999) 0.21 Sorubim lima Fish Goulding (1981) 0.11 Synbranchus marmoratus Terrestrial invertebrates Soares et al. (1986) 0.04 Tetragonopterus argenteus Invertebrates Silvano et al. (2001) 0.07 Tetragonopterus chalceus Aquatic invertebrates Goulding et al. (1988) 0.04 Trachelyopterus galeatus Fish Le Bail et al. (2000) 0.79 Triportheus albus Invertebrates 0.29 Triportheus angulatus Fruits Goulding (1980) 12.80

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30 0 10 20 Cumulative No. Species 30 40 FF FM 8 9 10 11 7 6 5 4 2 3 0 1 80 70 60 50 Samples Figure 3. Species richness saturation curves for flooded forest and floating meadows samples. Fish ranged from 38 mm to 740 mm (n=2789 individuals) and were caught with gill nets of 25, 40, 80, and 120 mm mesh size. Sampling was conducted during May 2003 at Cao Yarina, Pacaya Samiria National Reserve, Peru.

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31 1 10 100 1 A bundance (# individuals) 000 pees Rank of s 61 61 42ci 33 26 1 10 18 61 61 A Figure 4. Rank ordered abundances (absolute number of individuals per species) of 78 species of fish captured during May 2003 in Cao Yarina, Pacaya Samiria National Reserve, Peru. A) flooded forest (72 species). B) floating meadows (57 species). Mean species richness was equal in the two habitats (P= 0.9731). P value originated by ttest. Note the hollow curve where only three species account for 70% and 60% of the total abundance in flooded forest and floating meadows, respectively. See text for method of ranking species. 1 10 100 pecs 1000 A bundance (# individuals) 1 9 17 25 32ie 42 52 52 Rank of s B

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32 0100200300400500600700DianemalongibarbisPsectrogasterrutiloides** TriportheusangulatusPsectrogasteramazonicaCtenobryconspilurusPotamorhinaaltaamazonica Liposarcuspardalis* Hopliasmalabaricus* Cichlasomaamazonarum* MesonautamirificusSpeciesAbundance (# of individuals) FF FM Figure 5. Abundance per habitat of the 10 most abundant species (absolute number of individuals per habitat) captured during May 2003 in Cao Yarina, Pacaya Samiria National Reserve, Peru. ** indicates P>0.001, indicates P>0.05. Abbreviations: FF flooded forest, FM floating meadows.

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33 01000200030004000ENLNEDTime C 0481216ENLNED A 04080120ENLNED FF FMBaedbce Figure 6. Variation in abundance, biomass, and richness during three periods of capture (EN, LN, LD) during May 2003 in Cao Yarina, Pacaya Samiria National Reserve, Peru. Graphs A and C represent means combining the two habitats, graph B shows variation between habitats. Bars represent standard error. Abbreviations: EN early night: 18:00 24:00; LN late night: 24:00-06:00; ED early day: 06:00 12:00; FF flooded forest, FM floating meadows.

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34 00 50 11 00 50 200 250 300 35400 FF FM Abundance (# of individuals) Ctenobr y con spilurus Psectrogaster amazonica Triportheus angulatus Psectrogaster rutiloides Dianema longibarbis LN ED EN LN EN ED Samples Figure 7. Changes in abundance of the five most common species among three sampling periods in flooded forest and floating meadows. Sampling was conducted during May 2003 at Cao Yarina, Pacaya Samiria National Reserve, Peru. Abbreviations as in Figure 6.

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35 0100200300400500600700507090110130150200250300350400>400SL Interval (mm)Frequency FF FM Figure 8. Size frequency distributions of fishes captured per habitat during May 2003 in Cao Yarina, Pacaya Samiria National Reserve, Peru. Abbreviations: SL standard length, FF flooded forest, FM floating meadows.

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36 Depth Transparency HabitatFMFFNMS 1NMS 3 Figure 9. Non metric scaling ordination (NMS) of sampling locations in species space based on the abundances of 57 species that occurred in more than 5% of the sampling locations during May 2003 in Cao Yarina, Pacaya Samiria National Reserve, Peru. The vertical axis accounts for 32% while the horizontal axis accounts for 30% of the variation in the data. Filled triangles represent sampling locations in flooded forest (FF) and open triangles represent sampling locations in floating meadows (FM). The distances between triangles in the ordination are approximately proportional to the dissimilarities between the sampling locations. Environmental variables are join-plotted expressing its relationship with ordination scores. Length and angle of correlation vectors represents the strength of the correlation. The final stress value for a three dimensional representation was 11.367. Instability (0.000001) was the lowest out of the 10 repetitions.

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DISCUSSION Species Richness and Abundance The present study provides the first empirical data on the relative species abundance in an Amazonian flooded forest. Relative abundances of species often follow a hollow distribution with very few dominant species and a long tail of rare species (Hubbell, 2001; Magurran & Henderson, 2003). This hypothesis was corroborated in a study of fish assemblages in Amazonian floating meadows (Henderson & Crampton, 1997). In flooded forests of Cao Yarina, 70% of the total abundance was represented by three species: D. longibarbis, P. rutiloides and T. angulatus; while 25 species (35% of the total richness) only occurred once. In floating meadows 60% of the total abundance was represented by D. longibarbis, P. rutiloides and P. amazonica; while 12 species (21% of the total richness) occurred only once. What allows a species to be abundant in a habitat depends on characteristics of the habitat such as food resources and habitat structure, on ecological processes such as competition and predation, and on stochastic factors. Moreover, the identity of the dominant species in a habitat may change from place to place and year to year depending on migration and recolonization of the habitats (Hubbell, 2001). The reproductive biology of the callichthyid D. longibarbis, the dominant species in both habitats, is not well known. Callichthyids in general are reported as Kstrategists (sensu Pianka, 1970) (Junk et al., 1997) and Riehl & Baensch (1991) reported bubble nest-building behavior in D. longibarbis. Other species with nesting behaviors are defined as equilibrium life 37

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38 history strategist (sensu Winemiller, 1989), characterized also by prolonged breeding seasons, and parental investment in individual offspring, probably resulting in enhanced juvenile survivorship. This type of species usually has relatively stable sedentary local populations (Winemiller, 1989). This fact allows me speculate that D. longibarbis may be a year round resident of floating meadows that sends colonizers to adjacent flooded forest during the rise of the waters. To prove this hypothesis, a multiseason study in these habitats would be necessary. In this study, species richness was similar in both flooded forest and floating meadows (72 and 57 species respectively). First and second order Jackknife estimators (Heltshe & Forrester, 1983) estimate the number of species to be higher in both habitats than my absolute accounts and the number of species in flooded forest as higher (102 and 117 species respectively) than in floating meadows (74 and 79 species respectively). First order Jackknife estimators has been considered as the most accurate method for estimating species richness (Palmer 1990; Palmer 1991), however, it is susceptible to high number of rare species in the data set (Palmer 1990); indeed I found four times more rare species in flooded forest than in floating meadows. A higher species richness and biomass in flooded forest was expected because the flooded forest constitutes an expansion of available habitat in the floodplain during the flooded season, therefore is expected that many species would colonize this habitat with the rise of the water. The diversity and abundance of resources provided by flooded forests (Goulding 1980, Junk et al., 1997b) seems to be the driving force for colonization. For migratory fishes, flooded forests are a key habitat that provides enough food for them to accumulate fat reserves as

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39 preparation for subsequent reproductive migration at the low waters season (Junk et al., 1997b, Carvalho & Araujo-Lima, 2004). Biomass and Size Distributions Higher biomass along with higher species richness was expected in flooded forest because of increased availability of habitat during the flooded season. Total biomass was similar in the two habitats (73.3 kg vs. 51.1 kg for flooded forest and floating meadows respectively) and there was no difference in average biomass between habitats. Flooded forest however had a slightly higher CPUE than floating meadows (7.33 and 5.11 g fish m -2 net surface per sampled day, respectively). A comparable estimate of CPUE in flooded forest was much higher (104 g m 2 per day; Saint-Paul et al., 2000) than the CPUE value reported in the present study. Although the area covered during sampling in Cao Yarina was higher than the sampled area covered by Saint-Paul et al. (2000) (i.e., 1000.15 m 2 vs. 772 m 2 respectively), their study used a wider range of mesh sizes (13 different sizes, ranging from 12 to 200 mm) which in turn could caught a wider range of fish sizes, and therefore a highest biomass. For floating meadows, there is not comparable data since the data available are estimations of standing crop (Bayley 1983; Henderson & Hamilton 1995; Henderson & Crampton 1997). Body size distributions were highly similar between the two habitats, although differences were expected. A higher number of small fishes in floating meadows were expected, since this habitat is recognized as nursery habitat for juveniles of many species (Saint-Paul & Bayley 1979; Goulding 1980; Junk 1984, Sanchz-Botero & Araujo-Lima 2001), whereas in flooded forest small fishes could be more susceptible to predation; therefore bigger-sized fish were expected.

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40 The body size range found in Cao Yarina was truncated at the extremes (38 to 740 mm SL). This range most likely reflect the chosen mesh sizes (i.e., 25 to 120 mm) leading to under sample juveniles, small fish, and very large fish. Nevertheless, the shape of the distribution of body size frequencies of fishes in the sampled range was similar in both habitats. This result suggest that juveniles and small fishes (over 38 mm) of the species caught at Cao Yarina are using both flooded forest and floating meadows habitats in the same proportion. This contrast with previous hypothesis of floating meadows been a preferred habitat for juveniles because the availability of food and shelter (Junk, 1997). Large fishes (under 740 mm) seem also to be using both habitats in similar proportions despite the differences in habitat complexity. It is difficult to estimate what percentage of the community falls in and out of this range of body sizes found in Cao Yarina (38 to 740 mm SL). For example, in sandy beach communities in the Orinoco basin, Layman & Winemiller (2004) found that around 50% of the fishes were 50 mm SL. In a nutrient-poor river in the Amazon basin, Goulding et al. (1988) found that around 100 species (approximately 20% of total species richness) reach maturity at 30 mm, and all dominant species were adults 40 mm. In contrast, in the nutrient rich floodplain of the Solimoes-Amazonas River, juveniles of the medium sized fishes tend to dominate (Goulding et al., 1988). Goulding et al. (1988) hypothesize that in nutrient rich systems, the abundance of juveniles of bigger sized fishes would be higher compared with low nutrient systems, where early maturation seems to be a strategy due to competitive exclusion in food limited habitats. Indeed, in Cao Yarina (a nutrient rich system), the four dominant species were adults of medium

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41 sized fishes: D. longibarbis (46 to 103 mm), P. rutiloides (53 to 204 mm), T. angulatus (56 to 150 mm), and P. amazonica (54 to 131 mm). Diel Variation in Species Richness, Abundance and Biomass Diel changeover in species composition is known to occur in Neotropical fish assemblages (Arrington & Winemiller, 2003). In a study in the Orinoco floodplain, Arrington & Winemiller (2003) found nocturnal samples to be higher in number of species and individual abundances and explained the phenomenon as morphological trade-offs in foraging and anti-predator defenses. In Cao Yarina, the differences in species richness and abundance between day and night samples were impressive. In both flooded forest and floating meadows, species richness and abundance were much higher at night samples. While in flooded forest abundance was much higher at night samples (both early nigh and late night), in floating meadows abundance was higher at the early night samples but similar at both late night and early day samples species. Moreover, abundance at these time periods was higher than abundance at the early day samples in flooded forest. Differences in structural complexity between the flooded forest and floating meadows could be the mechanism allowing the higher abundance caught at day samples in floating meadows. The submerged portions of floating meadows are composed of a very dense matrix of stems and roots that may limit the pass of light during the day, whereas flooded forests, despite of having lots of stems and submerged branches, is a more open habitat. Consequently light penetration during the day time could be higher than in floating meadows. Therefore, predation risk at day time could be higher in flooded forests. It has been demonstrated that fishes are able to assess predation risk and

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42 modify their foraging behavior through foraging rate/mortality risk tradeoff mechanisms (Mittelbach, 1981; Werner et al., 1983; Mittelbach, 1984). Therefore predation avoidance behaviors may lead susceptible individuals to limit the use of rich habitats (Werner et al., 1983) as could be happening in flooded forest during the day time, but probably not in floating meadows where a more dense vegetation could reduce the risk of predation. However, the reduced abundance of fish predators in the samples does not provide evidence to support this hypothesis. The absence of diel change in biomass between habitats, despite of the differences in abundance, leads one to expect that a high amount of small fishes caught during night samples could weight as much as a few large fishes caught in day samples. Indeed, this was observed in flooded forest, where 21 individuals (23% of the total abundance) of 5 species (Arapaima gigas, C. macropomum, H. unitaeniatus, H. malabaricus, and L. pardalis) accounted for 85% of the total biomass in the day samples. In floating meadows, the trend was more dramatic. Twenty individuals (7% of the total abundance) of five species (C. macropomum, Electrophorus electricus, Leporinus trifasciatus, L. pardalis, and Osteoglossum bicirrhosum) accounted for 75% of the total biomass in the day samples. Fish Assemblages in Floodplains All quantitative analyses of community variables (i.e. abundance, biomass, species richness, and body size), without accounting for the species identity, did not provide evidence to separate the ichthyofauna of Cao Yarina in two assemblages. However, multivariate analysis of species abundances (i.e., MRPP and NMS ordination) suggested the presence of two subtly different fish assemblages in flooded forest vs. floating meadows. Floating meadows locations were characterized by deeper and more

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43 transparent waters than flooded forest. However, these habitats differ also in many other characteristics such as type of vegetation, structural complexity of the submerged portion of the vegetation, but especially in food resources. These variables are difficult to quantify and therefore it is hard to determine the individual effect on particular fish species. Specific habitat selection has been reported before in floodplain fish communities related to abiotic factors such as type of water (Saint-Paul at al., 2000), oxygen concentration (Junk et al., 1983; Winemiller, 1996), and water depth and transparency (Rodriguez & Lewis, 1994; Tejerina-Garro et al., 1998). In this last case, abundance of visually oriented predators (e.g., characids and cichlids) was positive correlated to water depth and transparency. In the present study, fish predators were scarce (19 and 4.6 % of the total species richness and abundance, respectively). Species of important predatory families such as Serrasalmidae were very scarce in the samples, indeed only two species were caught, Serrasalmus rhombeus and Pygocentrus nattereri. Only six juvenile specimens (45 to 76 mm) of S. rhombeus and 29 juvenile and adult specimens (63 to 205 mm) of P. nattereri were caught, mostly in the floating meadows. Pimelodids, another common family of silurid predators, was only represented by few specimens of Sorubim lima (n=3, 168 to 196 mm). Other common predators inhabiting flooded forest such as Cichla monoculus (Correa, 1999), were completely absent from the samples. Hoplias malabaricus, the most abundant predator species in Cao Yarina (1.4%), was scarcely found, mainly in floating meadows. A partial explanation for the low abundance of some large sized predator fishes is based on the limited efficiency of the gill nest. Surprisingly one individual of Arapaima gigas (a sub adult of 600 mm SL) was caught in this study when usually

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44 A. gigas are caught with harpoons or nets of mesh sizes bigger than 120 mm. However, it is unlikely that if other predator species, especially medium sized ones, were present in the system not even single individuals of each of those species were caught. Consequently an alternative explanation could be based on low productivity of the floodplain of Cao Yarina. CPUE in both flooded forest and floating meadows was low; therefore it is possible that low prey abundance could be limiting predators abundance as predicted by the Lotka-Volterra predation model (Begon et al., 1996). CPUE was low in the present study, but this fact alone does not provide sufficient evidence to support a low productivity hypothesis for Cao Yarina. Another possible explanation for the scarcity of some predator species (e.g., Cichla monoculus and Pimelodid species) could be based on over fishing in the area before the reserve was established. At the time this study was conducted, signals of illegal fishermen in the area were observed and indeed some of the gill nets used in this study were stolen. Variable food resources could also lead to differences in species composition between habitats. In both flooded forests and floating meadows, the majority of the species caught feed on invertebrates (aquatic and/or terrestrial), detritus, or fish. In flooded forest, invertebrate feeders and frugivores were dominant; whereas in floating meadows, invertebrate feeders and detritivores were dominant. These patterns were predicted, based on the differential food availability of flooded forests and floating meadows (seeds, fruits, and periphyton, vs. macrophytes, detritus, and authochtonous invertebrates) (Junk, 1997). Invertebrate feeders were expected to be an important guild in both habitats. Aquatic invertebrates are abundant in floating meadows (Junk, 1997) and terrestrial

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45 invertebrates are an important input of the terrestrial vegetation of the flooded forest (Goulding, 1980). The diversity of invertebrate feeders was high in both habitats but each species (except for D. longibarbis) presented a low abundance. It is possible that the ability of gymnotiform fishes to detect gill nets with their electric sense (W.G.R. Crampton, pers. commun.) may undermine the efficiency of gill nets in capturing them. Gymnotiforms are important components of floating meadows ichthyofauna (Crampton, 1996) but only seven species were found in this study, five of them been invertebrate feeders. Although a frugivore species (T. angulatus)was the second most abundant species in flooded forest, the total number of frugivore species found in Cao Yarina was surprising low. Serrasalminae, a sub-family with highly specialized frugivorous fishes (Goulding, 1980), was only represented by few individuals of Colossoma macropomum, and Mylossoma duriventre. Other frugivores found in flooded forest were Pterodoras granulosus, Auchenipterichthys longimanus, and Brycon cephalus all with low abundances as well (Table 4). The present study was conducted at the end of the high waters season; although fruit phenology of many species in Amazon flooded forests is strongly linked with the flooded season (Kubitzki & Ziburski, 1994), it is possible that the abundance of fruits in the flooded forests around Cao Yarina may have been low during the study period. If that was the case, then frugivore species could have migrated to other areas. However a study of fruit phenology in the area would be necessary in order to make inferences on relationships between fruits production and frugivore fish abundance.

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46 Detritivores was the second most abundant species guild in floating meadows, and was dominated by P. rutiloides. Detritivores fish such as curimatids and semaprochilodids gain energy mainly from phytoplankton (algae) present in the water column all over, while siluriform detritivores gain energy from other plant resources (e.g., tree leaves, tree seeds, wood, C 3 macrophytes and periphyton) (Araujo et al., 1986b). This suggests that detritivores catfish would be more abundant in flooded forest than in floating meadows. In Cao Yarina, detritivores catfish were scarce in both habitats; however the abundance was slightly higher in floating meadows. Light penetration is the primary factor limiting algae production especially for sessile periphyton (Putz & Junk, 1997). These authors hypothesized that higher light interception by the canopy of flooded forest would be limiting periphyton growth and that the root bunches of floating meadows could be a more favorable area for periphyton. This mechanism could explain the higher abundance of detritivores found in the floating meadows of Cao Yarina. Sampling Limitations Although there are limitations in sampling with gill nets (e.g., selectivity of body size (Jensen, 1986, 1990)), the method seems valid for comparing key assemblage characters between flooded forests and floating meadows habitats in a single floodplain. The use of a wider range of mesh size nets (e.g., 12 to 200 mm used by Saint-Paul et al. (2000)) in the future could yield a much more complete picture of the community. In order to increase the amount of research on fish community ecology in flooded forests, it is urgent to develop a reliable, inexpensive sampling technique for fishing in this habitat that can be easily adopted by local researchers. Such a method would yield a big amount

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47 of information of fish assemblages from flooded forest as it has been produced from floating meadows. Conclusion In conclusion, the fish assemblages found in the floodplain Cao Yarina were highly similar with subtly differences. Environmental factors explained part of the variation in fish assemblages. However, ecological processes such as competition and predation may also account for the observed variation in composition and abundance of species (e.g., Layman & Winemiller, 2004). The fact that several species were shared between the two habitats suggests movement of species between the flooded forest and the floating meadows of Cao Yarina during the high waters season. The higher number of rare species caught in the flooded forest suggest that unique resources provided by flooded forests may be critical for sustaining populations of species only present in this habitat during the high-waters season.

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APPENDIX SYSTEMATIC LIST OF SPECIES Table 5. Systematic list of species organized by order, family, genus and species based on gill netting sampling of the Cao Yarina floodplain, Pacaya Samiria National Reserve (PSNR), during May 2003. Taxon Total FF Total FM Osteoglossiformes Arapaimidae 0 0 Arapaima gigas (Schinz, 1822) 1 0 Osteoglossidae 0 0 Osteoglossum bicirrosum (Cuvier, 1829) 6 7 Characiformes Anostomidae (3) Leporinus trifasciatus Steindachner, 1876 2 1 Rhytiodus microlepis Kner, 1858 1 4 Schizodon fasciatus Spix & Agassiz, 1829 4 4 Acestrorhynchidae (3) Acestrorhynchus falcatus (Bloch, 1794) 1 0 Acestrorhynchus falcirostris (Cuvier, 1819) 7 0 Acestrorhynchus microlepis (Schomburgk, 1841) 1 0 Characidae (17) Astyanax bimaculatus (Linnaeus, 1758) 1 2 Brycon cephalus (Gunther, 1869) 1 0 Charax gibbosus (Linnaeus, 1758) 1 2 Colossoma macropomum (Cuvier, 1818) 11 7 Ctenobrycon spilurus (Valenciennes, 1850) 75 25 Cyphocharax cf. festivus 1 0 Gymnocorymbus thayeri Eigenmann, 1908 14 6 Moenkhausia cf. chrysagyrea 2 0 Mylossoma duriventre (Cuvier, 1818) 0 2 Pygocentrus nattereri Kner, 1858 2 27 Roeboides biserialis (Garman, 1890) 1 0 Serrasalmus rhombeus (Linnaeus, 1766) 1 5 Tetragonopterus argenteus Cuvier, 1816 2 0 Tetragonopterus chalceus Spix & Agassiz, 1829 1 0 Triportheus albus Cope, 1872 2 6 Triportheus angulatus (Spix & Agassiz, 1829) 309 48 Curimatidae (8) Curimata vittata (Kner, 1858) 4 1 Curimatella alburna (Muller & Troschel, 1844) 20 6 Curimatella meyeri (Steindachner, 1882) 11 18 Potamorhina altamazonica (Cope, 1878) 33 22 Potamorhina latior (Spix & Agassiz, 1829) 3 18 48

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49 Table 5. Continued Taxon Total FF Total FM Psectrogaster amazonica Eigenmann & Eigenmann, 1889 49 82 Psectrogaster essequibensis (Gunther, 1864) 9 3 Psectrogaster rutiloides (Kner, 1858) 202 209 Erythrinidae (3) 0 0 Erythrinus erythrinus (Bloch & Steindachner, 1801) 1 1 Hoplerythrinus unitaeniatus (Agassiz, 1829) 11 0 Hoplias malabaricus (Bloch, 1794) 6 32 Gasteropelecidae (1) 0 0 Gasteropelecus sternicla (Linnaeus, 1758) 1 1 Prochilodontidae (1) 0 0 Prochilodus nigricans Agassiz, 1829 9 7 Gymnotiformes Apteronotidae (1) Adontosternarchus sp. A 1 0 Parapteronotus hasemani (Ellis, 1913) 0 2 Electrophoridae (1) Electrophorus electricus (Linnaeus, 1766) 1 1 Gymnotidae (3) Gymnotus carapo Linnaeus, 1758 3 1 Gymnotus ucamara Crampton, Lovejoy & Albert, 2003 4 0 Gymnotus varzea Crampton, Thorsen & Albert, 2004 1 0 Sternopygidae (1) Eigenmannia limbata (Schreiner & Miranda Ribeiro, 1903) 0 1 Siluriformes Auchenipteridae (2) Auchenipterichthys longimanus (Gunther, 1864) 2 0 Trachelyopterus galeatus (Linnaeus, 1766) 15 7 Callichthyidae (7) Brochis splendens (Castelnau, 1855) 2 19 Callichthys callichthys (Linnaeus, 1758) 5 1 Dianema longibarbis Cope, 1872 596 436 Hoplosternum littorale (Hancock, 1828) 19 11 Lepthoplosternum sp. PUA (black belly) 0 1 Lepthoplosternum sp. PUB (pepper belly) 1 0 Megalechis thoracata (Valenciennes, 1840) 26 6 Doradidae (5) Agamyxis pectinifrons (Cope, 1870) 1 1 Anadoras grypus (Cope, 1872) 9 2 Doradidae sp. PUA 2 3 Doradidae sp. PUB 0 1 Pterodoras granulosus (Valenciennes, 1821) 16 4 Loricariidae (11) Ancistrus sp. PUA 2 0 Glyptoperichthys scrophus (Cope, 1874) 1 0 Hypoptopoma gulare Cope, 1878 2 10 Liposarcus pardalis (Castelnau, 1855) 19 24 Loricariichthys cf. acutus 3 6

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50 Table 5. Continued Taxon Total FF Total FM Loricariichthys cf. maculatus 7 3 Loricariichthys cf. nudirostris 2 5 Loricariichthys sp.1 3 2 Loricariichthys sp.2 (may be L. maculatus juvenile) 3 27 Loricariinae sp. Indeterminate TF03 1 0 Pseudorinelepis genibarbis (Valenciennes, 1840) 2 3 Pimelodidae (2) 0 0 Rhamdia quelen (Quoy & Gaimard, 1824) 1 2 Sorubim lima (Bloch & Schneider, 1801) 1 2 Perciformes Cichlidae (6) 0 0 Cichlasoma amazonarum Kullander, 1983 7 30 Crenicichla proteus Cope, 1872 1 18 Heros appendiculatus Heckel, 1840 1 1 Hypselacara temporalis (Gunther, 1862) 3 0 Mesonauta mirificus Kullander & Silfvergrip 1991 4 30 Satanoperca jurupari (Heckel, 1840) 5 4 Symbranchiformes Synbranchidae (1) Synbranchus marmoratus Bloch, 1795 1 0 Total number of individuals 1579 1210 Family followed by number of species in parenthesis. Total abundance for FF-flooded forest and FM-floating meadows is provided. Taxonomic nomenclature and authorities follows Reis et al. (2003)

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BIOGRAPHICAL SKETCH Sandra Bibiana Correa was born on November 11 th 1974, in Popayn, Cauca, Colombia. She is one of two children of Jose Alvaro Correa and Rosalba Valencia. She followed her passion for marine life and enrolled at the Universidad del Valle, where she earned her bachelors degree in biology with emphasis in marine biology. During the course of her career she found an opportunity to do research in Amazon fish ecology. Since then, she has devoted her studies to fish communities in different Amazonian locations. After graduating, she worked for several organizations to gain experience in tropical forest ecology. In 2002 she was accepted into the graduate program at the Department of Zoology at the University of Florida where was awarded a Master of Science degree in May 2005. She is still exploring and enjoying the magic of Amazonian ecosystems. 59


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Permanent Link: http://ufdc.ufl.edu/UFE0009463/00001

Material Information

Title: Comparison of Fish Assemblages in Flooded Forest versus Floating Meadows Habitats of an Upper Amazon Floodplain (Pacaya Samiria National Reserve, Peru)
Physical Description: Mixed Material
Copyright Date: 2008

Record Information

Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
System ID: UFE0009463:00001

Permanent Link: http://ufdc.ufl.edu/UFE0009463/00001

Material Information

Title: Comparison of Fish Assemblages in Flooded Forest versus Floating Meadows Habitats of an Upper Amazon Floodplain (Pacaya Samiria National Reserve, Peru)
Physical Description: Mixed Material
Copyright Date: 2008

Record Information

Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
System ID: UFE0009463:00001


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COMPARISON OF FISH ASSEMBLAGES IN FLOODED FOREST VERSUS
FLOATING MEADOWS HABITATS OF AN UPPER AMAZON FLOODPLAIN
(PACAYA SAMIRIA NATIONAL RESERVE, PERU)















By

SANDRA BIBIANA CORREA


A THESIS PRESENTED TO THE GRADUATE SCHOOL OF
THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE
REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE

UNIVERSITY OF FLORIDA


2005

































Copyright 2005

by

Sandra Bibiana Correa
































To my parents and my family. During the course of my life their love and belief in my
potential encouraged me to keep going and achieve my dreams. They supported me in all
of my decisions even when they separated us for long periods of time. I also dedicate this
thesis to Michael Goulding who inspired me to study Amazonian fishes and who is a
pioneer in showing their beauty and fragility to the world.
















ACKNOWLEDGMENTS

This thesis would not have been possible without the support and commitment of

my supervisory committee chair (Dr. James S. Albert) and my committee members (Dr.

Lauren J. Chapman and Dr. William G.R. Crampton). All of them put lots of energy and

time into the development of different aspects of this work. I thank James and Will for

the outstanding time we spent in Peru during the field work that was the basis for this

thesis. They were also crucial during the writing stage, and I enjoyed great discussions on

Amazonian fish natural history and ecology. Lauren was my main support during the data

analysis process. Without her, many of the results presented in this thesis may not have

come out.

I also want to thank my field assistant; B.Sc. Mario Escobedo, who worked on this

project as if it was his own; and with great dedication, made it possible to meet the goal

of 18 hours of continuous sampling per day. I also thank the fishermen who collaborated

on the sampling: Hitler Rodriguez, Wilson Lanza, and Marco from the village of Bretafia.

I thank the Instituto Nacional de Recursos Naturales (INRENA) for the research

permission (under J.S. Albert). I thank the graduate students of the Zoology Department

at University of Florida (UF) for great interactions. I thank Ann Taylor from the Editorial

Office at UF. Finally I want to thank N. Bynum, P. Coley, R. Prendeville, the Grunwald-

Seibel family, and C. Chapman for the valuable support that finally put me in graduate

school.









Last, I would like to acknowledge the support of the National Science Foundation

(NSF-DEB 0215388, PI: J.S. Albert) which founded Project Ucamara and made possible

the expedition for this thesis. This work was also supported with equipment donated by

Idea Wild. I would also like to thank Fish Base, for the incredible resource that this data

base represents to ichthyologists from all around the world.
















TABLE OF CONTENTS
page

A C K N O W L E D G M E N T S ......... .................................................................................... iv

LIST OF TABLES ........................... ...... ............. vii

L IST O F F IG U R E S ............................................................................... viii

ABSTRACT ........ .............. ............. ...... .......... .......... ix

INTRODUCTION .................................. .. ... .... ........ ............ ..

Characterization of Floodplains in the Amazon Basin................................................1
Importance of Flooded Forest and Floating Meadows for Fishes..............................3
Previous Studies on Amazonian Floodplain Fishes..........................................3
Specific Objectives .................. .................................... .. ................ .6

M A TERIALS AN D M ETH OD S............................................................................. 7

R E SU L T S ................................................................................16

D IS C U S S IO N ................................ .....................................................3 7

Species R ichness and A bundance......................................... .......................... 37
Biom ass and Size D istributions ................. _.............................. ............... 39
Diel Variation in Species Richness, Abundance and Biomass.................................41
Fish A ssem blages in Floodplains ........................................ .......... ............... 42
Sam pling L im itations........ ................................................................ ...... .... ..... 46
C onclu sion ......... .................. ..................................... ...........................47

APPENDIX: SYSTEMATIC LIST OF SPECIES ................................. ...............48

L IST O F R E F E R E N C E S ......... .. ............... ................. ................................................5 1

B IO G R A PH IC A L SK E TCH ..................................................................... ..................59















LIST OF TABLES


Table p

1 Abundance of 78 species in each of 10 locations in flooded forest and floating
meadows of the Caho Yarina, Pacaya Samiria National Reserve............................21

2 Mean values of different assemblage characteristics for flooded forest and
floating m eadow s ...................... ...................... .. .. .... ......... ......... 25

3 Comparison of day and night captures between flooded forest and floating
m eadow s ........ .................................................... .................. 26

4 Principal food items for 78 species and their relative abundance as a percentage
of the total abundance (n = 2793 individuals) in flooded forest and floating
meadows, during May 2003 at Caho Yarina, Pacaya Samiria National Reserve ....27

5 Systematic list of species organized by order, family, genus and species based
on gill-netting sampling of the Caho Yarina floodplain, Pacaya Samiria
N national Reserve, during M ay 2003 ............................................. ............... 48















LIST OF FIGURES


Figure p

1 Map of the study area in the Upper Amazon, Pacaya Samiria National Reserve,
P eru ....... ...........................................................14

2 Sam pling place in Location 1 ................................. .......................................15

3 Species richness saturation curves for flooded forest and floating meadows
sa m p le s ........................................................................................................ 3 0

4 Rank-ordered abundances (absolute number of individuals per species) of 78
species of fish captured during May 2003 in Caho Yarina, Pacaya Samiria
N national Reserve, Peru ..................................... ........ .. ...... ... ........ .... 31

5 Abundance per habitat of the 10 most abundant species (absolute number of
individuals per habitat) captured during May 2003 in Caho Yarina, Pacaya
Sam iria N national Reserve, Peru. ........................................ .......................... 32

6 Variation in abundance, biomass, and richness during three periods of capture
(EN, LN, LD) during May 2003 in Caho Yarina, Pacaya Samiria National
R e serv e, P eru .................................................... ................ 3 3

7 Changes in abundance of the five most common species among three sampling
periods in flooded forest and floating meadows ............................................... 34

8 Size frequency distributions of fishes captured per habitat during May 2003 in
Caho Yarina, Pacaya Samiria National Reserve, Peru.........................................35

9 Nonmetric scaling ordination (NMS) of sampling locations in species space
based on the abundances of 57 species that occurred in more than 5% of the
sampling locations during May 2003 in Caho Yarina, Pacaya Samiria National
R e serv e, P eru .................................................... ................ 3 6















Abstract of Thesis Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Master of Science

COMPARISON OF FISH ASSEMBLAGES IN FLOODED FOREST VERSUS
FLOATING MEADOWS HABITATS OF AN UPPER AMAZON FLOODPLAIN
(PACAYA SAMIRIA NATIONAL RESERVE, PERU)

By

Sandra Bibiana Correa

May 2005

Chair: James S. Albert
Major Department: Zoology

Flooded forests and floating meadows of Amazonian floodplains are important

habitats for fishes and yet the distribution and abundance of fish in these habitats are

poorly known. This study presents the first quantitative comparison of fish species

richness and composition, abundance, biomass and body-size distributions between

flooded forests and floating meadows in an Amazon floodplain. This study was

conducted in the floodplain of Reserva Nacional Pacaya Samiria, in the Peruvian

Amazon and sampled fish assemblages in both flooded forests and floating meadows,

using matched sets of gill nets of different mesh sizes. This represents the only reliable

method available of doing a quantitative assessment of fish assemblages in flooded

forest.

Species richness was higher in flooded forest because of to a high percentage of

unique species. Species abundances followed a hollow distribution in which three species

accounted for 70 and 60 % of the total abundance in flooded forest and floating









meadows, respectively. Diel variation in the abundance and richness of fishes was

observed in both habitats. Body-size distributions were very similar in the two habitats

(in the range of 70 to 400 mm). In both flooded forests and floating meadows, most of the

fishes caught were invertebrate feeders and frugivores (i.e., Dianema longibarbis and

Triportheus angulatus) in flooded forest; and detritivores (i.e., Psectrogaster rutiloides)

in floating meadows. Fish predators and frugivores were represented by few species.

Based on species composition and abundance, multivariate analysis suggested the

presence of two subtly different fish assemblages in flooded forest vs. floating meadows.

Water depth accounts for part of the differentiation in fish assemblages. However, the

fact that many species (including all the common ones) were shared between the two

habitats suggests movement of species between habitats in the floating meadows of Cafio

Yarina during the flood season.















INTRODUCTION

Amazon fishes are strongly influenced by the inundation pulse (Goulding, 1980;

Goulding et al. 1988; Junk, 1997). In the Amazon basin, floodplains are periodically

inundated by the overflow of rivers and lakes, or precipitation (Junk, 1997). Floodplains

offer a wide variety of resources for both plants and animals, and a complex nutrient and

energy cycle integrates the aquatic and terrestrial phases (Junk, 1997). The powerful

dynamics driven by the flood pulse make floodplains alternately suitable for aquatic and

terrestrial organisms. Therefore, floodplains are believed to have a principal role in

maintaining high biodiversity of both aquatic and terrestrial organisms (Goulding, 1980;

Junk, 1997; Gopal & Junk, 2000). Furthermore, the Amazon basin contains the most

diverse fish fauna (3,000 species) in the world (Val & Almeida-Val, 1995), which

accounts for about 10 % of the global fish fauna (Groombridge & Jenkins, 1998). Besides

the importance of floodplains in maintaining diversity, floodplains provide much of the

carbon consumed by several commercially important species (Araujo-Lima et at. 1986a;

Benedito-Cecilio et al., 2000). And fishes represent the main sources of protein and

commercial income to riverine people (Bayley, 1989; Crampton et al., 2004).

Characterization of Floodplains in the Amazon Basin

Floodplains are mosaics of lakes, channels (paranas), and levees (restingas), where

flooded forest and floating meadows (patches of aquatic macrophytes) are the principal

vegetation formations (Junk, 1984). Flooded forests are seasonally inundated by either

nutrient-rich rivers (called varzea in Brazil), or by nutrient-poor rivers (called igap6 in









Brazil) (Prance, 1980). Nutrient-rich rivers, originate in the Andes (Goulding et at. 2003).

They have a high concentration of suspended solids, with a near-neutral pH (6 to 7 pH),

and high electric conductivity (50 to 300 tS cm-1) (Junk, 1984). Nutrient-poor rivers,

originating in the Amazonian lowlands, have relatively high transparency, with water

color depending on the soils that they drain, and the concentration of organic compounds

in the water. The characteristic dark tea color of some waters is usually the result of local

podzol-based soil geochemistry. Here there is little sequestering of organic matters,

resulting in waters with a high concentration of humic substances (Leenheer, 1980). In

this type of water, acidity varies between 2 and 5 pH, transparency ranges from 1.30 to

2.90 m (Sioli, 1984), and electric conductivity is low (5 to 30 tS cm-1). In latosoils, the

sequester of humic compounds on the clay matrix results in highly transparent drainage

water (Leenheer, 1980) ranging from 1.10 to 4.30 cm (Sioli, 1984). Additionally, rivers

that drain ancient upland regions of granite shields (i.e., Brazilian and Guiana Shields),

where there is little erosion, are highly transparent (Goulding et at. 2003).

Chemical composition of each water type, nutrient levels, and the length of the

flood season determine the vegetation present in each forest type (Prance, 1979). Biomass

and local tree-species diversity (a-diversity) is higher in varzea than in igap6; however at

a regional scale (P-diversity), igap6 forests are more diverse (Kubitzki, 1989). Flooded

meadows are extensive rafts of floating macrophytes growing along the margins of rivers,

lakes, and channels. The specific composition of these meadows changes from area to

area, depending on water fluctuation. However, the most diverse macrophyte

communities are present in floodplains of nutrient-rich rivers (Junk & Howard-Williams,

1984).









Importance of Flooded Forest and Floating Meadows for Fishes

During the flood season, flooded forest and floating meadows are the most

important habitats for a variety of Amazonian fish species. Flooded forests offer a wide

variety of allochthonous food resources (e.g., terrestrial herbaceous plants, leaves,

flowers, seeds and fruits, and terrestrial invertebrates) (Goulding, 1980), while floating

meadows offer autochthonous food resources (e.g., phytoplankton, periphyton, aquatic

herbaceous plants, and aquatic invertebrates) (Junk et al., 1997). How fish species

partition resources and microhabitats in floodplains is only partially known, partly

because of the paucity of quantitative studies of fish assemblages in flooded forests. In

general, small fish species and juveniles are the main components of the fish assemblage

occupying floating meadows because of the availability of shelter and the abundance of

food among meadows' roots (Saint-Paul & Bayley, 1979; Goulding, 1980; Goulding &

Carvalho, 1982; Junk, 1984b; Sanchez-Botero & Araujo- Lima, 2001; Carvalho &

Araujo-Lima, 2004). When fishes become larger, they move toward the flooded forest or

into the main river channel (Junk, 1997).

Previous Studies on Amazonian Floodplain Fishes

Given the extremely high diversity of fishes contained in the Amazon basin, one of

the most striking questions in the study of Amazonian floodplains is the origin and

maintenance of species diversity. Henderson et al. (1998) summarized historical and

contemporary factors that aid in resolving this question in varzea floodplains. They

speculated that the floodplain fish fauna is characterized by species with wide

distributions and by low endemism. The spatial and temporal interconnection of habitats,

the ephemeral character of habitats at large temporal scale, and the obligate migration

during dry season, act as a group in limiting opportunities for speciation within









floodplains. They suggested that speciation may be occurring in river headwaters, with

subsequent colonization of whitewater floodplains. Moreover, floodplain habitats would

be selected for attributes of colonizing species resulting in more-simple body trends, and

wide phenotypic plasticity.

Other studies on Amazonian floodplain fishes, range from life history of

economically key species (e.g., Arapaima gigas (Hurtado, 1999), Colossoma

macropomum (Goulding & Carvalho, 1982), Cichla sp. (Cala et al. 1996), and pimelodid

catfishes (Arboleda, 1988; Rodriguez, 1991; Celis, 1994; Agudelo, 1994)) to community

ecology (see below), ecophysiology (Junk et al. 1983; Val et at. 1986; Val & Almeida-

Val, 1995), trophic ecology and nutrients flux (Araujo-Lima et al. 1986a; Forsberg et al.

1993; Yossa & Araujo-Lima, 1998; Benedito-Cecilio et al. 2000; Leite et al. 2002),

migration (Vazzoler & Menezes, 1992; Barthem & Goulding, 1997), larval movement

and recruitment (Araujo-Lima et at. 1994; Araujo-Lima & Oliveira, 1998; Carvalho &

Araujo-Lima, 2004), and fisheries (Bayley & Petrere, 1989; Bayley, 1995; Bayley, 1996;

Merona, 1990; Almeida etal. 2001; Crampton et al. 2004).

Most of the studies on community ecology of Amazonian fishes have focused on

associations between floating meadows and fishes (Bayley, 1983; Soares et al., 1986;

Araujo-Lima et al., 1986b; Henderson & Hamilton, 1995; Crampton, 1996; Henderson &

Crampton, 1997; Henderson & Robertson, 1999; Petry et al., 2003). In contrast, there are

few studies on fishes from flooded forests. The most comprehensive study was Goulding

(1980), who made a detailed account of feeding behavior (emphasizing the importance of

seeds and fruits in fish diet). Later, Goulding et al. (1988) made the most-intensive

survey of fish species in an Amazonian affluent (i.e., Rio Negro). They did a









nonsystematic sampling of the Rio Negro's flooded forest and provided a list of 184

species, and food habits of 140 species in this habitat. Henderson and Crampton (1997)

conducted a comparative study of fish richness and relative density in nutrient-poor and

richer floodplain habitats in the Tefe region, Brazil. They sampled both floating meadows

and flooded forests, at dry and flood season, and presented data on species abundance,

distribution, and biomass. However, the fact that they used a different sampling technique

at each habitat (seine nets in meadows, gill nets in flooded forest) did not allow direct

quantitative comparisons of abundance and standing crop. Probably the most complete

study of community ecology of fishes in flooded forests was conducted by Saint Paul et

al. (2000). They compared fish assemblages in nutrient-poor and richer flooded forests

near Manaus, Brazil, by using a wide range of gill-net mesh sizes. They reported 238

species; and contrasted species diversity, distribution, abundance, and biomass between

the two habitats. In all of the studies mentioned above, gill nets were used as the only

available method for quantitative sampling in flooded forests. Although fish length is

related to net mesh size (Jensen, 1990) and gill-net effectiveness varies with fish behavior

(Jensen, 1986), the structure of flooded forests does not allow the use of other sampling

techniques. The only means to improve the efficiency of gill nets is to use a combination

of nets of different mesh sizes (Jensen, 1990); usually arranged in batteries, with the

largest mesh sizes downstream (if there is any flow).

Data on biomass and production of Amazonian fishes are almost nonexistent

(Saint-Paul et at. 2000). There is an estimate of annual fish production of 31.2 g m2 in

marginal floating meadows, and 19.2 g m2 in drifting islands of meadows. The study was

conducted in Lake Mamiraua (Brazil), a nutrient-rich lake, during 8 days at the beginning









of the flood season (Henderson & Hamilton, 1995). Another study in the same lake

estimated a mean of 13.5 g fish m-2, based on 12 days of sampling during the flood

season (Henderson & Crampton, 1997). Both of these estimates were based on seine-net

sampling. In contrast, for flood forest there is an estimate of 33 g fish m-2 (of net area) per

day for a nutrient-poor lake (Lago do Prato, Anavilhanas, Brazil), and 104 g fish m-2 (of

net area) per day for a nutrient-rich lake (Lago do Inacio, Rio Manacapuru, Brazil)

(Saint-Paul et at. 2000). These estimates were based on 48 hours of sampling (over 2

years) during the flood season, and were based on gill-net sampling; therefore the authors

standardized by capture per unit effort (CPUE). The fact that all of the estimators

mentioned above were obtained with different sampling protocols makes them

noncomparable (e.g., seine nets give an estimator of biomass per m2 of meadow, whereas

gill nets give an estimator of biomass per m2 of net surface area). Moreover, fish

abundance and biomass differ from season to season in the same area (Henderson &

Hamilton, 1995; Henderson & Crampton, 1997; Saint-Paul et at. 2000). Thus, to

adequately compare fish biomass between habitats, sampling must be conducted during

the same season and using the same fishing techniques.

Specific Objectives

Given the apparent importance of flooded forest in maintaining fish diversity in

Amazonian floodplains, and the lack of knowledge on how fish species partition

resources and microhabitats in floodplains, the present study used a standardized

sampling technique to compare species richness and composition, abundance, biomass,

and size distribution of fishes in flooded forest with those of floating meadows in a

floodplain of the Peruvian Amazon.















MATERIALS AND METHODS

Study site. This study was conducted in the Cafio Yarina, a tributary of the Rio

Pacaya in the Pacaya Samiria National Reserve (PSNR). This reserve is located at the

confluence of the Marafion and Ucayali Rivers, in the Peruvian Amazon (5 20.575' S;

74 30.117' W). All specimens where collected within 10 km from one of the guard posts

in the reserve (Puesto de Vigilancia 2 "PV2", Fig. 1).

The PSNR is the largest continuous area of protected varzea floodplain in the

Amazon basin (21,500 km2) (INRENA, 2000) and therefore provides a good opportunity

to survey fish communities that have not been highly perturbed as is the case in most of

the lower and central Amazonian floodplains (with exceptions such as the Mamiraua

Reserve, Brazil). The PSNR is located within the Ucamara Depression, an active deposit

of marine and continental sediments dating back from the Late Tertiary to the present

(Bayley et al., 1992). Consequently, the area is an extensive floodplain that gets

inundated most of the year with a short dry season from July to September (INRENA,

2000). Indeed, 86% of the area is represented by inundated forest (51%), seasonally

flooded forest (34%), and rivers and oxbow lakes (1%) (Bayley et al., 1992), making the

landscape a complex mosaic of water bodies all interconnected during the inundation

season. Extensive beds of floating macrophytes cover approximately 30-40% of the total

open water surface area at high water (INRENA, 2000).

The taxonomic composition of macrophytes varies from one patch to another. The

most abundant plant species occurring in the sampled patches were Polygonium sp.,









Pistia stratiotes (L.), Eichhornia crassipes Solms., Paspalum sp., and an unidentified

legume species. Other taxa include Azolla sp., Neptunia sp., Ludvigia sp., Salvinia sp.,

Utricularia sp., and Echinochloeta sp. While most of these macrophyte species are

buoyant and drift with wind and water currents, herbaceous species such as Paspalum

repens Berg, and Polygonium sp.; and shrubs and trees such as Sena sp., and Cecropia sp.

are rooted to the bottom. Rooted plants made the sampling effort with gillnet very

difficult and in many occasions it was impossible to cut depth enough into the meadows

to clear a patch for the nets.

Cafio Yarina is a small affluent (no more than 100 m width) of the Rio Pacaya.

Conductivity ranges from 100 to 200 [t siemens cm-1 in the Pacaya River, similar to that

of "white" water, despite a dark coloration of the river which is more typical of

blackwaters. Water depth varied from 0.6 to 2.5 m in the sampling places in the flooded

forest and from 2.5 to 6.5 m in the floating meadows. Transparency was measured with a

Secchi disk and ranged from 0.8 to 2.2 m in flooded forest and from 1.5 to 2.8 m in the

floating meadows. Margins of the channel were covered by extensive beds of floating

meadows. Behind these macrophytes were patches of shrubs, palms and Cecropia sp.

trees which all together constitute the levee zone. The varzea flooded forest grows behind

this zone.

Fish sampling. This study was designed to quantify fish assemblages in a way that

would facilitate comparison between flooded forests and floating meadows. Fish

collection was conducted during 9 to 21 May 2003 in two habitats: flooded forests and

floating meadows. In each habitat 10 locations where chosen, separated by at least one

km, with the exception of Locations 7 and 8.









At each location I selected three sampling positions, no further than 100 m apart.

At each of these three positions a set of four gill nets of 25, 40, 80 and 120 mm mesh size

were deployed. In the flooded forest, the nets were located from the edge of the dry land

towards the open water, parallel to each other and with the bigger to the smaller mesh

sizes facing the current direction (Jensen, 1990). In floating meadows nets were located

around patches of vegetation and the order of mesh sizes was randomly assigned (Fig. 2).

Nets were 20 m long and depth varied from 3.5 to 5 m. Fishing was conducted for a

period of 18 h at each location. After selecting a location and recording its location with a

Global Positioning System (GPS) receiver, one team of two people went inside the

flooded forest and another team into the floating meadows to select the three sampling

positions. Nets were deployed around 18:00 h. At midnight all captured fishes were

removed from the nets and brought to the field station to be identified and measured. The

nets were visited again at 06:00 h and at noon, and the same procedure was followed each

time. Nets were removed at noon and brought to the field station to be repaired if

necessary and all were deployed again at 18:00 h at the next sampling positions.

Standard length (SL), weight (W), time at capture, and mesh size for all captured

fishes were recorded, and samples of each species were fixed in 10% formaldehyde and

preserved in 70% ethanol. All preserved materials were deposited at the Florida Museum

of Natural History (UF), University of Florida, USA.

Data analysis. Because of the low number of captures at individual sampling

positions, the specimens collected at all three sampling positions for a single location

were combined to produce a meaningful representation of the location. Therefore I

considered locations as sample units, representing each habitat. Statistical analyses were









performed using Stat View 5.0 and JPM 5.0.1, except for Jackknife richness estimator

and the multivariate analyses that were performed using PC-ORD software (McCune and

Mefford, 1999).

Species accumulation curves for each habitat were constructed based on number of

species caught per location. Each location represented 18 h of continuous sampling. Total

species per habitat were estimated with a first and second order Jackknife estimator

(Heltshe and Forrester, 1983) using PC-ORD.

Differences in assemblage characteristics between flooded forest and floating

meadows were evaluated by conducting t-tests on abundance (total number of

individuals), biomass (total weight of individuals), richness (total number of species),

mean SL, maximum SL, mean weight, and maximum weight of fish from 10 locations of

each habitat type. The same set of analyses was also conducted excluding individuals

over 400 mm SL. Biomass was also estimated as "capture per unit of effort" (CPUE) by

calculating the amount of grams of fish caught per meter square of net surface per day of

sampling.

To compare the distribution of abundance of species between flooded forest and

floating meadows, I ranked all species in a habitat based on abundance (n = 72 species in

flooded forest and n = 57 species in floating meadows), giving the rank = 1 to the most

abundant species and continuing the ranking in descending order. If more than one

species presented the same abundance I gave the same rank to all of those species and

then calculate the average rank (ties). I then plotted the ranks against the number of

individuals. Differences in median abundance between habitats for the 10 most abundant

species were also detected using the nonparametric Mann Whitney U test.









Temporal differences in abundance, biomass, and richness were explored using

repeated measures ANOVA. This analysis allowed me to explore the effects of time of

day, habitat, and their interaction on assemblage characters. Contrast analysis was

performed to evaluate differences between time periods.

Distribution of body size of all fishes captured in flooded forest versus floating

meadows was compared using the Kolmogorov-Smimov test for frequency distributions.

To study the distribution of species in the two habitats, multivariate procedures were

performed with a matrix of species abundance per location. The original matrix contained

20 locations x 78 species. Eighteen species present in only one location (5% of the

sampling units) were deleted from the matrix resulting in a new matrix of 20 locations x

60 species. This reduces the noise of uninformative rare species and is appropriate after

species richness and diversity has been analyzed from with the data (McCune and Grace,

2002). However, after deleting the rare species, the coefficient of variance and average

skewness for species were still very high (319.82%, and 2.55 respectively). Therefore the

abundance matrix was transformed using Logio (x + 1). This transformation is useful in

analyzing community data since it decreases the importance of highly abundant species

that would skew cluster and ordination procedures (McCune and Grace, 2002).

An outliers analysis (PC-ORD) pointed out a location from floating meadows

(FM5) as an outlier (standard deviation from average distance among sample units > 2)

(McCune and Grace, 2002). Similarly, a location from flooded forests (FF5) showed as

an isolated point in exploratory ordination analysis (near to the point representing FM5).

Although the standard deviation (Sd) value did not pointed out FF5 as an outlier (Sd =

1.77) I removed the two locations, based on the fact that these locations where highly









predated by piranhas, therefore the species richness and species abundances were

remarkable low in both. These two locations produced an artificial clumping effect in the

remaining 18 locations, over the axis correlated with species richness and diversity.

Finally, I removed from the matrix three species present in only one location after the

removal of locations FM5 and FF5.

The new matrix of 57 species and 18 locations of flooded forest and floating

meadows was analyzed with a nonmetric multi-response permutation procedure (MRPP)

in order to test the null hypothesis of no difference in species composition between the

two habitats. The MRPP (Mielke, 1984) is a nonparametric multivariate procedure to test

for differences between apriori defined groups (e.g., habitats). The nonmetric test

transforms the distance matrix in ranks prior to calculating the test statistic (T). T

describes the separation between groups and is associated to a p-value. A more negative

value of Tmeans a stronger separation between groups. MRPP also provides a measure

of "effect size" called A, which describes within group homogeneity, compared with the

random expectation. A ranges from 0 to 1, therefore, when A=0 the heterogeneity within

groups is not different from that expected by chance, whereas, when A = 1 all items are

identical within each group (McCune & Grace, 2002). McCune et al. (2000) point out

that in community ecology studies, A < 0.1 is common, even when groups are obviously

differ and A > 0.3 is considered high.

To represent the similarities in species composition between locations, I used

nonmetric multidimensional scaling (NMS; Kruskal, 1964; Mather, 1976). NMS is a

nonparametric ordination technique; starting from a matrix of species abundance per

sample, this procedure calculates coefficients of dissimilarity for each species, ranks









those coefficients, and "maps the samples in two or more dimensions, in which distance

between samples reflects similarity in species composition" (Clarke, 1993). The aim of

the analysis is to locate samples in positions (in the graph) that result in the lowest

"stress", stress been a measure of departure of sample positions in the graph from the

initial dissimilarity matrix (Clarke, 1993). Therefore in PC-ORD stress is scaled from

0 to 100 where zero means perfect agreement in rank orders (McCune & Grace, 2002).

NMS has advantages when analyzing community data by not assuming multivariate

normality and by being robust to large numbers of zero values (Clarke, 1993; McCune &

Grace, 2002). The analysis was performed through the autopilot slow-and-thorough

option of NMS in PC-ORD. This option performed 40 runs with real data and 50 runs

with randomized data to find a Monte Carlo test of significance for the best output.

The resulting ordination was rotated 15 degrees. Rotation maximizes the

percentage of variance explained by each axis (McCune & Grace, 2002). The variance

explained was expressed by the coefficient of determination between distances in the

ordination space and distance in the original species space. Sorensen distance was

selected in both cases since this distance measure has been recommended for analyzing

community data (McCune & Grace, 2002). Environmental and assemblage variables

were overlaid by using a joint plot, based on the individual correlations of those variables

with the axes of community ordination (see Clarke, 1993; McCune et al., 2000; and

Peterson & McCune, 2001 for examples of applying this ordination technique on

community data).




















































Figure 1. Map of the study area in the Upper Amazon, Pacaya Samiria National Reserve,
Peru. Number one indicates the location of guard post, Puesto de vigilancia 2
(PV2), around which the sampling was conducted. Map was made by
W.G.R. Crampton based on 1998 1:300,000 Landsat TM5 images; and used
with his permission.









































Figure 2. Sampling place in Location 1. Different mesh size nets are represented by
shades. Nets closer to Cafio Yarina, Pacaya Samiria National Reserve, Peru, are
located in floating meadows whereas nets located next to the land edge, are
placed into the adjacent flooded forest. Arrows represent the direction of the
water current.















RESULTS

A total of 2789 individuals representing 6 orders, 20 families, 61 genera and 78

species of fishes were captured in this study (See appendix). Characiformes (35 species)

was the most diverse taxon followed by Siluriformes (27 species), Perciformes (6 species

of Cichlids), Gymnotiformes (7 species), Osteoglossiformes (2 species), and

Synbranchiformes (1 species). Characiformes and Siluriformes alone accounted for 80%

of the total number of species. Seventy three species were caught in flooded forest, 21 of

which were exclusively found in this habitat. In floating meadows, 57 species were

caught and only five were exclusively found in this habitat (Table 1). Species richness

per location ranged from 9 to 33 in flooded forest (average 19.3 7.6) and from 14 to 30

in floating meadows (average 19.0 5.1).

Species richness approximates an asymptote in both habitats (Fig. 3), for the

species with individuals ranging from 38 to 740 mm SL. Total species estimates were

102 (first order jackknife) and 117 (second order jackknife) for flooded forest. In this

habitat, 32 species (44%) occurred in only one location. In floating meadows total species

was estimated in 74 (first order jackknife) and 79 (second order jackknife). In this habitat,

19 species (33%) occurred in only one location.

Assemblage characters, averaged across locations and including abundance (total

number of individuals), biomass (total weight of individuals), richness (total number of

species), mean SL, maximum SL, mean weight, and maximum weight offish were









similar in between flooded forest and floating meadows (Table 2). Similar results were

found when individuals over 400 mm SL were excluded from the analysis.

Total biomass was higher in flooded forest than in floating meadows (71.0 kg vs.

49.9 kg respectively). A total of 1000.22 m2 gill net surface area was deployed at each

habitat. Capture per unit effort was estimated as 7.10 g fish m-2 per day in flooded forest

and 4.99 g fish m-2 per day in floating meadows for fishes ranging between 38 to 740 mm

(n = 2789 individuals).

Species abundances at each habitat followed an expected hollow distribution

(Hubbell, 2001) where few species were very abundant and most of the remaining species

were represented by few individuals (Fig. 4). Indeed, in flooded forest 70% of the total

abundance was accounted for by three species: Dianema longibarbis, Psectrogaster

rutiloides, and Triportheus angulatus, while 25 species (35% of the total richness)

occurred only once. In floating meadows 60% of the total abundance was represented by

D. longibarbis, P. rutiloides, and Psectrogaster amazonica (Fig. 5), and 12 species (21%

of the total richness) occurred only once. Among the 10 most abundant species, only

T. angulatus was significantly more abundant in flooded forest than in floating meadows

(Mann Whitney, U= 6.0, P < 0.001; FF 30.9 19.0; FM 4.8 3.4), whereas Hoplias

malabaricus (U= 15.5, P < 0.01; FF 0.6 1.1; FM 3.2 2.7), Cichlasoma amazonarum

(U= 24.50, P < 0.05; FF 0.7 1.6; FM 3.0 2.8), and Mesonauta mirificus (U= 20.50,

P < 0.05; FF 0.4 0.7; FM 3.0 2.8) were significantly more abundant in floating

meadows (Fig. 5).

Numbers of individuals and species richness were higher at night; however biomass

did not show diel differences (Table 3). Abundance and richness were highest at the









18:00 to 24:00 period and the lowest at the 6:00 to 12:00 period (Fig. 6). Moreover, there

were temporal differences in abundance within each habitat, as well. In flooded forest the

abundance was highest during the 18:00 to 24:00 period and lowest at the 6:00 to 12:00

period (Contrast analysis for: 18:00 to 24:00 vs. 6:00 to 12:00, F= 45.31, df= 1,

P < 0.0001; 24:00 to 6:00 vs. 6:00 to 12:00, F= 9.72, df= 1, P < 0.01; Fig. 6). In floating

meadows the abundance was also highest at the 18:00 24:00 period, but it was equally

lower in the other two periods of time (Contrast analysis for: 18:00 24:00 vs. 6:00 to

12:00, F= 12.36, df= 1, P< 0.01; 24:00 to 6:00 vs. 6:00 to 12:00, F= 0.36, df= 1,

P = NS; Fig. 6). At the species level, the five most common species (D. longibarbis,

P. rutiloides, T. angulatus, P. amazonica, and Ctenobrycon spilurus) represented highest

abundances during the nocturnal samples (18:00 to 24:00 and 24:00 to 06:00) in both

flooded forests and floating meadows (Fig. 7). There were no temporal differences in

species richness within each habitat. Biomass was similar at day and night samples with

approximately 2 kg of fish per net area (2000.4 m2) in a 6 h period (Fig. 6).

Body size distributions were very similar in the two habitats. There were no

significant differences in the frequencies of SL intervals (Kolmogorov-Smirnov,

X2 = 0.667, P = NS) (Fig. 8).

Community-structure patterns. Flooded forest locations differed from floating

meadows locations in species composition (MRPP A = 0.15, P < 0.001, where A

describes within group homogeneity compared to the random expectation). Although the

A value was small, it is significantly different from zero and indicates that the

homogeneity within groups is higher than expected by chance, meaning that there is a

different assemblage at each habitat type.









Flooded forest locations separates from floating meadows locations along the

vertical ordination axis (Fig. 9). After rotation, the vertical axis explained 32% of the

variation in fish assemblages and was correlated with water depth (Pearson Correlation,

r = 0.68, P < 0.01), but not significantly correlated with water transparency (r = 0.40,

P = NS). The vertical axis was also positively correlated with the abundance of

C. amazonarum (r = 0.65, P < 0.005), H. malabaricus (r = 0.65, P < 0.005), Crenicichla

proteus (r = 0.59, P < 0.01), M. mirificus (r = 0.53, P < 0.05), and negatively correlated

with the abundance of T. angulatus (r = -0.86, P < 0.001), Clli hhnl 1y\ (lliL /n1yh.

(r = -0.71, P < 0.001), Gymnotus ucamara (r = -0.67, P < 0.005), Ancistrus sp. PUA

(r = -0.62, P < 0.01), and Hoplerythrinus unitaeniatus (r = -0.60, P < 0.01).

The horizontal axis explained 30% of the variation in fish assemblages and was

positively correlated with the abundance of P. rutiloides (r = 0.85, P < 0.001),

P. amazonica (r = 0.82, P < 0.001), Potamorhina altamazonica (r = 0.68, P < 0.005),

Curimatella meyeri (r = 0.58,, P < 0.05), Doradidae sp. PUA (r = 0.51, P < 0.05), and

negatively correlated with the abundance of Colossoma macropomum (r = -0.50,

P < 0.05) and Liposarcuspardalis (r = -0.49, P < 0.05). A third axis and explained 20%

of the variation in fish assemblages and was correlated with overall species richness

(r = 0.84, P < 0.001),

The coefficients of correlation with environmental and assemblage variables were

obtained from a joint plot and are in accordance with the highest values of depth

(t= -8.31, P < 0.0001; 144 cm + 47.8 in flooded forest vs. 403 cm 85.8 in floating

meadows). Species richness was similar in both habitats (t = 0.1, P = NS; 19.3 species

7.6 in flooded forest vs. 19.0 species 5.1 in floating meadows).









Literature review and personal observations by the author and W.G.R. Crampton,

were used to categorize the diet of caught species. In flooded forest, invertebrate feeders

(30% of 73 species) was the trophic guild with higher number of species, followed by

detritivores (20%), piscivores (15%), and frugivores (7%). Relative abundance, showed a

similar pattern, but in each guild, there was a single species that accounted for most of the

abundance in this habitat. These species were D. longibarbis (39% of the total

abundance), T. angulatus (20%), and P. rutiloides (13%) (Table 4).

In floating meadows, a similar pattern to the one found in flooded forest was

observed. Invertebrate feeders (40% of 57 species), detritivores (28%), and piscivores

(16%) were the trophic guilds with higher number of species. However the relative

abundance of invertebrate feeders, detritivores and piscivores was higher. Again a single

species accounted for most of the abundance in each guild. Dianema longibarbis (36% of

the total abundance), was the most abundant species, P. rutiloides (17% of the total

abundance), was the second most abundant species and the abundance of T. angulatus

(4% of the total abundance), was much lower than in flooded forest.













Table 1. Abundance of 78 species in each of 10 locations in flooded forest and floating meadows of the Cafio Yarina, Pacaya Samiria
National Reserve.


FM FM FM FM FM FM FM FM FM FM FF1 FF FF FF FF5 FF FF FF FF9 FF1 Total


1 2
Acestrorhynchus falcatus 0 0
Acestrorhynchus falcirostris 0 0
Acestrorhynchus microlepis 0 0
Adontosternarchus sp. A 0 0
Agamyxis pectinifrons 0 0
Anadoras grypus 0 0
Ancistrus sp. PUA 0 0
Arapaima gigas 0 0
Astyanax bimaculatus 0 0
Atli. //e'/u ti'l'/ i /11 /1 0 0
longimanus
Brochis splendens 0 0
Brycon cephalus 0 0
C, lli ///l.//y) i // l.//1y/ 0 0
Charax gibbosus 0 0
Cichlasoma amazonarum 7 3
Colossoma macropomum 1 0
Crenicichla proteus 1 0
Ctenobrycon spilurus 1 0
Curimata vittata 0 0
Curimatella alburna 0 2
Curimatella meyeri 9 2
Cyphocharax cf festivus 0 0
Dianema longibarbis 105 46


4 5
0 0
0 0
0 0
0 0
1 0
1 1
0 0
0 0
1 0
0 0

0 0
0 0
0 0
1 1
4 0
0 0
5 2
9 0
1 0
4 0
3 0
0 0
29 1


8 9 10
0 0 0
0 0 0
0 0 0
0 0 0
0 0 0
0 0 0
0 0 0
0 0 0
0 0 1
0 0 0


14 5 0 0
0 0 0 0
0000
0 1 0 0
0 0 0 0
4 0 2 8
0 5 0 1
3 0 5 2
1 0 1 1
0 0 0 0
0000






0 0 0 0
0 2 0 0
0 0 0 0
0000


3 4
0 1
0 6
0 1
0 0
1 0
0 3
1 0
0 0
0 0
0 0


0 0 0 0
0 0 0 0
0000
0 1 0 0
0 0 1 0
5 0 0 0
0 0 0 0
0 1 0 0
23 9 20 1
3 0 0 0
3000
6 1 10 0
8 0 1 0
0 0 0 0
0000


56 52 46 23 63 29 124 226 57 8


0 2
0 1
0 1
0 0
0 1
3 1
0 0
12 5
0 0
0 3
0 0
0 0


8
0 0
0 0
0 0
0 0
0 0
4 0
0 0
1 0
0 1
0 0

0 0
0 0
1 0
0 0
0 0
3 0
0 0
3 2
0 0
0 0
2 0
0 1


1
7
1
1
2
11
2
1
3
2

21
1
6
3
37
18
19
100
5
26
29
1


50 22 22 16 42 1032


Taxon













Table 1. Continued
Taxon


FM FM FM FM FM FM FM FM FM FM FF1 FF FF FF FF5 FF FF FF FF9 FF1 Total


Doradidae sp. PUA 2
Doradidae sp. PUB 1
Eigenmannia limbata 0
Doradidae sp. PUA 2
Doradidae sp. PUB 1
Eigenmannia limbata 0
Electrophorus electricus 0
Erythrinus erythrinus 0
Gasteropelecus sternicla 0
Gymnocorymbus thayeri 1
Gymnotus carapo 0
Gymnotus ucamara 0
Gymnotus varzea 0
Heros appendiculatus 0
Hoplerythrinus 0
unitaeniatus
Hoplias malabaricus 4
Hoplosternum littorale 3
Hypoptopoma gulare 0
Hypselacara temporalis 0
Leporinus trifasciatus 0
Lepthoplosternum sp. PUA 1
(black belly)
Lepthoplosternum sp. PUB 0
(pepper belly)
Liposarcus pardalis 0
Lot it iiL /h11,/) cf acutus 0


2 3
1 0
0 0
0 0
1 0
0 0
0 0
0 0
0 0
0 0
0 0
1 0
0 0
0 0
0 0
0 0

2 1
1 0
0 0
0 0
0 0
0 0


7 8
0 0
0 0
1 0
0 0
0 0
1 0
1 0
0 0
0 0
2 0
0 0
0 0
0 0
1 0
0 0

8 4
0 2
0 0
0 0
0 0
0 0


2 3 4
234
1 0 0
0 0 0
0 0 0
1 0 0
0 0 0
0 0 0
0 0 0
0 0 0
0 0 1
0 4 6
0 1 1
0 0 1
0 0 0
0 0 1
0 1 1

0 0 3
3 4 5
0 0 0
1 0 0
0 0 0
0 0 0
000


0 0 0 0 0 0 0 0 0 1 0 0 0


1 0 0
0 1 0
010


1 9 8
5 0 0
500


1 2 4
0 0 0
000


7 8
1 0 0
0 0 0
0 0 0
1 0 0
0 0 0
0 0 0
0 1 0
0 1 0
0 0 0
4 0 0
0 0 0
0 1 0
0 0 0
0 0 0
0 3 0

0 1 0
3 0 0
0 0 0
0 0 0
2 0 0
0 0 0


0 0 0 0 0


1 0 0
2 1 0
210


0
0 5
0 1
0 1
0 5
0 1
0 1
0 2
0 2
0 2
0 20
0 4
1 4
0 1
0 2
1 11

0 38
0 30
0 12
2 3
0 3
0 1

0 1













Table 1. Continued
Taxon

Lot ii I11 iiL /11/,) cf.
maculatus
Lot itI // iiL /11/)/ Cf.
nudirostris
Lot ii .I i/L /11/ij sp. 1
Lot ii 1 ti // ily sp.2
(possibly juvenile of
maculatus)
Loricariinae sp. Indet.
TF03


FM FM FM FM FM FM FM FM FM FM FF1 FF FF FF FF5 FF FF FF FF9 FF1 Total


2 3 4
234 0 1
001


0 0 0 0


0 0 2
002 3 0
530


0 0 0 0


Megalechis thoracata 0 1 0 0
Mesonauta mirificus 8 0 2 4
Moenkhausia cf 0 0 0 0
chrysagyrea
Mylossoma duriventre 0 0 0 0
Osteoglossum bicirrosum 0 0 0 1
Parapteronotus hasemani 1 0 0 1
Potamorhina altamazonica 5 3 4 2
Potamorhina latior 13 3 0 2
Prochilodus nigricans 0 0 0 0
Psectrogaster amazonica 7 14 39 2
PsectroPaster 0 3 0 0


7 8 9
789 0 0
200


0 0 5 0 0


0 0 0
000
001


0 0 0 0 0


2
0


essequibensis
Psectrogaster rutiloides 65 56 23 17 1
Pseudorinelepis genibarbis 0 0 0 0 2
Pterodoras granulosus 0 2 0 2 0
Ptei ygop~lQ1i l/l/,) scrophus 0 0 0 0 0
Pyvocentrus nattereri 4 0 2 4 1


0


3 1 0 0
5 0 0 2
0 0 0 0

0 0 2 0
0 3 1 0
0 0 0 0
4 1 0 2
0 0 0 0
4 3 0 0
9 1 5 3
0 0 0 0

20 2 5 5
0 1 0 0
0 0 0 0
0 0 0 0
0 7 0 0
5002
0000

0020
03 10
0000


4300

0000

20 2 5 5

0000

0700


10 2 3 4
0 0 1 0 1

0 0 0 0 0

0 0 0 0 0
17 0 1 2 0


0 0 0 1 0


1 0
6 1
0 0

0 0
2 0
0 0
1 0
0 0
0 4
0 0
0 0

15 1
0 1
0 0
0 1
0 0


4 4 0
440
0 0
001

0 0 0
000
2 0 1
0 0 0
000
18 1 5
3 0 0
0 0 0
000
21 0 0
7 0 0
700

140 1 3
0 0 1
16 0 0
0 0 0
0 0 1


6
0 4


7 8
1 0 0


0 0 2 0 0


0 0 0
0 0 0
000


0 0 0 0 0


0 5 1
0 0 0
0 1 0

0 0 0
0 0 0
0 0 0
7 1 0
0 0 0
5 0 0
12 8 7
2 0 0
000
010

000

000


500
12 8 7
200


26 4 24 1
0 0 0 0
0 0 0 0
0 0 0 0
0 0 0 0


0 7

0 5
0 31


0 1

3 3'
0 3,
0 2

0 2
1 1:
0 2
0 5:
0 2
0 11
0 1:
0 1i












Table 1. Continued
Taxon FM FM FM FM FM FM FM FM FM FM FF1 FF FF FF FF5 FF FF FF FF9 FF1 Total
1 2 3 4 5 6 7 8 9 10 2 3 4 6 7 8 0
Rhamdia quelen 0 0 0 2 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 3
Rhytiodus microlepis 0 0 0 0 3 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 5
Roeboides biserialis 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1
Satanopercajurupari 0 0 0 4 0 0 0 0 0 0 0 0 0 2 0 1 2 0 0 0 9
Schizodonfasciatus 0 0 1 2 0 0 0 1 0 0 0 0 0 1 2 0 0 1 0 0 8
Serrasalmus rhombeus 0 0 3 1 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 6
Sorubimlima 0 0 0 0 2 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 3
Synbranchus marmoratus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1
Tetragonopterus argenteus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 2
Tetragonopterus chalceus 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1
Trachelyopterus galeatus 2 2 0 0 0 0 1 0 0 2 1 5 2 4 0 0 1 2 0 0 22
Triportheus albus 0 1 1 0 1 1 0 0 2 0 0 0 0 0 1 0 0 1 0 0 8
Triportheus angulatus 3 10 1 10 3 5 0 7 4 5 37 43 55 22 5 20 63 33 23 8 357
Total 244 159 107 129 40 119 121 95 60 138 99 439 318 169 27 115 174 100 76 65 2789
Abbreviations: FM floating meadows; FF flooded forest. The numbers following the abbreviations correspond to the sampling
location in each habitat (1-10).






25


Table 2. Mean values of different assemblage characteristics for flooded forest and
floating meadows.
FF FM P
Mean abundance (total number of
fish) 157.9 128 121.4 56 0.4189
Mean biomass (g) 7099.5 4510 4994.1 + 4516 0.3107
Mean richness (species) 19.4 + 8 19.3 5 0.9731
Mean standard length (mm) 97.6 + 15 94.6 + 15 0.6474
Maximum standard length (mm) 527.8 164 366.4 196 0.0806
Mean weight (g) 62.9 + 50 45.7 + 47 0.4368
Maximum weight (g) 1717.0 + 1283 1156.1 + 1242 0.3339
Calculations are based on totals, means, or maximum values averaged across 10 locations
in each habitat type. Abbreviations: FF flooded forest, FM floating meadows. A t-test
was used to detect differences between the two habitats in Cafio Yarina, Pacaya Samiria
National Reserve.









Table 3. Comparison of day and night captures between flooded forest and floating
meadows. Repeated measures ANOVA was used to detect effects of time,
habitat and their interaction on abundance (loglOx+1 transformed), richness
(loglOx+1 transformed), and biomass (loglOx+1 transformed) of fishes with
three times of capture (18:00 24:00h, 24:00 6:00h, 6:00 12:00h) and two
habitats in Cafio Yarina, Pacaya Samiria National Reserve. For abundance,
Mauchley's criterion indicated rejection of the compound symmetry
assumption, therefore adjusted probability values (G, Greenhouse-Geisser and
H-F, Huynh-Feldt) are provided.
SOURCE OF Adj. G-
VARIATION df SS F P G H-F
Abundance


Between-subjects effect
Habitat
Error
Within-subjects effect
Time
Time*habitat
Error (time)

Biomass
Between-subjects effect
Habitat
Error
Within-subjects effect
Time
Time*habitat
Error (time)

Richness
Between-subjects effect
Habitat
Error
Within subjects effect
Time
Time*habitat
Error (time)


0.117
4.641

4.222
1.094
3.053



0.018
14.092

1.478
0.987
15.167



0.169
1.550

0.820
0.164
1.416


0.452 NS


24.894 <0.001
6.453 <0.005 <0.05




0.023 NS


1.754 NS
1.172 NS




1.963 NS


10.419 <0.001
2.087 NS


<0.01









Table 4. Principal food items for 78 species and their relative abundance as a percentage
of the total abundance (n = 2793 individuals) in flooded forest and floating
meadows, during May 2003 at Cafio Yarina, Pacaya Samiria National
Reserve. Food items were extracted from literature. Diet of species in which a
reference is not provided was assessed by W.G.R. Crampton and S.B. Correa.
Relative
Taxon Diet Reference abundance


Acestrorhynchusfalcatus
Acestrorhynchus
falcirostris
Acestrorhynchus
microlepis
Adontosternarchus sp. A
Agamyxis pectinifrons
Anadoras grypus
Ancistrus sp. PUA
Arapaima gigas

Astyanax bimaculatus
A, t he/ te'/ i/ h/ 1 /).
longimanus

Brochis splendens

Brycon cephalus
C, llit hil" /i ,lli' hil"c
Charax gibbosus

Cichlasoma amazonarum
Colossoma macropomum
Crenicichla proteus

Ctenobrycon spilurus

Curimata vittata

Curimatella alburna
Curimatella meyeri
Cyphocharax cf festivus
Dianema longibarbis
Doradidae sp. PUA
Doradidae sp. PUB
Eigenmannia limbata
Electrophorus electricus


Fish

Fish

Fish
Invertebrates
Invertebrates
Invertebrates
Detritus
Fish

Zooplankton

Fruits
Aquatic
invertebrates

Fruits
Zooplankton
Fish

Plants
Fruits
Fish

Zooplankton

Detritus

Detritus
Plants
Detritus
Invertebrates
Invertebrates
Invertebrates
Invertebrates
Fish


Planquette et al.
(1996)
Goulding et al.
(1988)
Planquette et al.
(1996)


Planquette et al.
(1996)
Manheimer et al.
(2003)

Burgess (1989)
Goulding (1980),
Anonymous (1981)
Mol (1995)
Winemiller (1989)
Stawikowski &
Werner (1998)
Goulding (1980)
Kullander (1986)
Mills & Vevers
(1989)
Val & de Almeida-
Val (1995)
Goulding et al.
(1988)
Soares et al. (1986)


0.04

0.25

0.04
0.04
0.07
0.39
0.07
0.04

0.11

0.07

0.75

0.04
0.21
0.11

1.32
0.64
0.68

3.59

0.18

0.93
1.04
0.04
36.95
0.18
0.04
0.04
0.07









Table 4. Continued


Taxon


Diet


Erythrinus erythrinus

Gasteropelecus sternicla
Gymnocorymbus thayeri
Gymnotus carapo

Gymnotus ucamara

Gymnotus varzea
Heros efasciatus
Hoplerythrinus
unitaeniatus

Hoplias malabaricus

Hoplosternum littorale
Hypoptopoma gulare
Hypselacara temporalis
Leporinus trifasciatus
Lepiih, q, ,/ ii 11111 sp. PUA
(black belly)
Lepih, l uMi 11111 sp. PUB
(pepper belly)

Liposarcus pardalis

Lot ii It ii /111.\ cf acutus
Lot it I iiL /11/1) cf.
maculatus
Lot it i iiL y11/ cf.
nudirostris
Lot it I// iiL /11/ 1 sp. 1
Lot i I / iiL //11/j sp. 2 (may
be maculatus juvenile)
Loricariinae sp. Indet.
TF03
Megalechis thoracata
Mesonauta mirificus
Moenkhausia cf
chrysagyrea leucopomis
Mylossoma duriventre


Fish
Aquatic
invertebrates
Invertebrates
Invertebrates
Aquatic
invertebrates
Aquatic
invertebrates
Plants
Aquatic
invertebrates

Fish
Aquatic
invertebrates
Detritus
Fish
Periphyton

Invertebrates

Invertebrates

Detritus
Aquatic
invertebrates


Reference
Planquette et al.
(1996)
Mills & Vevers
(1989)


Crampton et al.
(2003)


Planquette et al.
(1996)
Boujard et al.
(1997)


Yossa & Araujo-
Lima (1998)
Goulding et al.
(1988)


Detritus

Detritus
Detritus

Detritus


Detritus
Zooplankton
Periphyton

Invertebrates
Fruits


Mol (1995)


Goulding (1980)


Relative
abundance

0.07

0.07
0.72
0.14


0.14

0.04
0.07

0.39

1.36

1.07
0.43
0.11
0.11

0.04

0.04

1.54

0.29

0.39

0.25
0.18

1.07

0.04
1.15
1.22

0.07
0.07









Table 4. Continued


Taxon

Osteoglossum bicirrosum
Parapteronotus hasemani
Potamorhina altamazonica

Potamorhina latior
Prochilodus nigricans
Psectrogaster amazonica
Psectrogaster
essequibensis
Psectrogaster rutiloides
Pseudorinelepis genibarbis
Pterodoras granulosus
Ptei) 'gv li ln i scrophus

Pygocentrus nattereri

Rhamdia quelen
Rhytiodus microlepis
Roeboides biserialis
Satanopercajurupari

Schizodonfasciatus
Serrasalmus rhombeus
Sorubim lima

Synbranchus marmoratus

Tetragonopterus argenteus

Tetragonopterus chalceus

Trachelyopterus galeatus
Triportheus albus
Triportheus angulatus


Diet
Terrestrial
invertebrates
Invertebrates
Detritus

Detritus
Detritus
Detritus


Reference

Goulding (1980)

Soares et al. (1986)
Goulding et al.
(1988)
Soares et al. (1986)


Detritus
Detritus
Detritus
Fruits
Detritus


Fish

Fish
Plants
Fish scales
Invertebrates

Plants
Fish
Fish
Terrestrial
invertebrates

Invertebrates
Aquatic
invertebrates

Fish
Invertebrates
Fruits


Sazima & Machado
(1990)
Boujard et al.
(1997)
Soares et al. (1986)

Keith et al. (2000)
Planquette et al.
(1996)
Correa (1999)
Goulding (1981)

Soares et al. (1986)
Silvano et al.
(2001)
Goulding et al.
(1988)
Le Bail et al.
(2000)

Goulding (1980)


Relative
abundance


0.47
0.07
1.79

0.75
0.57
4.87

0.43
14.72
0.18
0.72
0.04

1.04

0.11
0.18
0.04
0.32

0.29
0.21
0.11

0.04

0.07

0.04

0.79
0.29
12.80











80

70- FF
70
F AFM
'5 60 o
Q,. A A
) 50 -
d A
Z 40 -
O
> 30 -A

20 -
E
3 10 -

0
0 1 2 3 4 5 6 7 8 9 10 11

Samples



Figure 3. Species richness saturation curves for flooded forest and floating meadows
samples. Fish ranged from 38 mm to 740 mm (n=2789 individuals) and were
caught with gill nets of 25, 40, 80, and 120 mm mesh size. Sampling was
conducted during May 2003 at Cafio Yarina, Pacaya Samiria National
Reserve, Peru.










-1000




0 10

1 . . . . .l l.. . .1 1 1.
1 10 18 26 33 42 61 61 61 61
Rank of species
A
1000
-a













100


10
-

















1 9 17 25 3 2 42 52 52

Rank of species
B


Figure 4. Rank ordered abundances (absolute number of individuals per species) of 78
species of fish captured during May 2003 in Cahio Yarina, Pacaya Samiria
National Reserve, Peru. A) flooded forest (72 species). B) floating meadows
(57 species). Mean species richness was equal in the two habitats (P= 0.9731).
P value originated by t- test. Note the hollow curve where only three species
account for 70% and 60% of the total abundance in flooded forest and floating
meadows, respectively. See text for method of ranking species.
-a




1C 111111111111111111....
< ( 7 53 4 25













meadows, respectively. See text for method of ranking species.












700

600

500

400

300

200

100

0


E '
c 5
Q|


cL 9

M
Oct
t1
*
ll
^1


E FF
O FM


SI .


Cs
0)
0C
r


'1)

0 M3 0 M

CL


-ao
0s
'16


NE -


62
So -g
ct!

-J


62
(to
- '
0-c
o -o
; ct
E


( E



c M
" N
C E
% (t


-S
3M
0 3
C.)
Q)
|E


Species


Figure 5. Abundance per habitat of the 10 most abundant species (absolute number of
individuals per habitat) captured during May 2003 in Cafo Yarina, Pacaya
Samiria National Reserve, Peru. ** indicates P>0.001, indicates P>0.05.
Abbreviations: FF flooded forest, FM floating meadows.










) 16

I. 12
0 8
I-I


0 0
Z
EN LN ED

A

S 120 a

b DFF
S 80 d FM

S 0e e
S 40

E
z 0
EN LN ED
B
4000
S3000
E 2000
0
mL 1000
0
EN LN ED

Time C

Figure 6. Variation in abundance, biomass, and richness during three periods of capture
(EN, LN, LD) during May 2003 in Cafio Yarina, Pacaya Samiria National
Reserve, Peru. Graphs A and C represent means combining the two habitats,
graph B shows variation between habitats. Bars represent standard error.
Abbreviations: EN early night: 18:00 24:00; LN late night: 24:00-06:00;
ED early day: 06:00 12:00; FF flooded forest, FM floating meadows.












S400 FF FM
S350 -
> 300 -
o Dianema longibarbis
250 o Psectrogaster rutiloides
200 A Triportheus angulatus
200 A
SA150 Psectrogaster amazonica
SA Ctenobrycon spilurus
100 -
50 t X
0I *I I

EN LN ED EN LN ED
Samples

Figure 7. Changes in abundance of the five most common species among three sampling
periods in flooded forest and floating meadows. Sampling was conducted
during May 2003 at Cafio Yarina, Pacaya Samiria National Reserve, Peru.
Abbreviations as in Figure 6.










700
FF
600
o FM
500

400

g 300
LL
200
100 -
oI I I-I I I I---
50 70 90 110 130 150 200 250 300 350 400 >400
SL Interval (mm)


Figure 8. Size frequency distributions of fishes captured per habitat during May 2003 in
Cafio Yarina, Pacaya Samiria National Reserve, Peru. Abbreviations: SL -
standard length, FF flooded forest, FM floating meadows.









A


A A Depth A
Transparency
A


NMS 3








Habitat
SFM
SFF
A
A
A
A

Habitat
AFM
AFF

NMS 1
Figure 9. Non metric scaling ordination (NMS) of sampling locations in species space
based on the abundances of 57 species that occurred in more than 5% of the
sampling locations during May 2003 in Cafio Yarina, Pacaya Samiria National
Reserve, Peru. The vertical axis accounts for 32% while the horizontal axis
accounts for 30% of the variation in the data. Filled triangles represent
sampling locations in flooded forest (FF) and open triangles represent
sampling locations in floating meadows (FM). The distances between
triangles in the ordination are approximately proportional to the dissimilarities
between the sampling locations. Environmental variables are join-plotted
expressing its relationship with ordination scores. Length and angle of
correlation vectors represents the strength of the correlation. The final stress
value for a three dimensional representation was 11.367. Instability
(0.000001) was the lowest out of the 10 repetitions.















DISCUSSION

Species Richness and Abundance

The present study provides the first empirical data on the relative species

abundance in an Amazonian flooded forest. Relative abundances of species often follow

a hollow distribution with very few dominant species and a long tail of rare species

(Hubbell, 2001; Magurran & Henderson, 2003). This hypothesis was corroborated in a

study of fish assemblages in Amazonian floating meadows (Henderson & Crampton,

1997). In flooded forests of Cafio Yarina, 70% of the total abundance was represented by

three species: D. longibarbis, P. rutiloides and T. angulatus; while 25 species (35% of the

total richness) only occurred once. In floating meadows 60% of the total abundance was

represented by D. longibarbis, P. rutiloides and P. amazonica; while 12 species (21% of

the total richness) occurred only once.

What allows a species to be abundant in a habitat depends on characteristics of the

habitat such as food resources and habitat structure, on ecological processes such as

competition and predation, and on stochastic factors. Moreover, the identity of the

dominant species in a habitat may change from place to place and year to year depending

on migration and recolonization of the habitats (Hubbell, 2001). The reproductive

biology of the callichthyid D. longibarbis, the dominant species in both habitats, is not

well known. Callichthyids in general are reported as K- strategists (sensu Pianka, 1970)

(Junk et al., 1997) and Riehl & Baensch (1991) reported bubble nest-building behavior in

D. longibarbis. Other species with nesting behaviors are defined as "equilibrium" life









history strategist (sensu Winemiller, 1989), characterized also by prolonged breeding

seasons, and parental investment in individual offspring, probably resulting in enhanced

juvenile survivorship. This type of species usually has relatively stable sedentary local

populations (Winemiller, 1989). This fact allows me speculate that D. longibarbis may be

a year round resident of floating meadows that sends colonizers to adjacent flooded forest

during the rise of the waters. To prove this hypothesis, a multiseason study in these

habitats would be necessary.

In this study, species richness was similar in both flooded forest and floating

meadows (72 and 57 species respectively). First and second order Jackknife estimators

(Heltshe & Forrester, 1983) estimate the number of species to be higher in both habitats

than my absolute accounts and the number of species in flooded forest as higher (102 and

117 species respectively) than in floating meadows (74 and 79 species respectively). First

order Jackknife estimators has been considered as the most accurate method for

estimating species richness (Palmer 1990; Palmer 1991), however, it is susceptible to

high number of rare species in the data set (Palmer 1990); indeed I found four times more

rare species in flooded forest than in floating meadows. A higher species richness and

biomass in flooded forest was expected because the flooded forest constitutes an

expansion of available habitat in the floodplain during the flooded season, therefore is

expected that many species would colonize this habitat with the rise of the water. The

diversity and abundance of resources provided by flooded forests (Goulding 1980, Junk

et al., 1997b) seems to be the driving force for colonization. For migratory fishes, flooded

forests are a key habitat that provides enough food for them to accumulate fat reserves as









preparation for subsequent reproductive migration at the low waters season (Junk et al.,

1997b, Carvalho & Araujo-Lima, 2004).

Biomass and Size Distributions

Higher biomass along with higher species richness was expected in flooded forest

because of increased availability of habitat during the flooded season. Total biomass was

similar in the two habitats (73.3 kg vs. 51.1 kg for flooded forest and floating meadows

respectively) and there was no difference in average biomass between habitats. Flooded

forest however had a slightly higher CPUE than floating meadows (7.33 and 5.11 g fish


-22
m-2 net surface per sampled day, respectively). A comparable estimate of CPUE in

flooded forest was much higher (104 g m2 per day; Saint-Paul et al., 2000) than the

CPUE value reported in the present study. Although the area covered during sampling in

Cafio Yarina was higher than the sampled area covered by Saint-Paul et al. (2000) (i.e.,

1000.15 m2 vs. 772 m2, respectively), their study used a wider range of mesh sizes (13

different sizes, ranging from 12 to 200 mm) which in turn could caught a wider range of

fish sizes, and therefore a highest biomass. For floating meadows, there is not comparable

data since the data available are estimations of standing crop (Bayley 1983; Henderson &

Hamilton 1995; Henderson & Crampton 1997).

Body size distributions were highly similar between the two habitats, although

differences were expected. A higher number of small fishes in floating meadows were

expected, since this habitat is recognized as nursery habitat for juveniles of many species

(Saint-Paul & Bayley 1979; Goulding 1980; Junk 1984, Sanchez-Botero & Araujo-Lima

2001), whereas in flooded forest small fishes could be more susceptible to predation;

therefore bigger-sized fish were expected.









The body size range found in Cafio Yarina was truncated at the extremes (38 to

740 mm SL). This range most likely reflect the chosen mesh sizes (i.e., 25 to 120 mm)

leading to under sample juveniles, small fish, and very large fish. Nevertheless, the shape

of the distribution of body size frequencies of fishes in the sampled range was similar in

both habitats. This result suggest that juveniles and small fishes (over 38 mm) of the

species caught at Cafio Yarina are using both flooded forest and floating meadows

habitats in the same proportion. This contrast with previous hypothesis of floating

meadows been a preferred habitat for juveniles because the availability of food and

shelter (Junk, 1997). Large fishes (under 740 mm) seem also to be using both habitats in

similar proportions despite the differences in habitat complexity.

It is difficult to estimate what percentage of the community falls in and out of this

range of body sizes found in Cafio Yarina (38 to 740 mm SL). For example, in sandy

beach communities in the Orinoco basin, Layman & Winemiller (2004) found that

around 50% of the fishes were < 50 mm SL. In a nutrient-poor river in the Amazon basin,

Goulding et al. (1988) found that around 100 species (approximately 20% of total species

richness) reach maturity at 30 mm, and all dominant species were adults < 40 mm. In

contrast, in the nutrient rich floodplain of the Solimoes-Amazonas River, juveniles of the

medium sized fishes tend to dominate (Goulding et al., 1988). Goulding et al. (1988)

hypothesize that in nutrient rich systems, the abundance of juveniles of bigger sized

fishes would be higher compared with low nutrient systems, where early maturation

seems to be a strategy due to competitive exclusion in food limited habitats. Indeed, in

Cafio Yarina (a nutrient rich system), the four dominant species were adults of medium









sized fishes: D. longibarbis (46 to 103 mm), P. rutiloides (53 to 204 mm), T angulatus

(56 to 150 mm), and P. amazonica (54 to 131 mm).

Diel Variation in Species Richness, Abundance and Biomass

Diel changeover in species composition is known to occur in Neotropical fish

assemblages (Arrington & Winemiller, 2003). In a study in the Orinoco floodplain,

Arrington & Winemiller (2003) found nocturnal samples to be higher in number of

species and individual abundances and explained the phenomenon as "morphological

trade-offs in foraging and anti-predator defenses". In Cafio Yarina, the differences in

species richness and abundance between day and night samples were impressive. In both

flooded forest and floating meadows, species richness and abundance were much higher

at night samples.

While in flooded forest abundance was much higher at night samples (both early

nigh and late night), in floating meadows abundance was higher at the early night

samples but similar at both late night and early day samples species. Moreover,

abundance at these time periods was higher than abundance at the early day samples in

flooded forest.

Differences in structural complexity between the flooded forest and floating

meadows could be the mechanism allowing the higher abundance caught at day samples

in floating meadows. The submerged portions of floating meadows are composed of a

very dense matrix of stems and roots that may limit the pass of light during the day,

whereas flooded forests, despite of having lots of stems and submerged branches, is a

more open habitat. Consequently light penetration during the day time could be higher

than in floating meadows. Therefore, predation risk at day time could be higher in

flooded forests. It has been demonstrated that fishes are able to assess predation risk and









modify their foraging behavior through foraging rate/mortality risk tradeoff mechanisms

(Mittelbach, 1981; Werner etal., 1983; Mittelbach, 1984). Therefore predation avoidance

behaviors may lead susceptible individuals to limit the use of rich habitats (Werner et al.,

1983) as could be happening in flooded forest during the day time, but probably not in

floating meadows where a more dense vegetation could reduce the risk of predation.

However, the reduced abundance of fish predators in the samples does not provide

evidence to support this hypothesis.

The absence of diel change in biomass between habitats, despite of the differences

in abundance, leads one to expect that a high amount of small fishes caught during night

samples could weight as much as a few large fishes caught in day samples. Indeed, this

was observed in flooded forest, where 21 individuals (23% of the total abundance) of

5 species (Arapaima gigas, C. macropomum, H. unitaeniatus, H. malabaricus, and

L. pardalis) accounted for 85% of the total biomass in the day samples. In floating

meadows, the trend was more dramatic. Twenty individuals (7% of the total abundance)

of five species (C. macropomum, Electrophorus electricus, Leporinus trifasciatus,

L. pardalis, and Osteoglossum bicirrhosum) accounted for 75% of the total biomass in

the day samples.

Fish Assemblages in Floodplains

All quantitative analyses of community variables (i.e. abundance, biomass, species

richness, and body size), without accounting for the species identity, did not provide

evidence to separate the ichthyofauna of Cafio Yarina in two assemblages. However,

multivariate analysis of species abundances (i.e., MRPP and NMS ordination) suggested

the presence of two subtly different fish assemblages in flooded forest vs. floating

meadows. Floating meadows locations were characterized by deeper and more









transparent waters than flooded forest. However, these habitats differ also in many other

characteristics such as type of vegetation, structural complexity of the submerged portion

of the vegetation, but especially in food resources. These variables are difficult to

quantify and therefore it is hard to determine the individual effect on particular fish

species. Specific habitat selection has been reported before in floodplain fish

communities related to abiotic factors such as type of water (Saint-Paul at al., 2000),

oxygen concentration (Junk et al., 1983; Winemiller, 1996), and water depth and

transparency (Rodriguez & Lewis, 1994; Tejerina-Garro et al., 1998). In this last case,

abundance of visually oriented predators (e.g., characids and cichlids) was positive

correlated to water depth and transparency.

In the present study, fish predators were scarce (19 and 4.6 % of the total species

richness and abundance, respectively). Species of important predatory families such as

Serrasalmidae were very scarce in the samples, indeed only two species were caught,

Serrasalmus rhombeus and Pygocentrus nattereri. Only six juvenile specimens (45 to 76

mm) of S. rhombeus and 29 juvenile and adult specimens (63 to 205 mm) of P. nattereri

were caught, mostly in the floating meadows. Pimelodids, another common family of

silurid predators, was only represented by few specimens of Sorubim lima (n=3, 168 to

196 mm). Other common predators inhabiting flooded forest such as Cichla monoculus

(Correa, 1999), were completely absent from the samples. Hoplias malabaricus, the most

abundant predator species in Cafio Yarina (1.4%), was scarcely found, mainly in floating

meadows. A partial explanation for the low abundance of some large sized predator

fishes is based on the limited efficiency of the gill nest. Surprisingly one individual of

Arapaima gigas (a sub adult of 600 mm SL) was caught in this study when usually









A. gigas are caught with harpoons or nets of mesh sizes bigger than 120 mm. However, it

is unlikely that if other predator species, especially medium sized ones, were present in

the system not even single individuals of each of those species were caught.

Consequently an alternative explanation could be based on low productivity of the

floodplain of Cafio Yarina. CPUE in both flooded forest and floating meadows was low;

therefore it is possible that low prey abundance could be limiting predators' abundance as

predicted by the Lotka-Volterra predation model (Begon et al., 1996). CPUE was low in

the present study, but this fact alone does not provide sufficient evidence to support a low

productivity hypothesis for Cafio Yarina. Another possible explanation for the scarcity of

some predator species (e.g., Cichla monoculus and Pimelodid species) could be based on

over fishing in the area before the reserve was established. At the time this study was

conducted, signals of illegal fishermen in the area were observed and indeed some of the

gill nets used in this study were stolen.

Variable food resources could also lead to differences in species composition

between habitats. In both flooded forests and floating meadows, the majority of the

species caught feed on invertebrates (aquatic and/or terrestrial), detritus, or fish. In

flooded forest, invertebrate feeders and frugivores were dominant; whereas in floating

meadows, invertebrate feeders and detritivores were dominant. These patterns were

predicted, based on the differential food availability of flooded forests and floating

meadows (seeds, fruits, and periphyton, vs. macrophytes, detritus, and authochtonous

invertebrates) (Junk, 1997).

Invertebrate feeders were expected to be an important guild in both habitats.

Aquatic invertebrates are abundant in floating meadows (Junk, 1997) and terrestrial









invertebrates are an important input of the terrestrial vegetation of the flooded forest

(Goulding, 1980). The diversity of invertebrate feeders was high in both habitats but each

species (except for D. longibarbis) presented a low abundance. It is possible that the

ability of gymnotiform fishes to detect gill nets with their electric sense

(W.G.R. Crampton, pers. commun.) may undermine the efficiency of gill nets in

capturing them. Gymnotiforms are important components of floating meadows

ichthyofauna (Crampton, 1996) but only seven species were found in this study, five of

them been invertebrate feeders.

Although a frugivore species (T. angulatus)was the second most abundant species

in flooded forest, the total number of frugivore species found in Cafio Yarina was

surprising low. Serrasalminae, a sub-family with highly specialized frugivorous fishes

(Goulding, 1980), was only represented by few individuals of Colossoma macropomum,

and Mylossoma duriventre. Other frugivores found in flooded forest were Pterodoras

granulosus, A41/ /henipi i kh1liy longimanus, and Brycon cephalus all with low

abundances as well (Table 4). The present study was conducted at the end of the high

waters season; although fruit phenology of many species in Amazon flooded forests is

strongly linked with the flooded season (Kubitzki & Ziburski, 1994), it is possible that

the abundance of fruits in the flooded forests around Cafio Yarina may have been low

during the study period. If that was the case, then frugivore species could have migrated

to other areas. However a study of fruit phenology in the area would be necessary in

order to make inferences on relationships between fruits production and frugivore fish

abundance.









Detritivores was the second most abundant species guild in floating meadows, and

was dominated by P. rutiloides. Detritivores fish such as curimatids and

semaprochilodids gain energy mainly from phytoplankton (algae) present in the water

column all over, while siluriform detritivores gain energy from other plant resources

(e.g., tree leaves, tree seeds, wood, C3 macrophytes and periphyton) (Araujo et al.,

1986b). This suggests that detritivores catfish would be more abundant in flooded forest

than in floating meadows. In Cafio Yarina, detritivores catfish were scarce in both

habitats; however the abundance was slightly higher in floating meadows. Light

penetration is the primary factor limiting algae production especially for sessile

periphyton (Putz & Junk, 1997). These authors hypothesized that higher light interception

by the canopy of flooded forest would be limiting periphyton growth and that the root

bunches of floating meadows could be a more favorable area for periphyton. This

mechanism could explain the higher abundance of detritivores found in the floating

meadows of Cafio Yarina.

Sampling Limitations

Although there are limitations in sampling with gill nets (e.g., selectivity of body

size (Jensen, 1986, 1990)), the method seems valid for comparing key assemblage

characters between flooded forests and floating meadows habitats in a single floodplain.

The use of a wider range of mesh size nets (e.g., 12 to 200 mm used by Saint-Paul et al.

(2000)) in the future could yield a much more complete picture of the community. In

order to increase the amount of research on fish community ecology in flooded forests, it

is urgent to develop a reliable, inexpensive sampling technique for fishing in this habitat

that can be easily adopted by local researchers. Such a method would yield a big amount









of information of fish assemblages from flooded forest as it has been produced from

floating meadows.

Conclusion

In conclusion, the fish assemblages found in the floodplain Cafio Yarina were

highly similar with subtly differences. Environmental factors explained part of the

variation in fish assemblages. However, ecological processes such as competition and

predation may also account for the observed variation in composition and abundance of

species (e.g., Layman & Winemiller, 2004). The fact that several species were shared

between the two habitats suggests movement of species between the flooded forest and

the floating meadows of Cafio Yarina during the high waters season. The higher number

of rare species caught in the flooded forest suggest that unique resources provided by

flooded forests may be critical for sustaining populations of species only present in this

habitat during the high-waters season.















APPENDIX
SYSTEMATIC LIST OF SPECIES

Table 5. Systematic list of species organized by order, family, genus and species based on
gill netting sampling of the Cafio Yarina floodplain, Pacaya Samiria National
Reserve (PSNR), during May 2003.
Total Total
Taxon FF FM
Osteoglossiformes
Arapaimidae 0 0
Arapaima gigas (Schinz, 1822) 1 0
Osteoglossidae 0 0
Osteoglossum bicirrosum (Cuvier, 1829) 6 7
Characiformes
Anostomidae (3)
Leporinus trifasciatus Steindachner, 1876 2 1
Rhytiodus microlepis Kner, 1858 1 4
Schizodonfasciatus Spix & Agassiz, 1829 4 4
Acestrorhynchidae (3)
Acestrorhynchusfalcatus (Bloch, 1794) 1 0
Acestrorhynchusfalcirostris (Cuvier, 1819) 7 0
Acestrorhynchus microlepis (Schomburgk, 1841) 1 0
Characidae (17)
Astyanax bimaculatus (Linnaeus, 1758) 1 2
Brycon cephalus (Gunther, 1869) 1 0
Charax gibbosus (Linnaeus, 1758) 1 2
Colossoma macropomum (Cuvier, 1818) 11 7
Ctenobrycon spilurus (Valenciennes, 1850) 75 25
Cyphocharax cf. festivus 1 0
Gymnocorymbus thayeri Eigenmann, 1908 14 6
Moenkhausia cf. chrysagyrea 2 0
Mylossoma duriventre (Cuvier, 1818) 0 2
Pygocentrus nattereri Kner, 1858 2 27
Roeboides biserialis (Garman, 1890) 1 0
Serrasalmus rhombeus (Linnaeus, 1766) 1 5
Tetragonopterus argenteus Cuvier, 1816 2 0
Tetragonopterus chalceus Spix & Agassiz, 1829 1 0
Triportheus albus Cope, 1872 2 6
Triportheus angulatus (Spix & Agassiz, 1829) 309 48
Curimatidae (8)
Curimata vittata (Kner, 1858) 4 1
Curimatella alburna (Muller & Troschel, 1844) 20 6
Curimatella meyeri (Steindachner, 1882) 11 18
Potamorhina altamazonica (Cope, 1878) 33 22
Potamorhina latior (Spix & Agassiz, 1829) 3 18










Table 5. Continued


Taxon
Psectrogaster amazonica Eigenmann & Eigenmann, 1889
Psectrogaster essequibensis (Gunther, 1864)
Psectrogaster rutiloides (Kner, 1858)
Erythrinidae (3)
Erythrinus erythrinus (Bloch & Steindachner, 1801)
Hoplerythrinus unitaeniatus (Agassiz, 1829)
Hoplias malabaricus (Bloch, 1794)
Gasteropelecidae (1)
Gasteropelecus sternicla (Linnaeus, 1758)
Prochilodontidae (1)
Prochilodus nigricans Agassiz, 1829
Gymnotiformes
Apteronotidae (1)
Adontosternarchus sp. A
Parapteronotus hasemani (Ellis, 1913)
Electrophoridae (1)
Electrophorus electricus (Linnaeus, 1766)
Gymnotidae (3)
Gymnotus carapo Linnaeus, 1758
Gymnotus ucamara Crampton, Lovejoy & Albert, 2003
Gymnotus varzea Crampton, Thorsen & Albert, 2004
Stemopygidae (1)
Eigenmannia limbata (Schreiner & Miranda Ribeiro, 1903)
Siluriformes
Auchenipteridae (2)
Auchenipterichthys longimanus (Gunther, 1864)
Trachelyopterus galeatus (Linnaeus, 1766)
Callichthyidae (7)
Brochis splendens (Castelnau, 1855)
Callichthys callichthys (Linnaeus, 1758)
Dianema longibarbis Cope, 1872
Hoplosternum littorale (Hancock, 1828)
Lepthoplosternum sp. PUA (black belly)
Lepthoplosternum sp. PUB (pepper belly)
Megalechis thoracata (Valenciennes, 1840)
Doradidae (5)
Agamyxis pectinifrons (Cope, 1870)
Anadoras grypus (Cope, 1872)
Doradidae sp. PUA
Doradidae sp. PUB
Pterodoras granulosus (Valenciennes, 1821)
Loricariidae (11)
Ancistrus sp. PUA
Glyptoperichthys scrophus (Cope, 1874)
Hypoptopoma gulare Cope, 1878
Liposarcus pardalis (Castelnau, 1855)
Loricariichthys cf. acutus


Total Total
FF FM
49 82
9 3
202 209
0 0
1 1
11 0
6 32
0 0
1 1
0 0
9 7


1 1


0 1










Table 5. Continued
Total Total
Taxon FF FM
Loricariichthys cf. maculatus 7 3
Loricariichthys cf. nudirostris 2 5
Loricariichthys sp.l 3 2
Loricariichthys sp.2 (may be L. maculatus juvenile) 3 27
Loricariinae sp. Indeterminate TF03 1 0
Pseudorinelepis genibarbis (Valenciennes, 1840) 2 3
Pimelodidae (2) 0 0
Rhamdia quelen (Quoy & Gaimard, 1824) 1 2
Sorubim lima (Bloch & Schneider, 1801) 1 2
Perciformes
Cichlidae (6) 0 0
Cichlasoma amazonarum Kullander, 1983 7 30
Crenicichlaproteus Cope, 1872 1 18
Heros appendiculatus Heckel, 1840 1 1
Hypselacara temporalis (Gunther, 1862) 3 0
Mesonauta mirificus Kullander & Silfvergrip 1991 4 30
Satanopercajurupari (Heckel, 1840) 5 4
Symbranchiformes
Synbranchidae (1)
Synbranchus marmoratus Bloch, 1795 1 0
Total number of individuals 1579 1210
Family followed by number of species in parenthesis. Total abundance for FF-
flooded forest and FM-floating meadows is provided. Taxonomic nomenclature and
authorities follows Reis et al. (2003)















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BIOGRAPHICAL SKETCH

Sandra Bibiana Correa was born on November 11th, 1974, in Popayan, Cauca,

Colombia. She is one of two children of Jose Alvaro Correa and Rosalba Valencia. She

followed her passion for marine life and enrolled at the Universidad del Valle, where she

earned her bachelor's degree in biology with emphasis in marine biology. During the

course of her career she found an opportunity to do research in Amazon fish ecology.

Since then, she has devoted her studies to fish communities in different Amazonian

locations. After graduating, she worked for several organizations to gain experience in

tropical forest ecology. In 2002 she was accepted into the graduate program at the

Department of Zoology at the University of Florida where was awarded a Master of

Science degree in May 2005. She is still exploring and enjoying the magic of Amazonian

ecosystems.